MAUREEN E. DOWN Starfishes

from the Caribbean

f and the Gulf of

Mexico

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 126
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Smithsonian Institution
SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 126

Maureen E. Downey Starfishes

from the Caribbean

and the Gulf of

Mexico

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1973
ABSTRACT

Downey, Maureen E. Starfishes from the Caribbean and the Gulf of Mexico.

Smithsonian Contribution to Zoology, number 126, 158 pages, 2 figures, 48 plates,

1973.—Asteroidea collected in the Gulf of Mexico and around the Caribbean by

vessels of the Bureau of Commercial Fisheries at Pascagoula, Mississippi, and

the Texas A&M University research vessel Alaminos numbered 95 species, 8 of

which, described elsewhere, were new. Fifty-six genera, of which three were new,

are represented in the collections. Keys to the orders, families, genera, and

species are provided. A brief historical resume and a general discussion of

zoogeography are included, as well as short descriptions of each genus and species.

Official publication date is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year.

Library of Congress Cataloging in Publication Data

Downey, Maureen E.

Starfishes from the Caribbean and the Gulf of Mexico.

(Smithsonian contributions to zoology, no. 126)

Bibliography: p.

1. Starfishes—Caribbean Sea. 2. Starfishes—Mexico, Gulf of. I. Title. II. Series: Smithsonian

Institution. Smithsonian contributions to zoology, no. 126.

QL1.S54 no. 126 [QL384.A8] 591'.08s [593'.93'0916364] 72-10019

For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402

Price $2.85 domestic postpaid, or S2.50 GPO Bookstore
Contents

Page

Introduction 1

Acknowledgments 2

Historical Review 2

Zoogeography 5

Station Data 9

Glossary 17

Key to the Orders of Asteroidea 21

Order Platyasterida 21

Family Luidiidae 21

Genus Luidia 21

Key to the Species of Luidia 22

L. senegalensis 22

L. clathrata 22

L. alternate 23

L. sagamina 24

L. barbadensis 24

L. elegans 25

Other Tropical-Subtropical Species of Luidia Not Represented in

This Collection 25

Order Paxillosida 25

Key to the Families of Paxillosida 26

Family Astropectinidae 26

Key to the Genera of Astropectinidae 26

Genus Astropecten 26

Key to the Species of Astropecten 27

A. americanus 27

A. articulatus 28

A. cingulatus 28

A. comptus 29

A. duplicatus 29

A. marginatus 29

A. nitidus 30

Other Species of Astropecten Not Taken in These Collections 31

Genus Psilaster 31

Key to the Species of Psilaster 31

P. cassiope 31

P. patagiatus 32

Other Atlantic Species of Psilaster 32

Genus Persephonaster 32

P. echinulatus 33

Other Atlantic Species of Persephonaster 33

Genus Plutonaster 34

P. intermedius 34

Genus Dytaster 35

D. insignis 35

Genus Blakiaster 36

B. conicus 36

Genus Tethyaster 36

T. grandis 37

Other Species of Tethyaster 37

iii
Page

Genus Dipsacaster 37

D. antillensis 38

Family Goniopectinidae 38

Key to the Genera of Goniopectinidae 38

Genus Goniopecten 38

G. demonstrans 38

Genus Prionaster 39

P. elegans 39

Family Benthopectinidae 39

Key to the Genera of Benthopectinidae 40

Genus Benthopecten 40

B. simplex 40

Genus Cheiraster 41

Key to the Species of Cheiraster 41

C. mirabilis 41

C. echinulatus 42

C. enoplus 42

Genus Pectinaster 43

Key to the Species of Pectinaster 43

P. mixtus 43

P. gracilis '. 44

Order Valvatida 44

Key to the Families of Valvatida 44

Family Odontasteridae 44

Genus Odontaster 45

Key to the Species of Odontaster 45

O. hispidus 45

O. setosus 45

Family Goniasteridae 46

Key to the Genera of Goniasteridae 47

Genus Goniaster 46

G. tessellatus 46

Genus Anthenoides 48

Key to the Species of Anthenoides 48

A. piercei 48

Genus Ceramaster 49

C. grenadensis , 49

Genus Peltaster 50

Key to the Species of Peltaster • 50

P. nidarosiensis 50

P. placenta 51

Genus Plinthaster 52

P. dentatus 52

Genus Tessellaster 53

T. notabilis 53

Genus Tosia 53

T. parva 54

Genus Rosaster 54

R. alexandri 54

Genus Circeaster 55

C. americanus 55

Genus Litonotaster 55

Key to the Species of Litonotaster 56

L. intermedius 56

Genus Paragonaster 56

Key to the Species of Paragonaster 56

P. subtilis 57

Genus Nymphaster 57

iv
Page

Key to the Species of Nymphaster 57

N. arenatus 58

N. subspinosus 59

Genus Pseudarchaster 59

Key to the Species of Pseudarchaster 59

P. gracilis 59

Family Oreasteridae 60

Genus Oreaster 60

O. reticulatus 60

Family Ophidiasteridae 61

Key to the Genera of Ophidiasteridae 62

Genus Chaetaster 62

C. nodosus 63

Genus Drachmaster 63

D. bullisi 63

Genus Tamaria 64

Key to the Species of Tamaria 64

T. floridae 64

T. halperni 65

T. passiflora 65

Genus Linckia 66

Key to the Species of Linckia 66

L. guildingii 66

L. nodosa 67

L. bouvieri 67

Genus Ophidiaster 68

Key to the Species of Ophidiaster 68

O. guildingii 68

Genus Narcissia 69

N. trigonaria 69

Order Spinulosida .-.- 70

Key to the Families of Spinulosida 70

Family Solasteridae 70

Key to the Genera of Solasteridae 70

Genus Solaster 70

Key to the Species of Solaster 70

S. notophrynus 71

S. caribbaeus 71

Genus Lophaster 71

L. verrilli 72

Family Pterasteridae 72

Key to the Genera of Pterasteridae 72

Genus Hymenaster 72

Key to the Species of Hymenaster 73

H. rex 73

H. modestus 74

H. anomalus 75

Genus Pteraster 75

Key to the Species of Pteraster 76

P. personatus 76

P. caribbaeus 77

P. rugosus 77

P. militaroides 78

P. acicula 79

P. species 79

Genus Calyptraster 79

C. personatus 80

Family Ganeriidae 80

Genus Leilaster 80

L. radians 80
Page

Family Poraniidae 81

Genus Poraniella 81

P. regularis 81

Genus Marginaster 82

M. pectinatus 82

Family Echinasteridae 82

Key to the Genera of Echinasteridae 88

Genus Henricia 83

Key to the Species of Henricia 83

H. sexradiata 83

H. antillarum 84

H. species 85

Genus Echinaster 85

Key to the Species of Echinaster 86

E. serpentarius 86

E. echinophorus 86

E. sentus 87

E. modestus 87

E. species A 88

E. brasiliensis 88

E. species C 89

E. species B 89

Genus Verrillaster 89

V. spinulosus 89

Order Forcipulatida 90

Family Asteriidae 90

Key to the Genera of Asteriidae 91

Genus Ampheraster 90

A. alaminos 91

Genus Sclerasterias 91

S. contorta 92

Genus Asterias 92

A. forbesi 93

Genus Coscinasterias 93

C. tenuispina 93

Genus Coronaster 94

C. briareus 94

Order Zorocallida 95

Family Zoroasteridae 95

Key to the Genera of Zoroasteridae 95

Genus Doraster 95

D. constellatus 95

Genus Mammaster 97

M. sigsbeei 97

Genus Zoroaster 97

Z. fulgens 97

Order Euclasterida 98

Family Brisingidae 98

Key to the Genera of Brisingidae 98

Genus Odinia 99

O. antillensis 99

Genus Midgardia 99

Af. xandaros 99

Literature Cited 100

Plates Ill

vi
Maureen E. Downey Starfishes

from the Caribbean

and the Gulf of

Mexico

Introduction

Two collections of Asteroidea, probably the most

comprehensive ever made in the Caribbean and the

Gulf of Mexico, have yielded 95 species, of 56 genera. Eight new species and three new genera have

been described from these collections. On the basis

of these collections, which cover a wider area,

greater depth range, and longer collecting period

than any previous starfish collections from this

region, certain revisions in the classification and

corrections to the taxonomy have been made possible. Keys to the taxa are given and brief descriptions of each species, as well as indications of the

distribution, both vertical and horizontal. Keys

are to be construed as applying only to the starfishes of this area. Despite the scope of these collections, about fifty nominal species previously

reported in the literature from the area were not

included in the present collections.

The source of the collections was the Bureau of

Commercial Fisheries (now the National Marine

Fisheries Service) at Pascagoula, Mississippi, where

a survey of the fauna of the Gulf and Caribbean

has been underway for about 20 years, first under

the direction of Dr. Stewart Springer, and later

under the direction of Harvey R. Bullis. The Department of Oceanography of Texas A&M University has, for the past six years, been surveying the

deepwater fauna of the Gulf of Mexico, under the

direction of Dr. Willis Pequegnat. Over a thousand

specimens of Asteroidea collected by these two

Maureen E. Downey, Department of Invertebrate Zoology,

National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

operations have been examined, as well as comparative material in the National Museum of Natural History, the Museum of Comparative Zoology

at Harvard, and material borrowed from other institutions and individuals.

The classification used is essentially that of Spencer and Wright (1966), with certain modifications.

The abundant material available has allowed the

solution of several taxonomic problems, the most

important being the separation of the family Zoroasteridae from the order Forcipulatida and the

establishment of a new order, Zorocallida (Downey,

1970), to contain it, and the abolition of the family

Chaetasteridae (only one genus)—the genus Chaetaster has been returned to the family Ophidiasteridae. Of the three new genera discovered in these

collections, Verrillaster (in the Echinasteridae) accommodates Echinaster spinulosus Verrill; the remaining species of the genus Echinaster form a

coherent group, from which spinulosus differs generically. Drachmaster (in the Ophidiasteridae)

has been added (Downey, 1970) for a new species

which clarified the systematic position of the genus

Chaetaster; and Midgardia (in the Brisingidae)

to contain a new species which could not be included in any of the existing genera. The figures

in parentheses in the list that follows represent the

total number of genera in the family (before the

diagonal) and the number of genera in each family

represented in these collections (after the diagonal):

Class Asteroidea (288/56)

Order Platyasterida (2/1)

Platasteriidae (1/0)

Luidiidae (1/1)

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Order Paxillosida (41/14)

Astropectinidae (20/8)

Goniopectinidae (4/2)

Porcellanasteridae (9/0)

Benthopectinidae (8/4)

Order Valvatida (93/23)

Sphaerasteridae (1/0)

Odontasteridae (5/1)

Archasteridae (1/0)

Goniasteridae (44/13)

Oreasteridae (20/1)

Ophidiasteridae (21/8)

Radiasteridae (2/0)

Order Spinulosida (72/11)

Helianthasteridae (3/0)

Solasteridae (7/2)

Korethrasteridae (4/0)

Pythonasteridae (3/0)

Pterasteridae (9/3)

Aster in idae (17/0)

Ganeriidae (8/1)

Poraniidae (13/2)

Echinasteridae (8/3)

Valvasteridae (1/0)

Acanthasteridae (1/0)

Mithrodiidae (1/0)

Metrodiridae (1/0)

Order Forcipulatida (82/5)

Heliasteridae (1/0)

Asteriidae (57/5)

Order Zorocallidae (7/3)

Zoroasteridae (7/3)

Order Euclasterida (17/2)

Brisingidae (17/2)

Nearly 20 percent of the known genera of starfishes

are represented in the restricted area covered by

this report. All of the orders are present, as are all

but two of the larger families.

ACKNOWLEDGMENTS.—It would be impossible to

adequately express my gratitude to all of the people

who have given me so much help and encouragement during the course of this study. First, I would

like to thank Mr. Harvey R. Bullis and Dr. Willis

Pequegnat for supplying the material for this study;

Dr. Pequegnat and Dr. Thomas Bright, of Texas

A&M University, and Dr. Ivan Goodbody, of the

University of the West Indies, for allowing me to

participate in collecting trips in the Gulf of Mexico, the Yucatan Peninsula, and Jamaica; Dr. Jerald

A. Halpern, of Hobart and William Smith College,

for the use of his unpublished PhD thesis and for

his invaluable help and advice on certain taxonomic problems; and last, but far from least, Dr.

Raymond B. Manning, Chairman of the Department of Invertebrate Zoology of the National Museum of Natural History, for permission to pursue

this study; Dr. David L. Pawson, Curator, Division

of Echinoderms, without whose sympathetic ear

and sound advice none of this could have been

done; and Dr. Daniel Cohen, of the NMFS Systematics Laboratory, who kindly read the manuscript

and offered valuable suggestions. I am deeply

grateful to those individuals and institutions who

loaned me specimens for comparison.

HISTORICAL REVIEW.—The first major report on

the Asteroidea of the Caribbean and Gulf of Mexico was published by Edmond Perrier, in 1884, on

the starfishes collected by the Blake between 1877

and 1879. Collections were made from Key West

to Havana, along the north coast of Cuba, Tortugas to Yucatan Bank at Cape Catoche and Cape

San Antonio, and off the mouth of the Mississippi

River during the first expedition (1877-1878). During the second expedition (1878-1879), the Blake

collected from Key West to Havana, Jamaica, the

Bahama Straits, the Windward Passage, St. Thomas,

Haiti, Puerto Rico, St. Croix, Saba Bank, Montserrat, St. Kitts, Guadeloupe, Dominica, Martinique,

St. Lucia, St. Vincent, Grenada, the Grenadines,

and Barbados. In all, 289 dredge hauls were made,

in depths varying from 14 to 2,412 fathoms. From

these collections, Perrier described 54 species, 46 of

them new.

W. Percy Sladen (1889), reporting on the Asteroidea collected by the Challenger expedition, included in his report 94 tropical-subtropical western

Atlantic species and extended the range of many

others.

The University of Iowa's Bahamas expedition of

1893 resulted in a report by A. E. Verrill (1915)

on the starfishes of the West Indies, Florida, and

Brazil. This comprehensive study covered mainly

the shallow-water Asteroidea (87 nominal species

and subspecies) and also listed 44 deepwater species.

Although it continues to be a useful work for the

area, it is somewhat outdated by nomenclatural

changes, the discovery of new forms, and the limited region covered.

In his Handbook of the Littoral Echinoderms of

Puerto Rico, H. L. Clark (1933) lists 17 nominal

species of Asteroidea from the West Indies, mostly

from shallow water. In 1941, he published a report

on the echinoderms of the West Indies collected by
NUMBER 126

the Havana-Harvard Atlantis expeditions which included 16 new species. Clark pointed out that 21

of the species were represented in the collections

by a single specimen.

Other reports on Caribbean and Gulf of Mexico

starfishes exist, of course, but they are generally

limited in area, or include species from a much

wider area, or are concerned only with certain

families. Reference to these reports will be found

in the literature cited.

Following is a list of the species in this and the

five above-mentioned collections (1, this report; 2,

Perrier, 1884; 3, Sladen, 1889; 4, Verrill, 1915; 5,

Clark, 1933; 6, Clark, 1941; italicized names are

junior synonyms, indented under valid name):

Species 1 2 1 4 i 6 Page

Luidia alternata x x x x x 23

L. alternata bicolor x

L. variegata x

Luidia barbadensis x x x x x 24

L. convexiuscula x x x

Luidia clathrata x x x x x 22

Luidia elegans x x x 25

Luidia sagamina x 24

Luidia senegalensis x x x 22

L. marcgravii x

Astropecten alligator x x x

Astropecten atnericanus x x 27

Archaster americanus x

Astropecten antillensis x x x

Astropecten articulatus x x x x x x 28

A. articulatus dubius x

Astropecten brasiliensis x x

Astropecten ciliatus x

Astropecten cingulatus x x 28

Astropecten comptus x x 29

Astropecten duplicatus x x x x 29

Astropecten marginatus x 29

Astropecten nitidus x x 30

A. nitidus forcipatus x

Astropecten nuttingi x

Astropecten richardi x

Psilaster cassiope x 31

P. squameus X

Psilaster florae

Archaster florae x

Psilaster sp., cf florae x

Psilaster patagiatus x 32

Persephonasler spinulosus x

Persephonaster echinulatus x x 33

Persephonaster leptactis x

Persephonaster pulcher x

Archaster pulcher x

Plutonaster pulcher x

Pontaster pulcher x

Species 1 2 3 4 5 6 Page

Plutonaster bifrons x

Plutonaster efflorescens x

Archaster efflorescens x x

Plutonaster intermedius x x x 34

Goniopecten intermedius x

Dytaster insignis x x 35

Archaster insignis x x

Dytaster exilis carinatus x

D. grandis x

D. madreporifer x

Dytaster sp., cf robustus x

Blakiaster conicus x x x x 36

Tethyaster grandis x 37

Sideriaster grandis x

Tethyaster vestitus

Astropecten vestitus x

Sideriaster vestitus x

Dipsacaster antillensis x 38

Leptychaster sp x

Goniopecten demonstrans x x x x x 38

Prionaster elegans x x x 39

Ctenodiscus crispatus x

Benthopecten simplex x x 40

Archaster simplex x

Pararchaster simplex x

Benthopecten spinosus x x

Cheiraster echinulatus x x 42

Archaster echinulatus x

Pontaster echinulatus x

Cheiraster enoplus x x 42

Luidiaster enoplus x

Cheiraster rairabilis x x x 41

Archaster coronatus x

Archaster mirabilis x

Cheiraster coronatus x

Pontaster coronatus x

Cheiraster mirabilis coronatus . x

Pontaster mirabilis x

Cheiraster planus x

Pectinaster forcipatus

Pontaster forcipatus x

Pectinaster dispar x

Pectinaster gracilis x x 44

Pectinaster mixtus x x 43

Luidiaster mixtus x

Pectinaster oligoporus x

Pectinaster vincenti x

Luidiaster dubius x

Odontaster hispidus x x x 45

Odontaster setosus x 45

Odontaster sp.. cf setosus x

Odontaster sp. (compact species) x

Goniaster tessellatus x 47

G. americanus x

Pentagonaster semilunatus .... x

Ceramaster grenadensis x x x 49

Pentagonaster affinis x x x

Pentagonaster grenadensis x x
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Species 12)45 6 Page

Sphaerodiscus bourgeti

Astrogonium bourgeti x

Cladaster rudis x

Plinthaster dentatus x x x 52

P. comptus x

P. nitidus x

Pentagonaster dentatus x x

Pyrenaster dentatus x

Tessellaster notabilis x x 53

Tosia parva x 54

Pentagonaster parvus x x

Anthenoides piercei x x x x x 48

Hippasteria caribbaeus x

Pseudarchaster gracilis x 59

P. concinnus x

Pseudarchaster granuliferus x

Pseudarchaster hispidus x

Pseudarchaster ordinatus x

Pseudarchaster parelii x

Paragonaster formosus

Archaster formosus x

Paragonaster grandis x

Paragonaster subtilis x x 57

Goniopecten subtilis x

Mediaster agassizi x

Mediaster bairdii

Archaster bairdii x

Mediaster pedicellaris x x

Goniodiscus pedicellaris x x

Nymphaster arenatus x x x x

N. basilicus X

N. ternalis x x

Pentagonaster arenatus x

Pentagonaster ternalis x

Nymphaster subspinosus x x x x

Pentagonaster subspinosus x

Asteroceramus brachyactis x

Circeaster americanus x

C. occidentalis x

Litonotaster intermedius x x

Pentagonaster intermedius x x x

Peltaster nidarosiensis x x

P. hebes x

Peltaster placenta x

Peltaster planus x

Rosaster alexandri x x x

Pentagonaster alexandri x x

Oreaster reticulatus x x x x

Pentaceros reticulatus x

Ophidiaster alexandri x x

Ophidiaster guildingii x x x x

Ophidiaster pinguis x

Fromia sp x

Linckia bouvieri x x x

Linckia guildingii x x x x

Linckia nodosa x x x

Narcissia trigonaria x x x

Drachmaster bullisi x

58

59

55

56

50

51

54

60

68

67

66

67

69

63

Species 1 2 3 4 5 6 Page

Tamaria fioridae x 64

Ophidiaster fioridae x x x

Tamaria halperni x 65

Tamaria passiflora x 65

Chaetaster longipes x x

Chaetaster nodosus x x x 63

Tremaster laevis x

Radiaster elegans x x x

Solaster abyssicola x

Solaster caribbaeus x x x 71

Solaster notophrynus x 71

Laetmaster spectabilis

Ctenaster spectabilis x x x

Lophaster furcifer x

Lophaster verrilli x 72

Lophaster radians x

Korethraster radians x

Korethraster hispidus

Remaster palmatus x x

Korethraster palmatus x x

Pteraster acicula x 79

Pteraster caribbaeus x x x x x 77

Pteraster militaroides x x 78

Pteraster personatus x 76

Pteraster rugosus x x 77

Pteraster stoibe x

Pteraster sp x 79

Calyptraster coa x x

Calyptraster personatus x 80

Hymenaster anomalus x 75

Hymenaster modestus x x 74

Hymenaster pellucidus x

Hymenaster regalis x

Hymenaster rex x 73

Hymenaster sp. 1 x

Hymenaster sp. 2 x

Asterina folium x x

Asterinides folium x

Asterina hartmeyeri x

Asterina lymani x x

Asterinopsis lymani x

Asterina minuta x

Asterina pilosa x x

Asterinopsis pilosa x

Stegnaster wesseli x x x

Asterina wesseli x

Asterinides modesta x

Enoplopatiria marginata x

Enoplopatiria siderea x

Leilaster radians x 80

Marginaster pectinatus x x x x x 82

Poraniella echinulatus x

Marginaster echinulatus x x

Poraniella regularis x x 81

Porania austera x

Porania grandis x

Henricia antillarum x x x 84

Cribrella antillarum x x
NUMBER 126

Species 1 2 3 4 5 6 Page

Henricia microspina x x

Henricia sexradiata x x x 83

Cribrella sexradiata x x

Henricia sp x 85

Echinaster brasiliensis x x x 88

Echinaster echinophorus x x x 86

Echinaster modestus x x x x x 87

Echinaster sentus x x x x x 87

Echinaster spinosus x

Echinaster serpentarius x x 86

Thyraster serpentarius x

Echinaster sps. A, B, C x 88-89

Verrillaster spinulosus x 89

Echinaster spinulosus x x x

Zoroaster fulgens x x x 97

Zoroaster ackleyi x x x x

Zoroaster diomedeae x

Mammaster sigsbeei x x x 97

Zoroaster sigsbeei x x

Doraster constellatus x 95

Asterias crassispina x

Asterias forbesii x x 93

Asterias tanneri x

Coscinasterias tenuispina x x 93

C. tenuispina atlantica x

Asterias (S.) tenuispina x

Stolasterias tenuispina x

Leptasterias hartii x

Leptasterias mexicana x

Sclerasterias contorta x 92

Asterias angulosa x x

Asterias contorta x x

Asterias linearis x x

Orthasterias contorta x

Coscinasterias linearis x

Orthasterias subangulosa x

Stephanasterias gracilis x

Asterias gracilis x x

Stephenasterias hebes x

Pedicellaster pourtalesi x x x x

Ampheraster alaminos x 91

Tarsaster fascicularis

Asterias fascicularis x x

Leptasterias fascicularis x

Coronaster briareus x x x 94

Asterias briareus x

Brisinga sp., cf costata x

Brisinga sp., cf elegans x

Craterobrisinga cricophora

Brisinga cricophora x x

Odinia antillensis x 99

Odinia pandina x

Freyella elegans x

Freyella trispinosa x

Freyella sp., cf sexradiata x

Midgardia xandaros x 99

Hymenodiscus agassizi x x x

Porcellanaster caerulus x

ZOOGEOGRAPHY.—Although at first glance it would

appear that over half of the species reported from

the Caribbean and the Gulf of Mexico are restricted to the area, such a conclusion would be

hasty on the basis of our present knowledge of

the Asteroidea and their distribution. Many of the

so-called endemic Gulf and Caribbean species are

known from few specimens, or even single specimens. The starfish fauna of Cuba comprises more

than sixty species and includes at least five valid

endemic species. Jamaica, on the other hand, separated from Cuba by the Cayman Trench, has a

very depauperate asteroid fauna, with less than a

dozen species, all common and widespread in the

tropical western Atlantic. The eastern extension

of the Cayman Trench separates Cuba and HaitiDominican Republic (the Island of Hispaniola),

but so little collecting has been done, at least in

recent years, around Hispaniola that its asteroid

fauna is practically unknown. Other major unexplored (or little-known faunistically) areas in the

region are the three basins of the Caribbean: the

Yucatan Basin, the eastern Colombian Basin, and

the Venezuelan Basin.

The order Platyasterida, with one family and one

genus, is well represented in the Gulf and Caribbean, as would be expected from this circumtropical shallow-water order, by nine species.

The order Paxillosida is also well represented;

three of the four families occur in the area. By

far the largest number of paxillosid species are

moderate to deepwater forms. Two families, the

Porcellanasteridae and the Benthopectinidae, are

found only in deep water, the Benthopectinidae

below 100 fathoms and the Porcellanasteridae below 500 fathoms. Benthopectinids, represented by

six species, of three genera, are abundant in the

deep waters of the Gulf and Caribbean, but no

specimens of porcellanasterids have been reported.

Madsen (1961) stated that, even when ripe, the

gonads of the Porcellanasteridae are quite small

and the eggs few. This would indicate either direct development or a demersal larval stage (see

also Pearse, 1969), which would certainly limit distribution; it is possible that the porcellanasterids

have been unable to cross the shallow rim of the

Gulf. This would not limit the Benthopectinidae

if, as has been suggested (Spencer and Wright,

1966, and others), the dorsal muscle bands on the
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

7P <? 6,0 55

• Bureau »f Csaaereial Fisheries Statlras

o Texas MM Umirersitj St«ti»a»

FIGURE 1.—Map showing stations at which Asteroidea were collected.

arms (present only in this family) enable them to

swim.

The other two paxillosid families, Astropectinidae (eight genera, fifteen species) and Goniopectinidae (two genera, two species), are mainly from

moderately deep waters, with the exception of the

abundant genus Astropecten, with seven species in

the area, which has much the same distribution as

the Luidiidae.

Of the seven families of the order Valvatida, five

are represented in the Gulf and Caribbean. However, by far the greatest number of species belong

to two families, Goniasteridae (15) and Ophidiasteridae (10). The goniasterids are widespread in

moderately deep waters all over the world, with the

exception of the polar and subpolar regions. Ophidiasterids are mainly tropical-subtropical shallowwater starfish, well represented in both the

Caribbean and the Indo-Pacific. Four of the other

five valvatid families are small, with a fairly limited distribution; the remaining family, Oreasteridae, is principally Indo-Pacific in distribution, and

is represented in the western Atlantic by only one

species, Oreaster reticulatus. Possibly this species

is either a Tethyan relict or a recent arrival in the

Caribbean; it does not appear to enter the Gulf of

Mexico.

Only six of the thirteen families in the order

Spinulosida occur in the Gulf and Carribean. The

Solasteridae, mostly a boreal family, are represented by two or three genera of rare occurrence.

The deepwater Pterasteridae, with nine species, are

the best represented spinulosid family; nearly half

of the known genera occur in the western Atlantic.

The Ganeriidae occur mostly in the Southern Hemisphere, with only one (doubtful) representative in
NUMBER 126

the tropical western Atlantic. The Poraniidae, a

mainly boreal, deepwater family, are represented

by two genera in the Gulf and Caribbean; both

include the smallest starfishes known, although

their more northern relatives reach a considerable

size. The family Echinasteridae is again mostly

boreal, in moderate depths, but the genus Echinaster is abundantly represented in the Gulf and

Caribbean in shallow water by eight species; most

of the known species of this genus occur here. The

cosmopolitan family Asterinidae is abundantly represented in tropical and subtropical shallow waters,

but surprisingly absent from the present collections; however, at least five species have been reported from the Gulf and Caribbean in the past,

two of them apparently endemic to Cuba.

The deepwater order Zorocallida (one family)

is well represented in the western Atlantic, nearly

half (3) of the known genera occurring here; the

rest are from the North Pacific and the Indian

Ocean. Two of the three species in these collections are apparently confined to the Gulf of

Mexico.

The order Forcipulatida is poorly represented

in the Gulf and Caribbean. The small family

Heliasteridae does not occur here. The very large

family Asteriidae, with nearly sixty known genera,

is represented in the area by only six or seven

genera; most of the species in these genera have

ranges extending to either the northern Atlantic

or the eastern Atlantic.

The order Euclasterida, of very wide distribution in deep waters of the world, is well represented in the Gulf and Caribbean by five of the

sixteen or seventeen known genera.

At least a dozen of the 95 species in these collections have never been taken in the Gulf of Mexico,

while about seven species have been reported only

from the Gulf. It would be premature to draw any

conclusions about the existence of a Gulf fauna, or

a Caribbean fauna, on the basis of our present

knowledge of asteroid distribution.

The greatest number of species (62) occurs at

depths of 50 to 200 fathoms. However, larger populations of fewer species probably occur in waters

below 1,000 fathoms, as this depth yields the greatest number of specimens per dredge haul.

In the present collections, all of the stations below 500 fathoms were occupied in the Gulf of

Mexico; the following deepwater species have been

collected in the Gulf, and undoubtedly occur also

in the deep basins of the Caribbean, which are

as yet largely unexplored.

Plutonaster intermedius (200-1000 fens)

Dytasler insignis (200-2000+ fms)

Dipsacaster antillensis (1000-2000-f- fms)

Benthopecten simplex (1000-2000 fms)

Litonotaster intermedius (1000-2000 fms)

Paragonaster subtilis (1000-2000 fms)

Hymenaster rex (1000-2000 fms)

Hymenaster modestus (500-2000 fms)

Hymenaster anomalus (1000-2000 fms)

Pteraster personatus (500-1000 fms)

Zoroaster fulgens (200-2000-j- fms)

TABLE I.—Depth ranges of Gulf and Caribbean Asteroidea (the number of stations at which each species

was taken is given, with number of specimens in parentheses)

Species Depth ranges in fathoms (total number of stations in parentheses)

0-25 25-50 50-100 100-200 200-500 500-1000 1000-2000 2000-f

(41) (43) (22) (37) (85) (16) (17) (S)

Luidia senegalensis

Luidia clathrata

Luidia alternata

Luidia sagamina

Luidia barbadensis

Luidia elegans

Astropecten americanus

Astropecten articulatus

Astropecten cingulatus

Astropecten comptus ....

Astropecten duplicatus

Astropecten marginatus

Astropecten nitidus

2(5)

6(10)

4(4)

3(4)

2(10)

6(35)

3(4)

3(3)

1(1)

5(10)

1(1)

1(12)

1(1)

1(1)

1(1)

3(7)

3(5)

5(18)

1(1)

1(8)

1(2)

6(18)
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 1 .—Continued

Species

Psilaster cassiope

Psilaster patagiatus

Persephonaster echinulatus

Plutonaster intermedius

Dytaster insignis

Blakiaster conicus

Tethyaster grandis

Dipsacaster antillensis

Goniopecten demonstrans

Prionaster elegans

Benthopecten simplex

Cheiraster mirabilis

Cheiraster echinulatus .

Cheiraster enoplus

Pectinaster mixtus

Pectinaster gracilis

Odontaster hispidus

Odontaster setosus

Goniaster tessellatus

Anthenoides piercei

Ceramaster grenadensis

Peltaster nidarosiensis

Peltaster placenta

Plinthaster dentatus

Tessellaster nobilis

Tosia parva

Rosaster alexandri

Circeaster americanus

Litonotaster intermedius

Paragonaster subtilis

Nymphaster arenatus

Nymphaster subspinosus

Pseudarchaster gracilis

Oreaster reticulatus

Chaetaster nodosus

Drachmaster bullisi

Tamaria floridae

Tamaria halperni

Tamaria passiflora

Linckia guildingi

Linckia nodosa

Linckia bouvieri

Ophidiaster guildingii

Xarcissia trigonaria

Solaster notophrynus

Solaster caribbaeus

Lophaster verrilli

Hymenaster rex

Hymenaster modestus

Hymenaster anomalus

Pteraster personatus

Pteraster caribbaeus

Pteraster rugosus

Pteraster militaroides

0-25

(41)

5(10)

1(2)

6(7)

1(1)

Depth

25-50

(43)

2(4)

10(72)

2(3)

1(1)

1(1)

2(2)

1(4)

1(2)

6(13)

10)

1(1 )

ranges in fathoms (total

50-100

(22)

1(1)

1(9)

1(2)

3(5)

10)

1(1)

1(2)

2(2)

1(1)

100-200

(37)

1(1)

1(1)

3(6)

1(1)

1(1)

2(7)

1(2)

1(1)

6(7)

1(1)

2(7)

1(1)

1(1)

3(3)

1(1)

1(1)

1(1)

5(8)

1(1)

2(2)

2(2)

number

200-500

(85)

5(7)

2(5)

13(30)

5(23)

2(7)

12 (27)

7(11)

2(2)

1(1)

1(1)

1(1)

1(1)

1(1)

23 (36)

7(11)

30 (80)

1(1)

11(23)

1(3)

1(1)

1(1)

1(1)

1(1)

3(3)

of stations

500-1000

(16)

4(26)

2(3)

4(9)

1(1)

1(1)

8(26)

1(3)

1(1)

in parentheses)

1000-2000 2000 +

(17) (?)

7 (18) 2 (3)

1 (1) 1 (1)

4(15)

1(1)

4(11)

2(6)

1(1)

1(1)

1(2)

1(1)
NUMBER 126

TABLE 1 .—Continued

Species Depth ranges in fathoms (total number of stations in parentheses)

0-25

(41)

25-50

(43)

5O-100

(22)

100-200

(37)

200-500

(85)

500-1000 1000-2000 2000 +

(16) (17) (S)

Pteraster acicula

Pteraster species

Calyptraster personatus

Leilaster radians

Poraniella regularis

Marginaster pectinatus ....

Henricia sexradiata

Henricia antillarum

Henricia species

Echinaster serpentarius

Echinaster echinophorus

Echinaster sentus

Echinaster modestus

Echinaster species A.

Echinaster brasiliensis

Echinaster species C.

Echinaster species B

Verrillaster spinulosus

Ampheraster alaminos

Sclerasterias contorta

Asterias forbesi

Coscinasterias tenuispina

Coronaster briareus

Doraster constellatus

Mammaster sigsbeei

Zoroaster fulgens

Odinia antillensis

Midgardia xandaros

10)

6(20)

1(1)

1(2)

1(1)

1(1)

1(1)

1(6)

1(2)

1(1)

1(1)

1(1)

4(8)

1(1)

4(13)

2(2)

4(6)

1(1)

1(8)

7(8)

1(1)

1(1)

1(1)

1(1)

2(3)

1(5)

1(1)

1(1) 2(2)

1(1)

4(12)

1(6)

1(5)

2(12)

1(1)

4(5)

2(11)

12 (40)

2(3)

1(1)

2(4) 3(6)

Station Data

Oregon Stations

2. 28°0O/ N, 94°49/ W. 50-52 fms. May 1950.

Bottom temperature: 69.2. Pacific otter trawl.

Astropecten cingulatus

33. 25°55' N, 83°55' W. 62 fms. June 1950.

Bottom temperature: 61.5. 55' shrimp trawl.

Pectmaster mixtus

35. 25°35' N, 83°42' W. 60 fms. June 1950.

Bottom temperature: 62.6. 55' shrimp trawl.

Henricia species

36. 28°30' N, 85°36' W. 120 fms. June 1950.

Bottom temperature: 54.3. 55' shrimp tr*wl.

Henricia sexradiata

102. 29°07.5' N, 88-50.5' W. 45 fms. October 1950.

Bottom temperature: 64.4. 100' Texas trawl.

Astropecten cingulatus

123. 28-46.5' N, 89845' W. 40 fms. September 1950.

Bottom temperature: 66.2. 100' Texas trawl.

Astropecten cingulatus

150. 27°43' N, 96°51' W. 15 fms. November 1950.

Bottom temperature: 66.8. 40* flat trawl.

Astropecten duplicatus

272. 29°16' N, 86°39' W. 216 fms. February 1951.

Bottom temperature: 50. 40* flat trawl.

Odontaster setosus

382. 29O11.5' N, 88-07.5' W. 190-210 fms. June 1951.

Bottom temperture: 83.5. 100' flat trawl.

Astropecten americanus

Doraster constellatus

Zoroaster fulgens

489. 27-44' N, 85°09' W. 254 fms. September 1951.

Bottom temperature: 825.

Plinthaster dentatus

Nymphaster arenatus

Henricia antillarum

491. 27°49' N, 84°59' W. 200 fms. September 1951.

Bottom temperature: 52.2. 40* flat trawl.

Henricia antillarum

1408. 28-02' N, 90° 15' W. 200 fms. October 1955.

Bottom temperature: 48.9. 40' flat trawl.

Odinia antillensis

1416. 28-06' N, 91°02' W. 29 fms. October 1955.

Bottom temperature: 71:6. 4' shell dredge.
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Chaetaster nodosus

1495. 28°48' N, 89°26' W. 45 fms. May 1956.

Bottom temperature: 65.3. 40' flat trawl.

Astropecten cingulatus 1920.

1505. 27°48' N, 94°55' W. 250 fms. May 1956.

Bottom temperature: 50. 40* flat trawl.

Nymphaster arenatus

1507. 27°43' N, 95°05' W. 300 fms. May 1956. 1929.

4(y flat trawl.

Persephonaster echinulatus

Cheiraster mirabilis 1934.

Plinthaster dentatus

1508. 27°40' N, 95°35' W. 200 fms. May 1956.

Bottom temperature: 53.1. 40* flat trawl.

Plinthaster dentatus 1936.

1509. 27°38' N, 95°35' W. 225 fms. May 1956.

Bottom temperature: 49.8. 80' balloon trawl.

Astropecten americanus 1937.

1510. 27°40' N, 95°45' W. 100 fms. May 1956.

Bottom temperature: 61.2. 40' flat trawl.

Luidia elegans

1513. 27°51' N, 94°53' W. 100 fms. May 1956.

Bottom temperature: 61.2. 40' flat trawl. 1938.

Luidia elegans

1514. 27°50' N, 94°45' W. 130 fms. May 1956.

Bottom temperature: 59. 40' flat trawl.

Luidia elegans 1946.

Astropecten nitidus

1537. 24°29' N, 83°27' W. 212 fms. June 1956.

Bottom temperature: 50.4. 72' balloon trawl. 2000.

Zoroaster fulgens

1538. 24°29' N, 83°32' W. 200 fms. June 1956.

Bottom temperature. 48.7. 72' balloon trawl. 2005.

Zoroaster fulgens

1648. 30°ll' N, 86°55' W. 17 fms. January 1957.

52-70 eastern trawl. 2027.

Astropecten comptus

1672. 29°41' N, 85°34' W. 135 fms. February 1957.

8' scallop dredge. 2164.

Echinaster serpentarius

1865. 16°2(y N, 83°2O' W. 42 fms. August 1957.

40' flat trawl. 2199.

Luidia clathrata

Echinaster brasiliensis

1885. 16-54' N, 81-18' W. 250 fms. August 1957. 2202.

40* flat trawl.

Nymphaster arenatus

Pteraster militaroides 2205.

1890. 16-35' N, 80-55' W. 100 fms. August 1957.

40' flat trawl.

Astropecten nitidus 2207.

1906. 12-19' N, 82°27' W. 325 fms. August 1957.

40' trawl.

Cheiraster mirabilis 2209.

1909. 12-35' N, 82-19' W. 350 fms. September 1957.

40' trawl.

Persephonaster echinulatus 2226.

Cheiraster mirabilis

1917. 13°20' N, 82°O2' W. 325 fms. September 1957.

40* shrimp trawl. 2240.

Persephonaster echinulatus

Cheiraster mirabilis

Pteraster militaroides

13-31' N, 81°54' W. 300 fms. September 1957.

40' shrimp trawl.

Persephonaster echinulatus

Nymphaster arenatus

13°22' N, 82-04' W. 300 fms. September 1957.

40* shrimp trawl.

Pteraster militaroides

16-00' N, 82°05' W. 21 fms. September 1957.

40' shrimp trawl.

Oreaster reticulatus

Echinaster sentus

15°54' N, 82°06' W. 21 fms. September 1957.

40' shrimp trawl.

Echinaster sentus

16-02' N, 82-07' W. 21 fms. September 1957.

40' shrimp trawl.

Luidia alternata

Oreaster reticulatus

Echinaster sentus

16-05' N, 82°05' W. 24 fms. September 1957.

40' shrimp trawl.

Oreaster reticulatus

Echinaster sentus

16-42' N. 82°36' W. 300 fms. September 1957.

40' trawl.

Nymphaster arenatus

07°34' N, 54-50' W. 45 fms. November 1957.

45' Ballerina.

Echinaster brasiliensis

07-34' N, 54-50' W. 200 fms. November 1957.

40* flat trawl.

Nymphaster arenatus

07-12' N, 53°02' W. 225 fms. November 1957.

40* flat trawl.

Anthenoides piercei

26-37.5' N. 82-53.5' W. 18-19 fms. April 1958.

41' midwater trawl.

Astropecten articulatus

24-42' N, 92-18' W. 2000 fms. June 1958.

40' flat trawl.

Dytaster insignis

28°52' N, 88°11' W. 625 fms. June 1958.

40' flat trawl.

Zoroaster fulgens

27-40' N, 82-50' W. 5-6 fms. July 1958.

40' midwater trawl.

Echinaster serpentarius

09-58' N, 61-11' W. 20-22 fms. August 1958.

40* flat trawl.

Luidia clathrata

09-48' N, 60-47' W. 20-22 fms. August 1958.

40* flat trawl.

Astropecten marginatus

08-32' N, 59-05' W. 23-33 fms. August 1958.

W flat trawl.

Astropecten marginatus

07°56' N, 58-20' W. 11 fms. August 1958.
NUMBER 126 11

40* flat trawl.

Echinaster brasiliensis

2243. 08°03' N, 58°24' W. 23-25 fans. August 1958. 2575.

40* flat trawl.

Astropecten duplicatus

2245. 08° 15' N. 58° 17' W. 39-50 fras. August 1958. 2643.

40* flat trawl.

Goniaster tessellatus

Echinaster brasiliensis 2644.

2248. 07°45' N, 57°34' VV. 30-35 fms. August 1958.

40' flat trawl.

Narcissia trigonaria 2650.

2249. O7°4O' N, 57°34' W. 27-30 fms. August 1958.

40* flat trawl.

Echinaster brasiliensis 2652.

2250. 07°38' N. 57°34' W. 26-27 fms. August 1958.

4C flat trawl.

Narcissia trigonaria 2771.

2267. 06°58' N. 56°02' W. 25 fms. September 1958.

40* flat trawl.

Echinaster brasiliensis

2272. 06°30' N, 55°52' W. 17 fms. September 1958. 2774.

40* flat trawl.

Luidia senegalensis

Luidia clathrata

2289. 07°25' N, 54°35' W. 75-80 fms. September 1958. 2775.

40* flat trawl.

Henricia sexradiata

2329. 06°40' N, 54°25' W. 25-27 fms. September 1958.

40* flat trawl. 2776.

Echinaster brasiliensis

2343. O7°34' N, 57°45' W. 19-20 fms. September 1958.

W flat trawl. 2779.

Echinaster brasiliensis

2345. 08° 12' N, 58°25' W. 34-35 fms. September 1958.

40' flat trawl. 2780.

Echinaster brasiliensis

2353. 11°35' N, 62°41' W. 212-250 fms. September 1958.

40* flat trawl. 2813.

Nymphaster arenatus

2547. 29° 15' N, 88°05' W. 100 fms. May 1959.

40' midwater trawl. 2814.

Ampheraster alaminos

2566. 26°24' N, 90°45' W. 1450-1500 fms. July 1959.

4C flat trawl. 2820.

Postlarval astropectenid

2567. 25°21' N, 91°02' W. 1725-1750 fms. July 1959.

30' balloon.

Calyptraster personatus

2571. 26°20' N, 90°31' W. 1300 fms. July 1959. 2821.

4C flat trawl.

Benthopecten simplex

Zoroaster fulgens

2572. 26°53' N, 89°56' W. 1325 fms. July 1959. 2824.

4C flat trawl.

Dytaster insignis

Benthopecten simplex

2574. 26°34' N, 89°53' W. 1450 fms. July 1959. 3210.

4C flat trawl.

Benthopecten simplex

Ampheraster alaminos

Zoroaster fulgens

27°06' N, 89°13' W. 1100-1200 fms. July 1959.

40' flat trawl.

Brisingid discs, unidentified

18°03' N, 64°27' W. 150-180 fms. October 1959.

40' semiballoon trawl.

Tamaria halperni

18° 12' N, 67°42' W. 260 fms. October 1959.

40' semiballoon trawl.

Henricia species

18° 16' N, 67°17' W. 250 fms. October 1959.

40' semiballoon trawl.

Zoroaster fulgens

18°18' N, 67-18.5' W. 300 fms. October 1959.

40' semiballoon trawl.

Zoroaster fulgens

11-40' N, 62"27' W. 220 fms. April 1960.

40* trawl.

Plinthaster dentatus

Pseudarchaster gracilis

11-32' N, 62-40' W. 195-212 fms. April 1960.

40* trawl.

Persephonaster echinulatus

Nymphaster arenatus

11-35' N, 62-37' W. 220-230 fms. April 1960.

40* trawl.

Plinthaster dentatus

Mammaster sigsbeei

11-36' N, 62-42' W. 235 fms. April 1960.

40* trawl.

Plinthaster dentatus

11-35' N, 62-59' W. 240-260 fms. April 1960.

40* trawl.

Zoroaster fulgens

11-36' N, 62-52' W. 215-230 fms. April 1960.

40* trawl.

Doraster constellatus

28°48' N, 87°57' W. 1000 fms. July 1960.

Trawl.

Dytaster insignis

28°53' N, 87-47' W. 950-1050 fms. July 1960.

Trawl.

Nymphaster arenatus

28°23' N, 88-21.5' W. 1000 fms. July 1960.

Trawl.

Plutonaster intermedius

Nymphaster arenatus

Hymenaster modestus

28-47.5' N, 87O57' W. 1000-1100 fms. July 1960.

Trawl.

Dytaster insignis

Hymenaster modestus

29°07.5' N, 88-04' W. 365-395 fms. July 1960.

Trawl.

Plinthaster dentatus

Zoroaster fulgens

29°50' N, 87°08' W. 66 fms. February 1961.

5' scallop dredge.

Verrillaster spinulosus
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

3555. 17° 17' N, 78°23' W . 15 fms. May 1962.

5' dredge. 4302.

Luidia clathrata

Luidia alternata

Astropecten duplicates 4392.

3560. 16°35' N, 80° 10* W. 315 fms. May 1962.

Bottom temperature: 45. 40' shrimp trawl.

Nymphaster arenatus 4398.

Zoroaster fulgens

3561. 16°35' N, 80°04' W. 400 fms. May 1962.

Bottom temperature: 46. 40' shrimp trawl.

Plinthaster dentatus

Zoroaster julgens 4413.

3573. 14° 18' N, 81°44' W. 410-420 fms. May 1962.

Bottom temperature: 41. 40' shrimp trawl.

Plinthaster dentatus

3574. 12°31' N, 82°21' W. 200 fms. May 1962. 4416.

40* shrimp trawl.

Nymphaster arenatus

3583. 09° 16' N, 81°37' W. 280 fms. May 1962.

40* shrimp trawl.

Persephonaster echinulatus

Doraster constellatus

3601. 09°07' N, 81MC W. 400 fms. May 1962. 4421.

Bottom temperature: 43. 40* shrimp trawl.

Plinthaster dentatus

3603. 12° 16' N, 82°54' W. 15-20 fms. June 1962.

5' dredge. 4459.

Astropecten duplicates

Oreaster reticulates

3608. 12°28' N, 82°28' W. 110 fms. June 1962. 4472.

40/ shrimp trawl.

Anthenoides piercei

3621. 16°00' N, 81°09' W. 120-130 fms. June 1962. 4474.

5' dredge.

Tethyaster grandis

3637. 17°13' N, 87°55' W. 120-170 fms. June 1962. 4480.

Bottom temperature: 51. 40* shrimp trawl.

Pectinaster mixtus

3654. 29-08.5' N, 88°00.5' W. 400-401 fms. July 1962.

Bottom temperature: 46.4. 40* flat trawl. 4574.

Psilaster cassiope

Plinthaster dentatus

Nymphaster arenatus 4702.

Zoroaster fulgens

4086. 27°46' N, 84°06' W. 31 fms. December 1962.

6' tumbler dredge.

Astropecten comptus 4703.

Coniaster tessellatus

Narcissia trigonaria

Echinaster species "B" 4731.

4089. 27-43' N, 83"34' W. 22 fms. December 1962.

6* tumbler dredge.

Astropecten duplicatus 4855.

4226. 00°18' N, 44-17' W. 150 fms. March 1963.

Bottom temperature: 47. 6' tumbler dredge.

Leilaster radians

Marginaster pectinatus

4300. 07-44' N, 54° 19' W. 300 fms. March 1963.

65' shrimp trawl. 4902.

Pseudarchaster gracilis

07-35' N, 54°25' W. 150 fms. March 1963.

6' tumbler dredge.

Astropecten americanus

12°32' N, 71°05' W. 40 fms. September 1963.

Bottom temperature: 81. 40' flat trawl.

Goniaster tessellatus

12-46' N, 70-41' W. 110 fms. September 1963.

Bottom temperature: 62. 6' tumbler dredge.

Astropecten nitidus

Pectinaster mixtus

Rosaster alexandri

11-53' N, 69-25' W. 350 fms. October 1963.

Bottom temperature: 46. 40' flat trawl.

Nymphaster arenatus

Zoroaster fulgens

11-54' N, 69-18' W. 500 fms. October 1963.

Bottom temperature: 44. 40' flat trawl.

Psilaster patagiatus

Plinthaster dentatus

Circeaster americanus

Nymphaster arenatus

Pseudarchaster gracilis

11-49' N, 69°24' W. 200 fms. October 1963.

40' flat trawl.

Anthenoides piercei

Pseudarchaster gracilis

m°bV N, 66-58' W. 53 fms. October 1963.

Bottom temperature: 73. 6' tumbler dredge.

Chaetaster nodosus

10-13' N, 65-23' W. 18 fms. October 1963.

40* flat trawl.

Luidia senegalensis

10-43' N, 64-29' W.

Bottom temperature: 63.

Astropecten duplicatus

11-10' N, 65°O7' W. 220 fms. October 1963.

40' flat trawl.

Persephonaster echinulatus

Odinia antillensis

23-13' N, 87-50' W. 210 fms. December 1963.

Bottom temperature: 50.2. 40' shrimp trawl.

Odinia antillensis

27-5O* N, 90-33' W. 400 fms. February 1964.

Bottom temperature: 45. 40' shrimp trawl.

Goniopecten demonstrans

Cheiraster mirabilis

27°55' N, 90-28' W. 300 fms. February 1964.

Bottom temperature: 50. 40' shrimp trawl.

Persephonaster echinulatus

27-35' N, 92°32' W. 250-300 fms. February 1964.

40' flat shrimp trawl.

Sclerasterias contorta

11° 16.5' N, 74-21.5' W. 400 fms. May 1964.

40' flat shrimp trawl.

Psilaster cassiope

Goniopecten demonstrans

Plinthaster dentatus

Nymphaster arenatus

09-02.4' N, 76-31' W. 400 fms. May 1964.

300 fms. October 1963.

40' flat trawl.
NUMBER 126 13

Goniopecten demonstrans 5920.

4928. 14°05' N, 81°21' W. 100 fms. June 1964.

6' tumbler dredge.

Astropecten nitidus 5923.

Tatnaria halperni

4938. 2O°31' N, 86° 12' W. 150-164 fms. June 1964.

40* flat trawl. 5929.

Linckia bouvieri

4939. 2O°25' N, 86° 13' W. 150 fms. June 1964.

6' tumbler dredge.

Linckia bouvieri

4994. 15°30' N, 63°38' W. 200-205 fms. September 1964.

40* shrimp trawl.

Pteraster rugosus

5023. 11°14' N, 60°51' W. 38 fms. September 1964. 5930.

6' tumbler dredge.

Narcissia trigonaria

Verrillaster spinulosus 5934.

5390. 17°45' N, 77°21' W. 8-9 fms. May 1965.

40' shrimp trawl.

Oreaster reticulatus 5937.

5438. 20-20* N, 69°59' W. 6-11 fms. May 1965.

6' tumbler dredge.

Ophidiaster guildingii 5970.

5456. 18°03' N, 67° 18' W. 12-14 fms. June 1965.

Bottom temperature: 82. 40' shrimp trawl.

Luidia alternata 6300.

Oreaster reticulatus

Echinaster echinophorus

5700. 12° 12' N. 72°26' W. 38 fms. October 1965.

40' shrimp trawl. 6403.

Narcissia trigonaria

5723. 09° 12' N, 76°20' W. 24 fms. October 1965.

65' shrimp trawl. 6696.

Echinaster brasiliensis

5739. 09°43' N, 79°20' W. 52 fms. October 1965.

Bottom temperature: 79. 40' shrimp trawl.

Narcissia trigonaria

Echinaster brasiliensis

5784. 24°28' N, 83°39' W. 310 fms. November 1965.

Bottom temperature: 48. 40' flat trawl.

Plinthaster dentatus

Nymphaster arenatus

5882. 28°59' N, 80°48' W. 10.5-115 fms. January 1966.

Bottom temperature: 60. 65' flat net. 6697.

Asterias forbesii

5894. 29°08.5' N, 8O°38' W. 13-14 fms. January 1966.

Bottom temperature: 61. 6' tumbler dredge.

Luidia clathrata 6699.

Luidia alternata

Astropecten articulatus

Asterias forbesii

5913. 18° 11' N, 63° 15' W. 162 fms. February 1966.

Tumbler dredge.

Astropecten nitidus 6703.

Tethyaster grandis

Cheiraster echinulatus

5915. H'lC N, 61°05' W. 900 fms. March 1966.

Tumbler dredge.

Tosia parva

15°37' N, 61°22' W. 34 fms. March 1966.

Tumbler dredge.

Astropecten duplicatus

15°36' N, 61°15' W. 40 fms. March 1966.

Bottom temperature: 78. Tumbler dredge.

Drachmaster bullisi

15°39' N. 61° 10' W. 355 fms. March 1966.

Shrimp trawl.

Goniopecten demonstrans

Plinthaster dentatus

Pseudarchaster gracilis

Solaster notophrynus

Pteraster militaroides

Zoroaster fulgens

15°38' N, 61°07' W. 442 fms. March 1966.

Shrimp trawl.

Circeaster americanus

15°31' N. 61° 12' W. 195 fms. March 1966.

Tumbler dredge.

Verrillaster spinulosus

15°34' N, 61°29' W. 50 fms. March 1966.

Tumbler dredge.

Pteraster rugosus

11° 13' N, 60°52' W. 34 fms. March 1966.

Shrimp trawl.

Drachmaster bullisi

29° 17' N, 81 "W W. 9 fms. November 1966.

Bottom temperature: 74. 60' Shrimp trawl.

Luidia clathrata

Asterias forbesii

17°24' N, 87-565' W. 143 fms. January 1967.

Bottom temperature: 59. 40* 4-seam shrimp trawl.

Anthenoides piercei

17°46' N, 62°59' W. 355-365 fms. May 1967.

Bottom temperature: 49. 40' flat net.

Persephonaster echinulatus

Goniopecten demonstrans

Cheiraster mirabilis

Ceramaster grenadensis

Peltaster placenta

Circeaster americanus

Nymphaster arenatus

Pseudarchaster gracilis

Pteraster caribbaeus

17-47' N, eS'OP W. 440-460 fms. May 1967

40' flat net.

Goniopecten demonstrans

Circeaster americanus

17-38.5' N, 62°16' W. 180-185 fms. May 1967

Bottom temperature: 60. 40' flat net.

Astropecten nitidus

Anthenoides piercei

Peltaster placenta

Tessellaster notabilis

16°53' N, 61-53' W. 410-460 fms. May 1967.

65' flat net.

Persephonaster echinulatus

Goniopecten demonstrans

Cheiraster enoplus

Circeaster americanus
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Nymphaster arenatus

Pseudarchaster gracilis

6715. 18°36' N, 63°27' W. 110-130 fans. May 1967.

Bottom temperature: 61. 6' tumbler dredge.

Blakiaster conicus

Tamaria halperni

Oregon II Stations

10184. 17°08' N, 80°28' W. 106 fms.

100/ Bat trawl.

Anthenoides piercei

10206. 12°44' N, 82° 16' W. 270 fms.

71' shrimp trawl.

Nymphaster arenatus

10212. 09°55' N. 76°OS' W. 70 fms.

15C flat trawl.

Echinaster brasiliensis

10353. 28°10' N, 92°11' W. 40 fms. January 1969.

71' shrimp trawl.

Anthenoides piercei

10491. 10°28' N, 60°04' W. 355 fans. April 1969.

40* shrimp trawl.

Plinthaster dentatus

Nymphaster arenatus

10499. 08°43' N, 59° 10* W. 40 fms. April 1969.

Bottom temperature: 79. 40* shrimp trawl.

Astropecten duplicatus

10505. 08° 13' N, W>W W. 20 fms. April 1969.

40' shrimp trawl.

Astropecten marginatus

10507. 08°10' N, 58-36' W. 20 fms. April 1969.

40* shrimp trawl.

Astropecten marginatus

10513. 08°26' N, 58° 11' W. 100 fms. April 1969.

52' flat trawl.

Rosaster alexandri

Tamaria halperni

10514. 08-06' N, 57-41' W. 202 fms. April 1969.

52' flat trawl.

Nymphaster arenatus

10517. 07-23' N, 57°04' W. 32 fms. April 1969.

40* shrimp trawl.

Narcissia trigonaria

10518. 07-03' N, 57-03' W. 20 fms. April 1969.

Bottom temperature: 81. 40* shrimp trawl.

Goniaster tessellatus

10533. 06-51' N. 55°47' W. 26 fms. April 1969.

40* shrimp trawl.

Goniaster tessellatus

10559. 06-54' N, 53°50' W. 36 fms. May 1969.

¥V shrimp trawl.

Telhyaster grandis

10561. 06-52' N, 53»46' W. 35 fms. May 1969.

40* shrimp trawl.

Echinaster brasiliensis

10602. 07-46' N, 54°35' W. 299 fms. May 1969.

Goniopecten demonstrans

Nymphaster arenatus

November 1968.

November 1968.

November 1968.

10628.

10633.

10636.

Pseudarchaster gracilis

Doraster constellatus

10619. O7°51' N, 54°23' W. 425 fms. May 1969.

100* shrimp trawl.

Zoroaster fulgens

10624. 07-30' N, 55°29' W. 440 fms. May 1969.

100' shrimp trawl.

Nymphaster arenatus

Pseudarchaster gracilis

21° W N, 85-02' W. 1 fm. June 1969.

Peltaster placenta

27'53' N, 85°13' W. 265 fms. June 1969.

204' shrimp trawl.

Plinthaster dentatus

Circeaster americanus

28-17' N, 86-21' W. 367 fms. June 1969.

204' shrimp trawl.

Plinthaster dentatus

Circeaster americanus

10667. 36-17' N, 74°49' W. 99 fms. July 1969.

40* shrimp trawl.

Odontaster hispidus

10715. 36-48' N, 74°40' W. 75 fms. July 1969.

Bottom temperature: 52. 8' tumbler dredge.

Odontaster hispidus

10728. 36-11' N, 74°49' W. 75 fms. July 1969.

Bottom temperature: 56. 8' tumbler dredge.

Odontaster hispidus

10794. 07°5O/ N. 54°15' W. 405 fms. November 1969.

191' shrimp trawl.

Nymphaster arenatus

10795. 07-48' N, 54°01' W. 392 fms. November 1969.

191' shrimp trawl.

Pseudarchaster gracilis

10802 07-26' N, 53° 16' W. 330-345 fms. November 1969.

191' shrimp trawl.

Plinthaster dentatus

10825. 15-42' N, 61°08' W. 35 fms. December 1969.

71' shrimp trawl.

Solaster caribbaeus

10827. 15-40' N. 61°09' W. 355 fms. December 1969.

71' shrimp trawl.

Peltaster nidarosiensis

Plinthaster dentatus

Pteraster species

10841. 71-24' N, 62-33' W. 352 fms. December 1969.

70' shrimp trawl.

Ceramaster grenadensis

10842 17-15' N, 62-22' W. 317 fms. December 1969.

40' shrimp trawl.

Ceramaster grenadensis

10843. 17-06' N, 62°17' W. 322 fms. December 1969.

40' shrimp trawl.

Nymphaster arenatus

10844. 17-06' N, 62°28' W. 344 fms. December 1969.

40' shrimp trawl.

Ceramaster grenadensis

Nymphaster arenatus

10849. 20-50' N, 73°23' W. 170 fms. December 1969.

Bottom temperature: 64. 40' shrimp trawl.
NUMBER 126 15

Tamaria halperni

10850. 20°49' N, 73°26' W. 170 fans. December 1969.

Bottom temperature: 64. 40' shrimp trawl.

Tamaria halperni

10858 22°59' N, 78°43' W. 152 fms. December 1969.

40* shrimp trawl.

Tamaria passiflora

10895. 23°04' N, 78°46' W. 198 fms. December 1969.

40* shrimp trawl.

Prionaster elegans

10872. 28°59' N, 87°09' W. 500 fms. January 1970.

150* TT.

Nymphaster arenatus

10876. 28°55.2' N, 87°23' W. 800 fms. January 1970.

70' shrimp trawl.

Plinthaster dentatus

Nymphaster arenatus

10877. 28°34' N, 87°26' W. 900 fms. January 1970.

70' shrimp trawl.

Nymphaster arenatus

10900. 28°50' N, 86°59' W. 400 fms. February 1970.

70' shrimp trawl.

Nymphaster arenatus

Silver Bay Stations

35. 29°20' N, 85°31' W. 25-30 fms. July 1967.

8' scallop dredge.

Luidia clathrata

Astropecten articulatus

41. 29° 19* N, %b"W W. 13 fms. July 1957.

8' scallop dredge.

Goniaster tessellatus

48. 29°00' N, 85° 15' W. 23-30 fms. July 1957.

8' scallop dredge.

Echinaster species "B"

281. 27°45' N, 95°46' W. 40 fms. February 1958.

52'4.5" nylon roller net w/8' brkt. doors.

Anthenoides piercei

961. 20°02' N, 91°53' W. 26 fms. January 1959.

54' fish trawl.

Poraniella regularis

1515. 33°37' N, 76°49 3' W. 48-55 fms. December 1959.

8' scallop dredge.

Luidia clathrata

Luidia sagamina

1564. 29°04' N, 80°45' W. 8-9 fans. January 1960.

8' scallop dredge.

Asterias forbesii

1710. 34°36' N, 75°35' W. 7.5-7 fms. March 1960.

Clam dredge.

Gonister tessellatus

Echinaster serpentarius

Asterias forbesii

1909. 29°06' N, 80°31' W. 13 fms. April 1960.

8' scallop dredge.

Echinaster modestus

1952 28°05' N, 80° 11' W. 16-16.5 fms. April 1960.

10' scallop dredge.

Luidia clathrata

Asterias forbesii

1963. 27°28' N, 80°08' W. 10-11 fms. July 1960.

8' scallop dredge.

Astropecten duplicatus

1970. 28°25' N, 80°01' W. 50 fms. April 1960.

8' scallop dredge.

Echinaster serpentarius

1972. 28°42' N, 80°08' W. 30-31 fms. April 1960.

8' scallop dredge.

Linckia guildingii

2009. 28° 15' N, 80°02' W. 40-41 fms. April 1960.

10' scallop dredge.

Goniaster tesselatus

2010. 28° 17' N, 80°01' W. 34-41 fms. April 1960.

10' scallop dredge.

Goniaster tessellatus

Linckia nodosa

Coscinasterias tenuispina

2020. 28°22' N, 80°02' W. 41-45 fms. April 1960.

10' scallop dredge.

Tosta parva

2390. 24°42' N, 80°44' W. 50 fms. October 1960.

40' balloon trawl.

Goniaster tessellatus

2432. 24°29' N, 83° 13' W. 33-35 fans. October 1960.

8' scallop dredge.

Astropecten cingulatus

2453. 23°47' N, 80°02' W. 160-180 fms. November 1960.

6' tumbler dredge.

Astropecten nitidus

Anthenoides piercei

2775. 29° 10' N, 80° 18' W. 20 fms. February 1961.

8' scallop dredge.

Astropecten comptus

Echinaster serpentarius

2863. 36°04' N, 74°49' W. 60 fms. March 1961.

40' balloon trawl.

Astropecten americanus

2926. 34°54' N, 75°32' W. 28 fms. March 1961.

6' tumbler dredge.

Goniaster tessellatus

3284. 27°55' N, 80°O3' W. 30 fms. July 1961.

Bottom temperature: 57. 8' tumbler dredge.

Goniaster tessellatus

5106. 27°59' N, 8O°O2' W. 35 fms. October 1963.

6' tumbler dredge.

Tethyaster grandis

Combat Stations

90. 28°52' N, 80°05' W. 65 fans. September 1956.

10* beam trawl.

Echinaster modestus

98. 29°40' N, 80° 17' W. 40 fms. September 1956.

40* flat trawl.

Goniaster tessellatus

180. 30°39' N, 78°39' W. 125 fms. November 1956.

40' balloon trawl.
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Astropecten nitidus

185. 29°55' N, 80° 10' W. 185 fms. November 1956.

40' flat trawl.

Odon taster setosus

Lophaster verrilli

339. 29° 19' N, 80° 18' W. 25 fms. June 1957.

10' beam trawl.

Coniaster tessellatus

Echinaster modestus

364. 34° 10* N, 75°59' W. 210 fms. June 1957.

Bottom temperature: 52.9. 40* flat trawl.

Plinthaster dentatus

450. 23°59' N, 79°43' W. 350 fms. July 1957.

10' beam trawl.

Plinthaster dentatus

Nymphaster arenatus

Nymphaster subspinosus

Mammaster sigsbeei

George M. Bowers Stations

43. 31°26' N, 8O°O5' W. 24 fms. March 1956.

Bottom temperature: 67. 4C flat trawl.

Goniaster tesselatus

85. Eleuthra, Bahamas (no data).

Echinaster species "C"

Alaminos Stations

Cruise 64-A-10

13. 27°52' N, 94°56' W. 62-96 fms. August 1964.

Tethyaster grandis

Cruise 65-A-9

15. 23°00' N, 86°48' W. 320 fms. July 1965.

Cheiraster echinulatus

Nymphaster arenatus

20. 25°00' N, 84°00' W. 67 fms. July 1965.

Luidia barbadensis

Astropecten nitidus

Cheiraster echinulatus

Anthenoides piercei

Coronaster briareus

21. 25°00' N, 84°00' W. 70 fms. July 1965.

Luidia barbadensis

Cruise 67-A-5

2H. 28°20' N, 88°22' W. 980 fms. July 1967

Nymphaster arenatus

Cruise 68-A-3

3B. 25°09/ N, 94° 11' W. 2000 fms. March 1968.

Skimmer.

Dytaster insignis

5B. 23°44' N, 95°36' W. 2100 fms. March 1968.

Dytaster insignis

7D. 23°50' N, 90°55' W. 2030 fms. March 1968.

Dytaster insignis

10B. 25°09' N. 96° 16' W. 530-550 fms. March 1968.

Zoroaster fulgens

Cruise 68-A-7

3C. 27°36' N, 87°41 5' W. 1500 fms. July 1968

Dytaster insignis

Plinthaster dentatus

Litonotaster intermedius

Ampheraster alaminos

Zoroaster fulgens

4A. 25°26.6' N. 86°06' W. 1750 fms. July 1968.

Dytaster insignis

Litonotaster intermedius

Paragonaster subtilis

Ampheraster alaminos

4E. 25°23.8' N, 86°06 7' W. 1750 fms. July 1968.

Dytaster insignis

Nymphaster arenatus

Ampheraster alaminos

9A. 29°25.5' N. 86°44' W. 200 fms. August 1968.

Astropecten americanus

10A. 29° 15 5' N, 86°55' W. 300 fms. August 1968.

Persephonaster echinulatus

Pseudarchaster gracilis

11 A. 29°14' N, 87°00' W. 400 fms. August 1968.

Psilaster patagiatus

Goniopecten demonstrans

Cheiraster echinulatus

12B. 29°21.5' N, 86°59.5' W. 500 fms. August 1968.

Dytaster insignis

Plinthaster dentatus

13A. 29°03' N, 87° 15' W. 600 fms. August 1968.

Plutonaster intermedius

Nymphaster arenatus

13D. 28°59' N, 87°23.3' W. 800 fms. August 1968.

Plutonaster intermedius

14C. 28°51' N, 87°36' W. 1150 fms. August 1968.

Hymenaster rex

15D. 29°10.3' N, 87°31.5' W. 600 fms. August 1968.

Plutonaster intermedius

15H. 29"10.5' N, 87°16' W. 500 fms. August 1968

Plutonaster intermedius

Plinthaster dentatus

Nymphaster arenatus

17B. 29°09.5' N, 87°02' W. 500 fms. August 1968.

Plutonaster intermedius

Cruise 68-A-13

1. 25°42' N, 96°04' W. 480 fms. November 1968.

Skimmer.

Psilaster cassiope

Nymphaster arenatus

Zoroaster fulgens

4. 25°38.4' N, 96° 15.3' W. 280 fms. November 1968.

Skimmer.

Persephonaster echinulatus

5. 26° 12.5' N, 96° 19.8' W. 150 fms. November 1968.

Skimmer.

Tethyaster grandis

7. 26° 17' N, 96° 18' W. 150 fms. November 1968.

Skimmer.

Astropecten americanus

8. 26° 15' N, 96°09' W. 400 fms. November 1968.

Skimmer.

Persephonaster echinulatus
NUMBER 126 17

Gonipecten demons trans

Cheiraster mirabilis

9. 25°14' N, 95°12' W. 1840 fms. November 1968.

Skimmer.

Dytaster insignis

Litonotaster intermedius

10. 25° 16' N, 95°09' W. 1850-1910 fms. November 1968.

2-meter dredge.

Litonotaster intermedius

11. 25°17' N, 95°57' W. 720-750 fms. November 1968.

Skimmer.

Nymphaster arenatus

12A. 25°31' N, 95°51' W. 580-720 fms. November 1968.

Skimmer.

Pteraster acicula

15. 27°85' N, 95°10' W. 360-470 fms. November 1968.

Skimmer.

Goniopecten demonstrans

Pseudarchaster gracilis

19. 27°44' N, 95°16' W. 200 fms. November 1968.

Skimmer.

Astropecten americanus

21. 27°37.5' N, 95°20' W. 350 fms. November 1968.

Skimmer.

Persephonaster echinulatus

Doraster constellatus

23. 27°36' N, 95°20' W. 400 fms. November 1968.

Skimmer.

Plutonaster intermedius

24. 28°29' N, 95°22' W. 480 fms. November 1968

Skimmer.

Psilaster cassiope

Plutonaster intermedius

Nymphaster arenatus

27. 27° 15' N. 95°06' W. 600-650 fms. November 1928.

Skimmer.

Plutonaster intermedius

Cruise 69-A-ll

27. 18-54' N, 94-53.8' W. 425-450 fms. August 1969

Skimmer.

Psilaster cassiope

Dytaster insignis

Goniopecten demonstrans

58. 19-02.6' N, 95-27.5' W. 260 fms. August 1969.

Large skimmer.

Midgardia xandaros

78. 21-30' N, 96-55' W. 370-400 fms. August 1969.

Skimmer.

Nymphaster arenatus

87. 21°44' N, 96°46' W. 970 fms. August 1969.

Large skimmer.

Nymphaster arenatus

Pteraster personatus

Cruise 69-A-13

4. 26-25' N, 91°55' W. 1085 fms. October 1969.

Skimmer.

Benthopecten simplex

Hymenaster anomalus

9. 24-40' N, 90-39' W. 2015 fms. October 1969.

Skimmer.

Dipsacaster antillensis

29. 25-30' N, 86-09' W. 1765 fms. October 1969.

Skimmer.

Dipsacaster antillensis

Paragonaster subtilis

Cruise 70-A-10

25. 16-43' N, 82°38' W. 235-335 fms. July 1970.

57' trawl.

Henricia antillarum

Glossary

Abactinal, dorsal, aboral=surface opposite mouth.

Aboral—see abactinal.

Aciculate spine=slender, pointed spine.

Actinal, ventral, oral=surface containing mouth.

Actinal face (of adambulacral plate) ^external surface of adambulacral plate exclusive of furrow

margin.

Actinal intermediate area, oral intermediate area=

interradial area of actinal surface.

Actinolateral plates=plates between the adambulacrals and the inferomarginals.

Actinolateral spines-see lateral spines.

Actinostome—see mouth.

Adradial, dorsolateral=the row or rows of plates

between the carina and the superomarginals.

Adambulacra, adambulacral=plate immediately

below the ambulacral plate, forming the border

of the furrow.

Alveolar pedicellaria-see sugar-tongs pedicellaria.

Alveolus=an elongate pit in a plate containing a

sugar-tongs pedicellaria.

Ambitus=the periphery of a starfish.

Ambulacra, ambulacral, ambulacral groove, furrow=channel on actinal surface containing the

lube feet; the paired series of plates lining said

channel.

Ambulacral channel-see ambulacra.

Ambulacral groove-see ambulacra.

Ampulla=the bulblike portion of the tube foot,

internal to the ambulacral plate; it may be single

or double.

Anus, anal pore=small opening in center of disc

dorsum, not present in all starfishes.

Aperture papilla=small flattened and expanded

spine or flap of tissue covering a tiny aperture

(segmental pore) of unknown function between

the adambulacrals in Pterasteridae.

Apical spines-see preoral spines.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Ann, ray=radial portion of starfish exclusive of

disc.

Bipinnaria=planktonic larval stage in the development of a starfish.

Bivalved pedicellaria, valvate pedicellaria=pedicellaria with two low, broad valves lying in a

depression in a plate and somewhat resembling

a clamshell.

Carina, carinal ridge=median ridge or row of

plates on dorsal arm.

Carinal ridge—see carina.

Carinate adambulacral=adambulacral plate with

a shoulder or ridge projecting into the furrow.

Costae, skeletal arches=rows of plates forming

arches across the arm. Found only in the Brisingidae.

Costal plates=plates of the costae.

Cribriform organs=specialized channels between

adjacent marginal plates, of unknown function.

Cruciform plates=plates with 3—5 flattened, expanded basal lobes, found usually in the Pterasteridae.

Diplacanthid=with two spines.

Disc=central portion (body) exclusive of arms.

Distal=away from the disc center.

Dorsal—see abactinal.

Dorsolateral—see adradial.

Dorsum=abactinal disc surface exclusive of marginals.

Duck-billed pedicellaria=pedicellaria - with two

valves expanded at base and tip, but with slender

shank; one valve usually larger than the other.

Found only in the Zoroasteridae.

Epioral spines, suboral spines=spines on actinal

surface of mouth plates exclusive of margin.

Epiproctal cone=a small blind tube projecting

from the center of the dorsum in some starfish.

Excavate pedicellaria-see sugar-tongs pedicellaria.

Fasciculate pedicellaria, spiniform pedicellaria=

simple pedicellaria formed by 2-5 straight or

slightly curved upright movable spines.

Fascioles=channels between series or rows of

plates, frequently lined with spinules.

Feliped pedicellaria=pedicellaria formed of a basal

piece and two valves, toothed on the margin,

resembling small clasped hands or paws.

Fissiparous=reproducing by fission.

Furrow—see ambulacrum.

Furrow spines, groove spines=spines on the furFIGURE 2.—A, Major divisions, directions, and measurements of

a starfish (schematic) [Ar=arm, D=disc, d=distal, Ir=interradial, p=proximal, R=major radius, r=minor radius], B,

Abactinal surface (schematic) [a=anus, m=madreporite, o=

ocular plate, Sm — superomarginal plates], c, Actinal surface

(schematic) [amb=ambulacra, Im=inferomarginal plates, mp

— mouth plates], D, Cross section through a starfish arm (schematic) [ad=adambulacral plates, al=actinolateral plates, am

=rambulacral plates, c = carinal plate, dl = dorsolateral (or

adradial) plates, Im=inferomarginal plates, sa=superambulacral plates, Sm=superomarginal plates, l=pointed tube feet

with simple ampulla, 2 = suckered tube feet with double ampulla. Typical starfish pedicellarie: E, crossed pedicellaria of the

type found mainly in the Brisingidae; F, sugar-tongs pedicellaria

(in alveolus) typical principally of the Ophidieasteridae

[a=erect, b=open and lying flat within the alvellus]; c,

straight pedicellaria of the type found in the Luidiidae and

Astropectinidae; H, pectinate pedicellaria found only in the

Benthopectinidae; I, feliped pedicellaria found in the order

Forcipulatida [a = frontal view, b = side view]; J, valvate pedicellaria found only in starfishes of the order Valvatida; K,

straight pedicellaria consisting of a basal piece and two valves

found in the orders Forcipulatida and Zorocallida; L, duckbilled pedicellaria found only in the Zoroasteridae.

row margin of the adambulacrals, or within the

furrow.

Glassy tubercles=small, low, transparent, pebbly

projections on some plates of certain starfish.

Granule=a small, deciduous, spherical or hemispherical external ornamentation.

Groove-see ambulacrum.

Groove spine-see furrow spine.

Gyri=irregular channels on the surface of the

madreporite.

Inferomarginal plates, inferomarginals=marginals

pertaining to the actinal surface.

Interbrachial-see interradial.

Interbrachial septum, interradial strut=an internal

interradial strut of muscle tissue or of fused

plates between the mouth and the interradial

margin.

Intermarginal=between the two series of marginals.

Interradial, interbrachial=the area of the disc between the arms.

Interradial strut-see interbrachial septum.

Jaw plate-see mouth plate.

Lateral fringe=a fringe of tissue, usually not supported by spines, around the ambitus of certain

Pterasteridae.
NUMBER 126 19

1 2

G H I K L
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Lateral spines (as in Pterasteridae)=long spines

projecting from the adarabulacrals and supporting the actinal surface in Pterasteridae.

Madreporite=:a specialized plate on the abactinal

surface, forming a sievelike opening for the water

vascular system.

Major radius—see R.

Marginal mouth spines=spines on the margin of

mouth plates exclusive of preoral spines.

Marginal plates, marginals=one or two rows of

plates generally defining the ambitus and usually

larger and more regular than the other plates.

Marginal spines=spines ornamenting the marginals.

Median furrow=suture between the paired mouth

plates.

Minor radius-see r.

Monacanthid=with one spine.

Mouth, peristome, actinostome=opening to digestive system, in center of actinal surface.

Mouth plate, jaw plate=paired plates at apex of

actinal interradial area, around the mouth.

Ocular plates, oculars=the terminal plate on the

abactinal surface of the arms.

Oscular valves=valves surrounding osculum.

Osculum=central opening in the supradorsal

membrane, found in Pterasteridae.

Oral—see actinal.

Oral intermediate-see actinal intermediate.

Oral spines=spines on the mouth plates.

Ossicle—see plate.

Papulae=respiratory organs, fingerlike or glovelike

projections of soft tissue issuing from pores between plates, and usually confined to abactinal

surface.

Papular area=area containing papulae.

Papular pores=pores through which papulae project.

Papularia=small specialized areas on abactinal surface containing papulae; usually found in Benthopectinidae.

Paxilla, paxillate, paxilliform=a more or less

columnar plate with, usually, an expanded base

and bearing spines or granules on the top.

Paxillar column—see pedicel.

Pectinate pedicellaria=pedicellaria usually lying

between two plates and formed by two opposing

rows of short, curved spines, like the teeth of a

rake. Found only in the Benthopectinidae.

Pedicel, paxillar column=the column, or shank,

of a paxilla.

Pedicellariamsmall pincerlike organ on body surface, used to catch and hold paniculate matter.

Peristome—see mouth.

Plate, ossicle=any single skeletal element except

the spines and pedicellariae.

Podia—see tube feet.

Preoral spine, apical spine=spine or spines at apex

of mouth plate.

Primary plates=the first-formed, usually large,

plates of the disc, generally obscured or lost in

development, but still plainly distinguishable in

certain starfish, such as the Zoroasteridae.

Proximal=toward the disc center.

R. major radius=the distance from the center of

the disc to the tip of an arm.

r, minor radius=distance from center of disc to

interradial margin.

Ray-see arm.

Recurved hyaline epioral spine=large, heavy, semitransparent spine on the surface of the mouth

plate, directed away from the mouth. Found in

Odontasteridae.

Reticulate=skeleton of plates arranged in a loose

network surrounding irregular plateless spaces.

Sacculate=enveloped in tissue, as sacculate spines.

Secondary plates=smaller plates between the primary or larger plates.

Segmental pore=a tiny aperture between the adambulacrals in Pterasteridae.

Skeletal arches—see costae.

Skeleton=the hard, calcite elements of the starfish

body.

Spine=a spikelike, thornlike, or peglike external

ornamentation.

Spinelet, spinule=a small spine.

Spiniform pedicellaria—see fasciculate pedicellaria.

Spiraculae=small round or irregular holes in the

supradorsal membrane.

Subambulacral=plate next to adambulacral on actinal surface.

Suboral spine—see epioral spine.

Sugar-tongs pedicellaria, excavate pedicellaria, alveolar pedicellaria=small pedicellaria with two

slender, toothed valves, usually lying in an alveolus. Found mostly in the Ophidiasteridae.

Superambulacral plate=an internal plate above the

juncture of the ambulacral and adambulacral.
NUMBER 126 21

Superomarginal plates, superomarginals=marginal

pertaining to the abactinal surface.

Supradorsal membrane=the external dorsal surface above the true abactinal surface. Found

only in the Pterasteridae.

Tessellate=skeleton of flat plates fitting closely together, like a mosaic.

Tube feet, podia=soft locomotor organs, in two

or four rows, within the ambulacral groove.

Tubercle=a large conical or low, rounded projection on a plate, not deciduous.

Valvate pedicel laria—see bivalved pedicellaria.

Ventral—see actinal.

Key to the Orders of Asteroidea

1. Marginal plates usually large, conspicuous, defining ambitus; pedicellariae spiniform, pectinate,

valvate. or excavate, never of basal piece and two valves; papaulae restricted to abactinal

surface (except in some Ophidiasteridae); abactinal surface paxillate, tessellate, or imbricate, never reticulate or absent 3.

Marginal plates usually small, inconspicuous, not defining ambitus; pedicellariae straight

and/or crossed, or absent; papulae frequently on all surfaces; abactinal surface reticulate,

imbricate, or absent, never tessellate or truly paxillate 4

2. Only one series of marginal plates; actinal plates in single transverse series, separated by

continuous spinous fasdoles from furrow to margin; arms five to many, flat, straplike;

abactinal surface always paxillate Platyasterida

Two series of marginal plates; actinal plates not in single transverse series; arms always five;

abactinal surface usually tessellate or imbricate (rarely paxillate) S

3. Mouth plates keeled, prominent, with median furrow; marginals either alternate or with

intermarginal channels; pedicellariae usually spiniform or pectinate; abactinal surface

usually paxillate Paxillosida

Mouth plates inconspicuous; marginals corresponding and without intermarginal channels;

pedicellariae valvate, sunken in ossicles; abactinal surface usually tessellate or imbricate ....

Valvatida

4. Mouth plates prominent, broad, unkeeled; no pedicellariae Spinulosida

Mouth plates not particularly broad; straight and/or crossed pedicellariae present and usually

abundant 5

5. Dorsal skeleton reticulate, plates not in regular series; ambulacral and adambulacral plates

crowded and compressed; papulae on all surfaces Fordpulatida

Dorsal skeleton imbricate or in costae, never reticulate; papulae absent or confined to

dorsal surface 6

6. Disc domed, with definite arrangement of enlarged primary plates; arm plates in compact

and imbricating longitudinal and transverse series; adambulacral plates alternating carinate

and noncarinate; arms always five, not sharply distinct from disc Zorocallida

Disc plane, no enlarged primary plates; arm plates, not extending beyond proximal third of

arm, usually in costae; adambulacral plates not carinate; arms always more than five,

sharply distinct from disc Eudasterida

Order PLATYASTERIDA Spencer, 1951

Plates in transverse series, at least on actinal surface; fascioles continuous from furrow to margin;

only one series of marginals (none in certain fossils). Luidiidae is only living family.

Family LUIDIIDAE Verrill, 1899

Arms five to many, flat, strap shaped. Superambulacral plates present. Abactinal surface paxillate.

Genus Luidia Forbes

Luidia Forbes, 1839:123. [Type, by original dsignation, L.

fragilissima Forbes=L. ciliaris (Philippi) .]

Hemicnemis Muller and Troschel, 1840:105. [Type, by original designation, L. ciliaris (Philippi).]

Petalaster Gray, 1840:183 [Type, by original designation,

P. hardwickii Gray.]

The genus Luidia is distinguished by its long, flat,

straplike arms, small disc, absence of superomarginal plates, paxillar abactinal surface, and elongate

inferomarginal plates extending from the ambitus
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

to the adambulacral plates. Inferomarginals, with

a conspicuous fringe of inferomarginal spines, are

present. This is mainly a shallow-water genus, species in this collection ranging from shore to 130

fathoms. All of the species are sand or sand/mud

dwellers. They are generally more active and agile

than other starfishes, and they feed principally on

small molluscs and on other echinoderms.

Key to the Species of Luidia

1. Arms nine L. senegalensis

Arms five or six 2

2. Paxillae square or rectangular; skeleton compact and sturdy L. clathrata

Paxillae rounded; skeleton loose or fragile 3

3. Arms banded or blotched with black L. alternate

Arms not banded or blotched with black 4

4. Abactinal surface mostly black; marginal spines large, erect; pedicellariae numerous

L. sagamina

Abactinal surface gray or brown; marginal spines fine, not usually erect; pedicellariae few

or none 5

5. Inferomarginal plates with single long marginal spine and numerous small spines

L. barbadensis

Inferomarginal plates with three long tapering spines L. elegam

Luidia senegalensis (Lamarck)

PLATE 1: FIGURES A, B

Asterias senegalensis Lamarck, 1816:255.

Luidia senegalensis.—Mutter and Troschel, 1842:78, pi. 5: fig.

4.-Perrier, 1876a:262; 1878:34, 66-67, 91, 96-97-Rathbun,

1879:149.-Sladen, 1889:246, 742.-H. L. Clark, 1898a:5;

1919:54, 71; 1933:20-Doderlein and Hartmeyer, 1910;151.

-Doderlein, 1920:238, 249-250, figs. 9, 20.-A. H. Clark,

1939:442.-Engel, 1939:3, 7.-Bernasconi, 1943:5-6.-Tommasi, 1958:9-11, pi. 2: fig. 2; 1970:8, fig. 21.-Cherbonnier,

1959:170.-Durand, 1959:21-22, fig. 2.-Engel and Schroevers, 1960:5.-Ummels, 1963:94-95, pis. 10, 11.

Luidia marcgravii Lutken, 1859:43-46.—Verrill, 1867:343:

1915:208-209 -Boone, 1933:76-77, pis. 33-36.

There are nine arms, and the abactinal surface is

compact. The paxillae are square, in regular longitudinal and transverse rows, except in the midregion of the arms and disc, where they are small

and irregular. The center of the paxilla is compactly covered with small granules, and the

periphery bears numerous fine spinules. The inferomarginal plates bear one or two (two in larger

specimens) rather short, acute marginal spines,

thickened near the base, sometimes curved, not at

all flattened, usually more or less appressed to the

arm. Below, the inferomarginals are covered with

short flattened spines and spinules. The adambulacral plates bear two flattened, curved furrow

spines, one above the other, the inner furrow spine

more slender than the outer one, and behind them

are two stout spines, side by side. There are no

actinal interradial plates, the very elongate inferomarginals in this region exetnding to the mouth.

The jaws are narrow, projecting across the peristome, and armed with a tuft of stout spines. No

pedicellariae were noted. The madreporite is concealed by paxillae.

Specimens in this collection are from 10—20 fathoms, from the northeast coast of South America.

The species is known from Florida, the Antilles,

and Senegal.

MATERIAL EXAMINED.—Oregon Stations: 2272 (4)

[R=50 mm, r = 9 mm; Rr=l:5]; 4472 (1) [R=?

(all arms broken), r=20 mm; this specimen ingested a large Cardium, which caused the disc to

appear much inflated].

Luidia clathrata (Say)

PLATE 1: FIGURES C, D

Asterias clathrata Say, 1825:142.

Luidia clathrata.-Lutken, 1859:37-39.-Gray, 1866:4.-Verrill,

1867:343; 1895:134; 1901:36; 1914:7, 31, pi. 103; 1915:200,

pi. 24: fig. 2.-A. Agassiz, 1877:116, 117-119, pi. 2O.-Perrier. 1878:34, 91, 95, 96-Sladen, 1889:253, 742.-H. L. Clark,

1898a:5; 1901, p. 339, 343; 1919:54, 55, 71; 1933:19-20.-

Doderlein and Hartmeyer, 1910:150-151.-Doderlein, 1920:

238, 239, 251-252, figs. 1, 21.-A. H. Clark, 1939:442.-Caso,
NUMBER 126 23

1943:46-50, pi. 12: figs. 1-2, pi. 13: figs. 1-2; 1961:39-41-

Bernasconi, 1943:6-7, pi. 2: fig. l.-Tommasi, 1958:9, pi. 2:

fig. 1; 1970:8, fig. 22.-Cherbonnier, 1959:170.-Durand,

1959:21-23, pi. 3.-Engel and Schroevers, 1960:5.-Ummels,

1963:95-97, pis. 10-11.—Gray, Downey, and Cerame-Vivas,

1968:139, fig. 8.

There are five arms, and the skeleton is compact.

The paxillae are square, in regular longitudinal

and transverse rows except on the midregion of

the arms and disc. The center of the paxillae is

covered with granules, the periphery with fine

spinules. The broad midregions of the arms and

disc have small irregularly shaped and arranged

paxillae. The inferomarginal plates bear one moderately long, curved, flattened marginal spine and

one or two small spines above it, not appressed to

the arm. Below, there is a covering of very short

spines and spinules. The adambulacral plates have

two furrow spines, one behind the other, laterally

flattened and somewhat curved, the inner spine

more slender than the outer one. Behind these are

two spines, side by side, flattened in the plane of

the furrow. A few actinal interradial plates are

present, one series extending a short distance on

the arm. The oral spines form a thick cluster, and

the jaws are covered with fine spinules. No pedicellariae were noted. The madreporite is concealed

by paxillae.

Specimens in this collection are from 0—45 fathoms, from Cape Hatteras to northeastern South

America and from the Gulf of Mexico. The known

range is from New Jersey to Brazil. The color is

usually light gray, with a darker gray stripe down

the middle of the arm; however, specimens from

the southern part of the range may be pastel. Also,

the arms have a broader base; there are usually two

or even three marginal spines, and an extra pair of

side-by-side adambulacral spines may occur, especially proximally. These are not specific differences

but geographic variations, of less than subspecific

importance.

MATERIAL EXAMINED.—Oregon Stations: 1865 (1)

[R=100 mm, r=12 mm, Rr=l:10]; 6300 (5) [R

=77 mm, r=12.5 mm, Rr=l:7j; 3555 (1) [R=20

mm, r=6 mm, Rr=l:4]; 2272 (1) [R=54 mm, r

=9 mm, Rr=l:5]; 2207 (1) [R=42 mm, r=9

mm, Rr=l:4]; 5894 (1) [R=67 mm, r=9 mm,

Rr=l:6]; Silver Bay Stations: 1515 (1) [R=24

mm, r=6 mm, Rr=l:4]; 35 (1) [R=90 mm, r =

11 mm, Rr=l:9]; 1952 (1) [R=120 mm, r=6 mm,

Rr=l:20].

Luidia alternate (Say)

PLATE 2: FIGURES A, B

Asterias alternata Say, 1825:144-145.

Luidia alternata.-Lutken, 1859:42-43; 1871:301.-Verrill.

1867:343.-Perrier, 1876a:254; 1878:34, 91, 96.-Sladen. 1889:

250-251, 740.-Ives, 1890:326.-H. L. Clark, 1898a:5; 1919:54,

55, 71; 1933:20, pi. l.-Doderlein and Hartmeyer, 1910:151-

Doderlein, 1920:241, 267-268, figs. 7, ll.-Caso, 1943:56-57,

pi. 16: figs. 1-2, pi. 17: figs. 1-2; 1961:44-47, figs. 11-12.-

Engel and Schroevers, i960:— Ummels, 1963:97-98, pis. 8,

10.—Gray, Downey and Cerame-Vivas, 1968: fig. 6.—Tommasi 1970: pi. 8: fig. 24.

Luidia granulosa Perrier, 1869:109.

Luidia variegata Perrier, 1876b:257.

This species has five arms, and the skeleton is not

compact. The paxillae are small, high, and well

separated, in regular rows along the side of the

arm, irregular and compact in the middle of the

arm and disc. The sides of the arms appear higher

in this species because the paxillae of the second,

third, or fourth row above the inferomarginals

bear a stout acute spine. Similar spines may appear on a few other paxillae as well. Paxillae without a central spine have 1-8 central clavate spinules, and all paxillae have numerous subclavate

peripheral spinules. The inferomarginal plates

bear 2 or 3 long, acute, erect marginal spines, not

at all flattened, and below, 2-5 moderately long,

somewhat flattened spines; the channels between

the inferomarginals are deep and bordered on each

side by numerous fine spinules. The adambulacral

plates bear a long, fine, curved, laterally flattened

furrow spine and 2 or 3 other spines, long, acute,

and not flattened. There may be small supplementary spines as well. The very few actinal interradial

plates usually bear pedicellariae. The mouth plates

are prominent, each plate bearing a double row

of short, stout, acute spines. Actinal pedicellariae,

stout and three- or four-valved, are usually present.

The madreporite is hemispherical, with fine shallow channels.

This is the only nonbrooding starfish showing

what I believe to be sexual dimorphism. The female (the specimen from Oregon Station 5894) is
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

much larger, looser of skeleton, and bears many

more pedicellariae.

The color in life is distinctive: blotched or

banded in creamy white and black or brown. Specimens in this collection came from 0-20 fathoms,

from Florida and the Caribbean. The range of the

species is from Cape Hatteras to northern South

America; it is not known from the western Gulf of

Mexico.

MATERIAL EXAMINED.—Oregon Stations: 5894 (1)

[R=115 mm, r=7 mm, Rr=l:17]; 1937 (1) [R=44

mm, r=6 mm, Rr=l:6]; 5456 (1) [R=56 mm, r

=10 mm, Rr= 1:5.6]; 3555 (1) [R=7 mm, r=3

mm, Rr=l:3].

Luidia sagamina Doderlein

PLATE 2: FIGURES C, D

Luidia sagamina Doderlein, 1920:290, fig. 37.

Luidia aciculata Mortensen, 1933a:425, fig. 7, pi. 20; figs. 7-12.

Luidia sagamina aciculata.—Madsen, 1950:199, figs. 6-7.

There are five long arms, and the skeleton is moderately compact, the body flat. The paxillae are

in irregular longitudinal and transverse rows, with

a short, stout central spinule and 6 or 7 glassy,

very thorny peripheral spinules. The inferomarginal plates have 2 or 3 long, rough-textured marginal spines, usually alternating 2-3-2-3. The

upper spine is longest and is usually black at the

base; paxillae also are mostly black. The channels

between the inferomarginals are wide, bordered by

long, fine spinules. The adambulacral plates have

one slender, curved furrow spine, not flattened, and

one other very much larger, long, acute, evenly

tapered spine. There is only one actinal interradial

plate, which bears a large three-valved pedicellaria;

the valves of the pedicellariae are somewhat flattened at the tip. The jaws are armed with 4-6 long,

rough spines and within the peristome on each side

are two small, subequal blunt spines arising from

a common furcate base. The madreporite is concealed by paxillae.

The distribution of this species is of considerable

interest. Previously, it was known only from

Sagami Bay, Japan (the type locality); Durban,

South Africa; St. Helena; from Gaboon to Senegal

in tropical West Africa; and the Cape Verde Islands. Madsen (1950) states that he could find no

difference between the Japanese L. sagamina and

Mortensen's (1933) L. aciculata from St. Helena,

but kept them separate for geographical reasons

only. With this further extension of range, I do

not believe such separation is justifiable. Although

not common, L. sagamina may have a nearly worldwide distribution. A possible clue as to how this

species became so widely distributed may be found

in a report by Tokioka (1942) of a gigantic Luidia

bipinnaria (12 mm) taken in the plankton near

Seto, Japan. Mortensen (1939) stated that luidiids

of the ciliaris group (to which L. sagamina belongs), with very large bipinnariae, have a freeswimming larval planktonic life of some months.

These bipinnariae are, moreover, active swimmers,

and it is possible that under particularly favorable

conditions of currents, wind, etc., large actively

swimming larvae could become very widely distributed, even though the adults are rather rare.

Another specimen, examined after the above description was written, has been collected by the

Duke University R/V Eastward off North Carolina.

MATERIAL EXAMINED.—Silver Bay Station 1515 (1)

[R=8 mm, r=3 mm, Rr= 1:2.5].

Luidia barbadensis Perrier

PLATE 3: FIGURES A, B

Luidia barbadensis Perrier, 1876a:29; 1884:267, pi. 10: figs.

7, 8.-Verrill, 1915:205, pi. 24: fig. l.-John and Clark,

1954:145.

Luidia convexiuscula Perrier, 1881a:30; 1884:268, pi. 10: fig.

6.-Verrill, 1915:207.-John and Clark, 1954:145.

This species has six long, slender arms. The skeleton is fragile. Paxillae are in regular longitudinal

and transverse rows except in the midregion of

the arms and disc; they have a uniform covering

and peripheral border of 20-30 short, thorny, subclavate spinules. The inferomarginal plates have

two long, tapering, acute spines, the upper one

smaller than the lower, both somewhat curved. Below are 3-6 small spines; the channels between the

inferomarginals are wide and shallow, with the

bordering spinules long and few. The adambulacral plates bear a very long, slender furrow spine,

behind which is a row of three obliquely placed

unequal spines, and behind them another row of

two spines, frequently replaced by a large threevalved pedicellaria. The actinal interradial area

has 3-6 large, stout three- or four-valved pedicel-
NUMBER 126 25

lariae, around the bases of which are fringes of

spinules. The jaws are concealed by a bundle of

numerous long stout spines. The madreporite,

barely visible beneath the paxillae, is flat and covered with deep gyri.

Specimens in this collection were collected in

65-80 fathoms, in the Gulf of Mexico west of the

Florida Keys. The known range is from Florida

to Venezuela.

MATERIAL EXAMINED.—Alaminos Stations: 20/65—

A-9 (1) [R=75 mm, r=5 mm, Rr=l:15]; 21/65-

A—9, arm fragments.

Luidia elegans Perrier

PLATE 3: FIGURES C, D

Luidia elegans Perrier, 1876a:256; 1884:269.-Verrill, 1880:403;

1895:134; 1915:203, pi. 16: figs. 4, 4a, pi. 29: fig. l.-Gray,

Downey, and Cerame-Vivas, 1968:140, fig. 9.

This species has five arms, and the skeleton is loose

and fragile. The paxillae are small, high, in

vaguely transverse rows on a narrow portion of each

side of the arms, but otherwise quite irregularly

distributed. They have one central and 7—15 peripheral slender spinules of moderate length, not

acute; the central spinule on certain disc paxillae

is sometimes replaced by a small, blunt, two-valved

pedicellaria. The inferomarginal plates have two

or three (alternating 2-3-2-3) long, stout, straight,

evenly tapered, acute marginal spines, and below

a few tiny spinuks and sometimes one or two small

spines. The adambulacral plates have a stout,

curved furrow spine, and behind it two long, stout,

tapering spines, one above the other. Between the

adambulacral and inferomarginal plates there is

usually a stout, blunt-tipped, two- or three-valved

pedicellaria. The actinal interradial area bears a

number of these short, blunt, two-valved pedicellariae. The jaws have two very large, stout, oral

spines, behind which are two rows of 3—7 spines

diminishing in size distally, the outer row much

smaller than the inner one. Within the peristome,

each mouth plate bears a large pedicellaria. The

madreporite is hemispherical, covered with bumps

rather than gyri.

This species was taken in 100-130 fathoms, in

the western Gulf of Mexico, a westward extension

of the range. It does not occur in very shallow

water. The previously known range was from Martha's Vineyard to the eastern Gulf of Mexico and

the West Indies.

MATERIAL EXAMINED.—Oregon Stations: 1514 (2)

[R=77 mm, r=10 mm, Rr=l:7]; 1519 (2) [R=83

mm, Rr=l:10]; 1513, arm fragments.

Other Tropical-Subtropical Species of Luidia

Not Represented in This Collection

Only three other Luidias, not taken in these collections, are reported from the area here included. L.

bernasconiae A. H. Clark (1945) is a perfectly valid

species, despite A. M. Clark's placing it in the

synonymy of L. alternata in her otherwise excellent paper (1951a). It differs from L. alternata in

having 5-10 clavate peripheral spinules on the

paxillae vs 10—16 slender, acute ones on L. alternata, and there are two adambulacral spines in L.

bernasconiae, vs 3—4 in L. alternata. Pedicellariae

are abundant on L. bernasconiae, and are shorter

and broader than in L. alternata.

The other two species, interestingly enough, were

collected in the same dredge-haul by the Rosaura

expedition. L. rosaurae John and Clark (1954) is

closely allied to L. barbadensis, but where L. barbadensis has six arms, L. rosaurae has only five. L.

rosaurae has both dorsal and ventral pedicellariae,

but L. barbadensis has only ventral ones. L. barbadensis has two marginal spines, L. rosaurae only

one. The third species, L. barimae John and Clark

(1954), is ten-rayed; the skeleton is loose and

fragile, and none of the paxillae are arranged in

regular rows.

A fourth species, L. atlantidae Madsen (1950),

from western and northwestern Africa, has not been

reported in the literature from the western Atlantic, but I have seen specimens collected off the coast

of North Carolina south of Cape Fear, and specimens have also been taken in the Florida area

(Halpern, personal communication).

Order PAXILLOSIDA Perrier, 1884

Mouth plates prominent, with keel and median

furrow. Actinal interradial area with small ossicles. Plates not in transverse series. Ambulacral

areas not compressed. Tube feet in two rows.
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to the Families of Paxillosida

1. Aboral surface paxillate; oral interadial area with flat ossicles extending onto arms; cribiform

organs or specialized marginal fasciolcs present; papulae scattered over most of abactinal

surface; no dorsal muscle bands on arms; tube feet in two rows, pointed 2

Abactinal surface with simple flattened plates carrying few spinelets; oral interradial plates

not extending onto arms; marginals alternate, without fascioles or cribriform organs;

papulae confined to base of arms and disc, frequently in localized papularia; pair of dorsal

muscle bands on arms; tube feet in two rows, suckered Benthopectinidae

2. Cribriform organs present, continuous with actinal channels bordered by webbed spinelets;

spinelets of abactinal paxillae webbed; marginals covered with smooth membrane; disc

broad \ Goniopectinidae

Marginal fascioles present; paxillar spinelets not webbed; marginals not covered with smooth

membrane; disc small Astropectinidae

Family ASTROPECTINIDAE Gray, 1840

Disc small; arm long, pointed, straight-sided (rarely

petaloid). Marginal fascioles present. Aboral surface paxillate. Oral interradial areas with flat ossicles extending onto arms. Superambulacral plates

present. Tube feet pointed, ampullae double.

Key to the Genera of Astropectinidae

1. Madreporite large, and superomarginals confined to aboral surface, with inferomarginals

defining the ambitus Dipsacaster

Either one of above conditions, but not both 2

2. Inferomarginals not separated from adambulacral plates by actinal intermediate plates

Astropecten

Inferomarginals separated from adambulacrals for part or whole of ray by actinal intermediate

plates 5

3. Superomarginals thin and confined to side of ray Dytaster

Superomarginals not thin and confined to side of ray 4

4. Actinal interradial areas small 6

Actinal interradial areas large 5

5. Madreporite concealed by paxillae Plutonaster

Madreporite naked and large Tethyaster

6. Actinal spinelets sacculate Psilaster

Actinal spinelets not sacculate 7

7. Odd interradial actinal intermediate plates present Blakiaster

No odd interradial actinal intermediate plates Persephonaster

Astropecten Gray

Astropecten Gray [from Linck, 1733], 1840:180. [Type, by subsequent designation, Asterias aranciacus Linnaeusz=A aurantiacus Tiedemann (Fisher, 1908:93).]

Stellaria Nardo, 1834:716. [Type, by monotypy, S. aurantiaca.

Invalidated by Stellaria M0ller.]

Asterias L. Agassiz [non Linnaeus], 1835b: 168. [Type, by original designation, A. aurantiaca Linnaeus.]

Astropectinides Verrill, 1914:321. [Type, by original designation. A. mesacutus (Sladen).]

Conspicuous marginal plates, ambital fringe of

spines, paxillate dorsal surface, few or no actinal

interradial plates; inferomarginal spines born on

horseshoe-shaped tubercles; inferomarginal and

superomarginal plates opposite, usually massive,

never overlapping; superomarginals encroach more

or less upon aboral surface, inferomarginals frequently project beyond them; arms normally five,

body flat; distinct fascioles between marginal plates;

equal number of adambulacral, inferomarginal,

and superomarginal plates; adambulacral plates

bear 3 or 4 furrow spines in a longitudinal row,

additional spines on actinal face of plate; pedicellariae rare.

There are over 150 known species of Astropecten, mostly from tropical and subtropical waters.

They occur in shallow to moderate depths, princi-
NUMBER 126 27

pally on sandy or shelly bottoms. Their food consists mainly of small molluscs. Doderlein monographed the genus in 1917, but unfortunately, this

paper is now somewhat out of date. The genus has

an extensive fossil record, reaching back possibly

to the Upper Cretaceous.

Key to the Species of Astropecten

1. Marginals spinous, superomarginals concave A. americanus

Marginals granulose, superomarginals convex 2

2. Two basal superomarginals (and frequently other proximal superomarginals) with single

large stout spine A. duplicatus

No spine or tubercle on proximal superomarginals 3

3. Single spine on distal superomarginals A. articulatus

No spines or tubercles on any superomarginals 4

4. Body extremely flat, arms broad-based, inferomarginals projecting far beyond superomarginals A. marginatus

Body not particularly flat, arms not broad-based, inferomarginals scarcely or not at all

projecting beyond superomarginals 5

5. Superomarginals very numerous (over 40) A. cotnptus

Superomarginals few (25 or less) 6

6. Adambulacral spines small A. cingulatus

Adambulacral spines long, slender A. nitidus

Astropecten americanus (Verrill)

PLATE 4: FIGURES A, B

Archaster americanus Verrill, 1880:402; 1885:542.

Astropecten americanus.-Verrill, 1894:255; 1895:133; 1915:184,

pi. 16: figs. 1, la.-Doderlein, 1917:106, pi. 2: fig. 11, pi. 9:

figs. 9, 9a, 96.-Gray, Downey, and Cerame-Vivas, 1968:143,

figs. 13. 14.

The five arms are long, narrow, and taper evenly

to an acute terminus. Paxillar areas on the arms

are very narrow, less than the width of the supermarginals. Paxillae are small, high, set in the dorsal membrane, and not usually in contact with one

another. The 6—10 moderately long peripheral

spinules of the paxillae are not acute; in preserved

specimens they are usually erect, and paxilla and

spinules together look like a paint brush. The

papular pores between the paxillae are moderately

large. At the side of the arms, the paxillae are in

transverse rows, and they diminish in size from the

disc to the end of the arm. The superomarginal

plates are tumid and wedge shaped in the interradius, but on the arms they are square and flat

or slightly concave. They have a sparse covering

of tiny spinules. The inferomarginals project

strongly beyond the superomarginals; they bear two

long, slender, acute marginal spines, somewhat

curved and usually appressed to the arm and directed outward and upward. The lower spine is

generally smaller than the upper one. In the interradial area, these spines are flattened and bladelike. Below, the inferomarginals are covered with

tiny acute spines and spinules. The three adambulacral furrow spines are flattened laterally, the central one is the largest, and behind them is a small

group of spines, sometimes replaced by a large

three-valved pedicellaria, especially proximally.

The mouth plates are long, narrow, and prominent; each bears a row of small spinules, and on

either side of the jaw are two rows of moderately

long fine spinules.

In life, this species is a lively reddish brown. It

is perhaps the most abundant western Atlantic species; in places, it literally paves the ocean floor.

Specimens in this collection came from 150-230

fathoms. This species does not occur in shallow

water; it was taken in the Gulf of Mexico, off Cape

Hatteras, and off Guyana.

MATERIAL EXAMINED.—Alaminos Stations: 10/68-

A-13 (1) [R=72 mm, r=9 mm, Rr=l:7]; 9A/

68-A-7 (1) [R=56 mm, r=9 mm, Rr=l:5]; 7/68-

A-13 (2). Oregon Stations: 382 (1) [R=82 mm,

r:=12 mm, Rr=l:8]; station unknown (1) [R=81

mm, r=9 mm, Rr=l:8]; 4302 (13) [R=20 mm,

r=4 mm, Rr=l:5]; 1509 (3). Silver Bay Station

2863 (1) [R=24 mm, r=5 mm, Rr=l:5].
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Astropecten articulatus (Say)

PLATE 4: FIGURE C, D

Asterias articulatus Say, 1825:141.

Astropecten articulatus.—Muller and Troschel, 1842:72.— Lutken, 1864:128, 129.-Verrill, 1868:343; 1885:40, 77; 1895:

133; 1915:159.-Perrier, 1876b:29O.-A. Agassiz, 1877:114, pi.

19: figs. l-8.-lves, 1891:337, pi. 16: figs. 4-8.-H. L. Clark,

1898a:4.-Fontaine, 1953:181, fig.-Cherbonnier, 1959:320-

322, pi. 7.—Engel and Schroevers, 1960:2.—Hulings and

Hemlay, 1963:354-359.-Dragovich and Kelly, 1964:91.-

Gray, Downey and Cerame-Vivas, 1968:143, fig. 15.—

Tommasi, 1970:6, fig. 17.

Astropecten dubius Gray, 1840:182; 1866:4.

Astropecten articulatus duplicatus.—Ron, 1967:277, fig. 6.

The five arms of this species are moderately long,

high on the side, and not at all acute. The paxillar areas on the arms are about twice the width of

the marginals and very compact. The paxillae are

in regular transverse rows except in the midregion

of the arm and on the disc; they have a uniform

covering of granuliform spinules and are very close

together, giving an even granulate appearance to

the entire dorsum. Superomarginals are higher

than wide, with a uniform covering of granules.

The interradial superomarginals never bear spines,

but some on the arms may bear a small tuberculate

spine. The inferomarginal plates do not project

beyond the superomarginals. They bear a pair of

flattened spines, side by side, and there is frequently

a third tiny spine proximal to them. Below, they

are covered with small spines and spinules. There

are a few tiny actinal interradial plates (never more

than six), bearing a small, central spine and surrounded by tiny fine spinules. The adambulacral

plates have 3 or 4 slender furrow spines, side by

side; behind them are two larger, flattened, truncate spines, also side by side, and behind them are

three smaller similar spines. The mouth plates are

prominent, elongate, and covered with small short

spines. On either side there is a double row of

spinules, and at the apex a cluster of short, thick,

blunt spines. The madreporite is conspicuous,

round, flush with the dorsal surface, and covered

with fine gyri.

The known range of this species is from Chesapeake Bay to Colombia, including the Gulf of Mexico. Specimens in this collection came from 0-100

fathoms, mostly from around Florida. In life, this

species is usually bright blue or purple, with conspicuous yellow or orange marginals.

MATERIALS EXAMINED.—Oregon Stations: 5894 (1)

[R=84mm, r = 14 mm, Rr= 1:8]; 2164 (2) [R=70

mm, r=14 mm, Rr=l:6]. Silver Bay Station 35 (1)

[R=73 mm, r=14 mm, Rr=l:7].

Astropecten cingulatus Sladen

PLATE 5: FIGURES A, B

Astropecten cingulatus Sladen, 1889:218, pi. 35: figs. 5, 6, pi.

39: figs. l-3.-Verrill, 1915:178.-Bernasconi, 1941:42, pi. 7:

figs. 1-2; 1958a:130-131, pi. 3: figs. 1-2; 1961:22, pi. 1: fig.

2; 1966:161-162.—Engel and Schoevers, 1960:3.—Tommasi,

1970:5, fig. 16.

This compact five-armed species has a relatively

small paxillar area. The paxillar area on the arms

is less than half the width of the massive superomarginals, and somewhat sunken below them. The

paxillae are high and, on the arms, flattened longitudinally. They are compact on the disc, less so on

the arms. They have 1-3 central granules and 7—11

peripheral clavate spinules. The superomarginals

are massive and tumid and encroach quite far on

the dorsal surface; they are covered with granules,

those on the outer half of the plate usually larger

than those on the inner half (but sometimes the

other way around). The channels between the plates

are bordered with many fine setiform spinules. The

inferomarginals project very slightly beyond the

superomarginals and bear three flattened marginal

spines of moderate length. Below, the plates are

bare, save for a few acute spinules. The adambulacral plates bear 3 or 4 furrow spines parallel to

the furrow, the central one large, sturdy, and curving slightly inward toward the furrow.

The actinal face of the plate bears one long acute

tapering spine, and, usually, 1-3 tiny spinelets.

The mouth plates are small and narrow, covered

with spinules, and with a cluster of small, blunt

oral spines. The madreporite is very small, round,

covered with deep gyri, and almost touching the

interradial marginals.

The depth range of this species in this collection

is 30-50 fathoms, and all are from the Gulf of

Mexico. The range is Cape Hatteras to Brazil.

MATERIAL EXAMINED.—Oregon Stations: 1495, (2)

[R=31 mm, r=10 mm, Rr=l.S]; 2, (1) [R= 8 mm,

r=5 mm, Rr=l:1.5]; 102 (3) [R=55 mm, r=10

mm, Rr=l:5]; 123 (3) [R=50 mm, r=10 mm,

Rr=1.5]. Silver Bay Station 2432, (1) [R=23 mm,

r=8 mm, Rr=l:S].
NUMBER 126 29

Astropecten comptus Verrill

PLATE 5: FIGURES C, D

Astropecten comptus Verrill, 1915:176, pi. 12: figs. 3-3c, pi. 22:

fig. 1.

The five arms in this species are moderately long

and slightly constricted at the base. The very even

granular appearance of the dorsum and superomarginal plates is strongly reminiscent of A. articulatus, but the arms are longer and narrower and taper

to a more acute point. The paxillae have 4—10

central granules and 9—15 peripheral clavate

spinules. The first three or four series of paxillae

on the arms are in regular transverse rows. The

superomarginal plates, evenly covered with granules, are higher than wide, particularly in the

interradius, forming a high side to the arms. The

inferomarginal plates project slightly beyond the

superomarginals and bear two flattened curved

spines, side by side. Below, they bear numerous

small spines and spinules. The adambulacral plates

bear three long slender furrow spines, flattened in

the plane of the groove, and behind them are two

shorter but broader truncate spines, side by side;

behind these are about three small spines. The

mouth plates are short and narrow, covered with

moderately long spines and with a cluster of short,

flat oral spines at the apex. The madreporite is

small, round, and covered with coarse gyri.

Specimens in this collection are from 10—30

fathoms, from around Florida. The known range is

Cape Hatteras to Cuba, and the Gulf of Mexico.

MATERIAL EXAMINED.—Oregon Stations: 1648 (1)

[R=61 mm, r=12 mm, Rr=l:6; 4806 (1) [R=53

mm, r=8 mm, Rr=l:7. Silver Bay Station 2775 (4)

[R=16 mm, r=5 mm, Rr=l:3].

Astropecten duplicatus Gray

PLATE 6: FIGURES A, B

Astropecten duplicatus Gray, 1840:185; 1866:3.—Perrier,

1876b:271.-Verrill, 1915:165, pi. 22: fig. 2, pi. 23: figs. 1, 2.

- H . L. Clark, 1898a:5.-Gruvel, 1909:1018.-Boone, 1928:

5.-Fontaine, 1953: pi. 181; fig.-Cherbonnier, 1956:322-323;

1959:108-109, fig. l.-Caso, 1961:27-29, fig. 4.-Gray, Downey and Cerame-Vivas, 1968:144, fig. 16.

Astropecten valenciennii Muller and Troschel, 1842:68.

Astropecten variabilis Lutken, 1859:51.—Verrill, 1867:343.—

A. Agassiz, 1869,-Field, 1893:82.-Loriol, 1899:16, pi. 2.

Astropecten articulatus duplicatus.—Ron, 1967:277, fig. 6.

The five arms of this species are of moderate length

and taper to a rather acute point. The paxillar area

on the arms is about twice the width of the superomarginal plates. Paxillae are in regular transverse

rows on the arms, and bear 15-25 capitate spinules.

The superomarginal plates are higher than wide,

extending above the paxillar surface, and the

midportion of each plate is covered with spherical

granules, while the sides bear numerous uniform

fine blunt spinules. In the interradius, the first

pair of superomarginals (and usually a few on each

side of them) bear a stout tuberculate spine on the

inner edge of the upper surface; superomarginals

on the arms may bear a similar smaller spine on

the outer edge of the upper surface. The inferomarginal plates extend beyond the superomarginals;

each bears a single large, flattened spine and beside

it a much smaller spine. Below, the inferomarginal

plates bear a dense covering of flattened spines and

spinules. The central furrow spine of the adambulacral plates is laterally flattened, with a rounded

tip, and flanked by two shorter truncate spines.

Behind the furrow spines are two truncate spines,

the distal one large and flattened in the plane of

the groove. Behind these are 2 or 3 small spines.

The mouth plates are thickly covered with small

blunt spines. The madreporite is small, round,

evenly covered with deep gyri.

This spinous Astropecten ranges from shallow

water to over 300 fathoms; it occurs all over the

Gulf of Mexico and the Caribbean, and north to

Cape Hatteras.

MATERIAL EXAMINED.—Oregon Stations: 3602 (1)

[R=19 mm, r=7 mm, Rr=l:3]; 4089 (2) [R=15

mm, r=4 mm, Rr=l:4]; 2243 (1) [R=20 mm, r =

5 mm, Rr=l:4]; 3555 (2) [R=29 mm, r=7 mm,

Rr=l:4]; 5920 (12) [R=34 mm, r=9 mm, Rr=

1:4]; 4472 (2) [R=33 mm, r=6 mm, Rr=l:5]; 150

(27) [R=15 mm, r=5 mm, Rr=l:3]. Oregon II

Station 10499 (3) [R=2 mm, r=5 mm, Rr=l:4].

Silver Bay Station 1953 (2) [R=45 mm, r=ll mm,

Rr=l:4].

Astropecten marginatus Gray

PLATE 6: FIGURES C, D

Astropecten marginatus Gray, 1840-Kenk, 1944:185, figs. 2,

5.-Cherbonnier, 1956:320.-Bernasconi, 1957:67-68, pi. 2:

figs. 3, 4.-Tommasi, 1959:14, pi. 2: fig. 5; 1970:5, fig. 15.-
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Brito, 1960:8, pi. 2: fig. l.-A. M. Clark, 1962; fig. 30a.-

Roa. 1967:267.

An extremely flat and thin-bodied species, this

five-armed starfish has a larger disc area than most

astropectinids. The short arms taper rapidly to an

acute point. The small paxillae are in regular

transverse rows except for a narrow section on the

midarm and disc. The paxillae have 3—6 central

granules and 8-16 peripheral subclavate spinules.

The superomarginal plates are higher than wide

and confined to the dorsal surface; they are covered

with an even coating of small spaced granules, and

the channels between are bordered by fine uniform

spinules. The inferomarginals project strongly

beyond the superomarginals and the upper edge

of the plate thus exposed is covered with granules

continuous with those of the superomarginals. The

inferomarginal plates bear two subequal, slightly

curved marginal spines, not flattened, and proximally, one tiny blunt spine. Below, the long narrow

inferomarginals bear a row of 4 or 5 moderate

long acute spines on their outer edge (under the

marginal spines) and a few scattered acute spines

elsewhere. A single row of small spinules borders

the channels between the inferomarginals. The

adambulacral plates bear three blunt slender furrow spines, the middle one slightly longer, and one

acute spine of about the same length on the actinal

face. On either side of the plate is a row of small

spinules continuous with and similar to those

bordering the channels between the inferomarginals.

The narrow mouth plates are covered with short

stout spines and there are two rows of longer

truncate oral spines. The madreporite is moderately

large, flat, round, and covered with radiating gyri.

All these specimens occurring in this collection,

from 20-30 fathoms, are from northeastern South

America; the normal range also includes Puerto

Rico, so presumably the species will be found also

in the Lesser Antilles.

MATERIAL EXAMINED.—Oregon Stations: 2209 (1)

[R=55 mm, r=15 mm, Rr=l:5]; 2226 (1). Oregon

II Stations: 10502 (2) [R=45 mm, r=12 mm,

Rr=l:4]; 10507 (1) [R=41 mm, r=10 mm, Rr=

1:4].

Astropecten nitidus Verrill

PLATE 7: FIGURES A, B

Astropecten nitidus Verrill, 1915:179, pi. 10: fig. 2.—Gray,

Downey, and Cerame-Vivas, 1968:145, fig. 17.

Astropecten nitidus forcipatus Verrill, 1915:180, pi. 16: fig. 3,

pi. 20: fig. 1.

The five arms of this species are long, narrow, and

tapered very little. The paxillar areas are narrow,

about the width of the superomarginals, and the

paxillae are in regular transverse rows on the arms.

The paxillae have 1—9 blunt central spinules, and

6-15 peripheral ones, also blunt. The superomarginal plates are higher than broad, not forming

high sides to the arms but encroaching quite far on

the dorsal surface. They are covered with numerous,

small, flattened granules and, around the edge,

with many fine spinules. The inferomarginal plates

extend beyond the superomarginals and each bears

two long, acute, slightly curved marginal spines,

appressed to the arm and directed upward and

distad. Below, they bear a few long, slender, acute

spines and some fine tiny spinules. The adambulacral plates have three subequal, flattened furrow

spines and behind them are two groups of smaller

spines or one group of spines and a large conical

pedicellaria. There is a single row of actinal interradial plates, not extending beyond the first inferomarginals, which are usually armed with either a

large conical pedicellaria or a group of small

spines. The short narrow jaws are covered with

small spines and spinules and bear at their apex a

pair of flattened oral spines, with a group of shorter

flattened spines behind them. The madreporite,

located very near the border, is tiny and concealed

by paxillae.

Verrill (1915) distinguished a variety which he

called Astropecten nitidus forcipatus; it is by no

means a subspecies, as it varies from the species

only in number of pedicellariae (a very unstable

character in astropectinids) and its vertical and

horizontal ranges fall well within the limits for the

species. The range of this species is from Cape

Hatteras to the Gulf of Mexico, in 20-200 fathoms.

MATERIAL EXAMINED.—Oregon Stations: 1514 (1)

[R=15mm,r=4mm,Rr=l:4].5913 (2); 4928 (1);

6699 (1) [R=38 mm, r=7 mm, Rr=l:5]; 4398

(12); 1890 (1) [R=27 mm, r=6 mm, Rr=l:4].

Alaminos Station 20/65-A-9 (5) [R=40 mm, r=9
NUMBER 126 31

mm, Rr=l:4]. Silver Bay Station 2453 (1) [R=33

mm, r=7 mm, Rr=l:5]. Combat Station 180 (1).

Other Species of Astropecten Not Taken in

These Collections

Of the four other Astropectens reported from the

area included in this report, the most puzzling

"absentee" from this collection is A. nuttingi

Verrill, 1915. This common species ranges from

North Carolina to Florida, in depths of 90-225

fathoms. It somewhat resembles A. americanus, but

the shape is more regularly stellate, with broader

arms and a wide paxillar area. The superomarginals

bear 1 or 2 small spines on the upper side, and the

paxillae are stellate.

A. alligator Perrier, 1881a, may be the same

species, but Perrier's description is not clear, and

I have not had an opportunity to examine the

type. The same is true of A. richardi Perrier, 1876b,

which may possibly be a synonym of A. articulatus,

and A. antillensis Lutken, 1859, which Verrill believed to be a local variety of A. duplicatus and

H. L. Clark considered a synonym of A. marginatus.

Also, A. ciliatus Grube, 1857, is probably a junior

synonym of A. marginatus', it has been reported

from Puerto Rico, which is in the known range of

A. marginatus.

Psilaster Sladen

Psilaster Sladen, 1885:611. [Type, by original designation,

Astropecten andromeda Muller and Troschel.]

Phidiaster Koehler, 1909a:28. [Type, by monotypy, P. agassizi

Koehler.]

Superomarginals massive, unarmed, encroaching

far onto dorsal surface, covered with granules or

scales; two series of spines or spinules behind furrow series on adambulacral plates; paxillae rather

crowded, at least on disc; inferomarginal spines

usually appressed to arm; actinal interradial area

of moderate size, its plates covered with spinules;

fascioles quite shallow; madreporite small.

The genus is distributed worldwide, in moderately deep waters. These starfishes, like most

astropectinids, feed mainly on small molluscs.

Key to the Species of Psilaster

Actinolaterals extending far out on arm P. cassiope

Actinolaterals not extending beyond the 5th or 6th inferomarginal P. patagiatus

Psilaster cassiope Sladen

PLATE 7: FIGURES C, D

Psilaster cassiope Sladen, 1889:228, pi. 41: figs. 1, 2, pi. 7:

figs. 9, lO.-Madsen, 1950:187.

Psilaster andromeda var. cassiope. Grieg, 1907b: 13.

Psilaster squameus H. L. Clark, 1941:17-18, pi. 2: fig. 1.

This robust species has five arms that taper

smoothly to an acute point. The disc is broad, and

the paxillar area on the arms, relatively broad at

the base, becomes extremely narrow by the end of

the arm. The oval paxillae are in regular transverse rows across the arm, not compact, and small.

They are short and bear a flat even covering of

15—25 granules. The massive superomarginal plates

are about twice as high as broad and encroach quite

far on the dorsal surface. They are covered with

spaced squamiform granules, and the shallow

channels between. the plates are filled with fine

spinules. The inferomarginal plates, with similar

but fewer granules, bear, near their edge, an

oblique row of 2 or 3 small, acute, flattened spines,

appressed to the arm and directed upward. The

interradial actinal areas, of moderate size, are

covered with flattened, squamiform spinules. A

single row of actinolateral plates extends more than

two-thirds the length of the arm. The furrow

margin of the adambulacral plates bears a chevron

of 7 or 8 flattened spines, and behind them, on the

actinal face, are a few more flattened, much shorter

spines. The mouth plates bear, in the center, a few

granuliform spinules and, on either side, two uniform rows of small, short, contiguous spines. There

are two short, flattened, oral spines. The madreporite is small, round, and covered with deep gyri.

Occurring at about 400-500 fathoms, one specimen off the Mississippi Delta, one off Venezuela,

and several in the western Gulf, this species was

previously reported from Cuba by H. L. Clark, as

P. squameus. The differences between his sped-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

mens and Sladen's are too minor to constitute

specific characters; Clark also seems to have misinterpreted Sladen's figures of P. cassiope. However,

Clark did concede that his specimens were possibly

P. cassiope, which, of course, they are. Although

Grieg (1907b) regarded P. cassiope as a variety

of P. andromeda, the two species are clearly different, on the basis of granulation of the marginals,

armature of the adambulacrals, and general facies.

The specimen from off Colombia (Oregon Station 4855) has unusually broad superomarginal

plates, encroaching much furthrer onto the abactinal surface than they do in the specimens from

the Gulf of Mexico. It is an altogether more robustlooking starfish, and the arms, although broken,

obviously did not taper as sharply to an acute

point as in the Gulf specimens. Despite the difference in general appearance, however, there are

no specific differences.

MATERIAL EXAMINED.—Oregon Stations: 3654 (1)

[R=22 mm, r=7 mm, Rr=l:3]; 4855 (1) [R=82

mm, r=19 mm, Rr=l:4]. Alaminos Stations:

24/68-A-13 (1) [R=22 mm, r=6 mm, Rr—1:3];

1/68-A-13 (3) [R=82 mm, r=17 mm, Rr=l:7];

27/69-A-ll (1) [R=47 mm, r=ll mm, Rr=l:4].

Psilaster patagiatus Sladen

PLATE 8: FIGURES A, B

Psilaster patagiatus Sladen, 1889:232, pi. 41: figs. 3, 4, pi. 7:

figs. 11, 12.

Psilasteropsis patagiatus. Koehler, 1907:29.—Farran, 1913:7.—

Grieg, 1932:19.

This robust species has five arms of moderate

length, tapering smoothly to an acute point. The

paxillar area on the medium-sized disc is somewhat

reduced by the intrusion of the massive superomarginal plates, and still further reduced on the

arms. The small paxillae are in regular transverse

rows on the arms; they have 1-6 central and 12-18

peripheral spinules, small, acute, and frequently

all erect and in contact, like a wet paintbrush. The

superomarginal plates are large and encroach far

onto the dorsal surface. They are higher than wide

and are covered sparsely with small round granules,

mostly concentrated in the center of the plate. The

fascioles between the plates are shallow, and, on

the arms, the plates fit so closely that there are no

fascioles. The inferomarginal plates have granules

similar to those of the superomarginals and, in

addition, bear 2 or 3 small, flattened, acute spines,

appressed to the arm and directed upward. The

adambulacral plates, nearly square on their actinal

face, bear a furrow series of 7 or 8 slender contiguous spines and behind them a few small spinelets. The actinal interradial areas have about six

chevrons of polygonal plates, none extending

beyond the fifth or sixth inferomarginal. They are

covered with small spinules. The jaws are prominent, with regular series of small, blunt spines

and, at their apex, two larger, heavy, blunt, oral

spines.

Inside the mouth, on the abactinal surface of the

jaw plates above the oral spines, is a row of spines

similar to the furrow spines of the adambulacrals.

These in no way resemble the small group of

supraoral spines in Persephonaster except in position. The madreporite is small and surrounded by

a series of slightly enlarged paxillae.

The two lots in this collection are from off

Louisiana and off Venezuela, in 400-500 fathoms.

It seems odd that both this species and the previous

one should be recorded by Sladen from the Cape

Verde Islands; I would suspect some error in

labeling.

MATERIAL EXAMINED.—Oregon Station 4416 (2)

[R=64 mm, r=18 mm, Rr=l:5]. Alaminos Station

11A/68-A-7 (3) [R=24 mm, r=9 mm, Rr=l:3].

Other Atlantic Species of Psilaster

Other species of Psilaster reported from the North

Atlantic include P. andromeda (Muller and Troschel) and P. florae (Verrill), from the east and west

North Atlantic respectively, north of 35°, and thus

outside the range covered by this report; they are

probably one and the same species. They differ

from P. cassiope and P. patagiatus mainly in having

the marginals vertical, almost entirely confined to

the sides of the arms.

Persephonaster Wood-Mason and Alcock

Persephonaster Wood-Mason and Alcock, 1891:430. [Type, by

original designation, P. croceus.]

Psilasteropsis Fisher, 1906:1023. [Type, by original designation, P. cingulatus Fisher.]

For the present, it was thought best to leave this

genus and the two species in the previous genus
NUMBER 126

(Psilaster) in the genera assigned by the original

authors; however, considerable confusion exists

as to what really constitutes Persephonaster or

Psilaster. Psilaster is probably a perfectly good

genus, and most of the species in it are probably

correctly assigned, but Persephonaster, ill-defined

to begin with, has had assigned to it by various

authors some species which rightly belong to Psilaster, some which may be correctly assigned, and a

few which probably constitute a third genus. The

problem is insoluble without examination of the

types of the species in these two genera; unfortunately, type specimens of many of the species,

including the three species originally assigned to

Persephonaster by Alcock, are in the Indian

Museum and therefore unavailable to anyone outside India. Alcock (1893a) restricted the definition

of the genus without solving these problems.

Persephonaster echinulatus Clark

PLATE 8: FIGURES C, D

Persephonaster echinulatus H. L. Clark, 1941:20, pi. 3: fig. 2.

The five slender arms taper evenly to an acute

point. The abactinal surface is plane, and the

paxillae are not crowded; on the arms, there are

frequently large bare spaces between them. The

small paxillae bear 4—8 long slender spinules; occasionally, especially on the proximal parts of the

arms, a paxilla may be replaced by a two-, three-,

or four-valved simple pedicellaria of stout bent

spines. The superomarginal plates are, at least in

the interradial area, confined almost wholly to the

sides of the disc, and the inner margin bears a stout

erect spine or group of coalesed spinules (on the

arms, the position of the superomarginals depends

on the stretching or contraction of the abactinal

membrane). The rest of the plate is covered with a

fine furlike covering of slender spinules, especially

fine, numerous, and long in the wide sulci between

the plates. The inferomarginals, with a similar

but sparser covering of fine spinules, have their

distal edge slightly overlapping the plate ahead,

and a ridge down the center of the plate bears 2-4

long slender spines appressed to the arm and

directed forward, overlapping the plate ahead.

The actinal interradial area has four rows of

elongate imbricate plates; beyond them are several

rows of two or three rounded plates, and a single

row of actinolateral plates continues almost to the

end of the arm; all these plates are clothed in long

fine spinules and one or more stout pedicellariae

are usually present. The adambulacral plates bear a

curved fan of about five long, slender furrow spines;

behind them are spinules similar to those on the

actinolateral plates. The prominent, narrow mouth

plates bear two rows of spinules, one row on each

edge, and there are two large, flattened oral spines,

one on each jaw half. Inside the mouth, above the

oral spines, is a group of 5 or 6 short spinules. The

small hemispherical madreporite is covered with

deep gyri. In the abactinal center of the disc, an

epiproctal cone is usually conspicuous in preserved

specimens.

Ranging in depth from about 200 fathoms to

about 450 fathoms, this species has a much wider

distribution in the Gulf of Mexico, the Caribbean,

and the Lesser Antilles than Clark indicated.

MATERIAL EXAMINED.—Oregon Stations: 2771 (1)

[R=7 mm, r=3 mm, Rr=l:3]; 6696 (4) [R=20

mm, r=5 mm, Rr=l:4]; 1917 (2) [R=29 mm,

r=7 mm, Rr=l:4]; 4480 (1) [R=55 mm, r=7

mm, Rr=l:6 (specimen differs from others examined in having one very long inferomarginal

spine, a large spine on the superomarginal, and

flattened, leaf like oral spines)]; 4703 (2) [R=34

mm. r=8 mm, Rr=l:4]; 1507 (2) [R=29 mm,

r=7 mm, Rr=l:4]; 3583 (1) [R=45 mm, r=8 mm,

Rr=l:5]; 1920 (3) [R=22 mm, r=5 mm, Rr=l:5];

1909 (1) [R=28 mm, r=7 mm, Rr=l:4]. Alaminos

Stations: 21/68-A-13 (5) [R=30 mm, r=5 mm,

Rr=l:6]; 8/68-A-13 (1); 4/68-A-13 (1) [R=43 mm,

r=6 mm, Rr=l:8]; 10A/68-A-7 (5) [R=50 mm,

r=ll mm. Rr=l:5].

Other Atlantic Species of Persephonaster

The other two western Atlantic species of Persephonaster, P. leptactis and P. spinulosus, were

described by H. L. Clark (1941) from Cuba; the

first is known only from one specimen, and the

second is probably Psilaster patagiatus. P. leptactis

differs from P. echinulatus principally in having

very narrow paxillar areas on the arms, granulous

rather than spinous marginals, and 6-8 furrow

spines on the adambulacral plates; it lacks pedicellariae.
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Plutonaster Sladen

Plutonaster Sladen, 1885:610; 1889:81. [Type-species, by original designation, Archaster bifrons Wyville Thomson.]

Although at least seven species of Plutonaster have

been described from the Atlantic (two from the

east coast of North America, two from the Azores,

and one from the South Atlantic near Ascension

Island), examination of over a thousand specimens

in the National Museum of Natural History,

mostly labelled P. agassizi, led me to conclude that

there are probably no more than two or three species. The four small specimens of P. notatus (Sladen, 1889) in the National Museum of Natural

History, from the mid-Atlantic ridge south of the

Azores, represent a clear-cut species, evidently with

a very restricted distribution. On the basis of comparisons between the National Museum of Natural

History Plutonaster material and descriptions

and/or specimens of the other Atlantic Plutonasters, I suspect that they constitute one quite variable species. The spinulation of the marginals is

certainly variable, even in specimens from the same

or adjacent stations. This is true also for length

and width of arms, width of paxillar area on arms,

size of marginals, and armature on actinal face of

the adambulacrals, as well as granulation (or spinulation) of the interradial actinal plates.

It is interesting to note that frequently specimens

from one station will have short, broad arms with

massive marginals and little or no spinulation on

the superomarginals, granulose abactinal interradial plates, and a large prominent spine on the

actinal face of all the adambulacral plates, while

those from an adjacent station a few miles away and

at comparable depths have long, narrow arms,

with smaller marginals, one or more stout spines

on both series or marginals, a slender peglike spine

on many of the actinal interradial plates, and a

single large spine on the actinal face of only a few

distal adambulacrals. However, no correlation could

be discovered with range, depth, bottom type, or

other known factors, and other specimens from the

same general area show all variations between

these two extremes. An examination of the types of

the described species and of collections of Plutonaster in other museums would probably resolve

these into one abundant, variable, amphi-Atlantic

species. Until this question is resolved, the name

P. intermedius, to which the western Atlantic

Plutonasters from 0° to 30° N have usually been

referred, will be used for the purposes of this report.

Plutonaster intermedius (Perrier)

PLATE 9: FIGURES A, B

Goniopecten intermedius Perrier, 1884:251, pi. 7: figs. 1, 2.

pi. 4: fig. 4.

Plutonaster intermedius.—Terrier, 1894:316.

This rather large five-armed species has a muchflattened body form, with a large disc and moderate

to long arms. The paxillar area is broad on the

disc and narrow on the arms, with the only regularity in arrangement of paxillae occurring around the

interradial arcs. The polygonal paxillae are compact and much smaller in the center of the disc

than elsewhere. They are flat-topped and covered

with small granules which become spiniform

around the periphery of the paxillae. The superomarginals are broad and covered with a uniform

coating of small granules which become slightly

spiniform at the narrow, deep fascioles. Frequently,

each superomarginal, at least proximally, bears

a stout erect spine, bulbous at the base, on the outer

edge. These spines may occur all the way out to the

tip of the arm, or may be confined to the superomarginals of the interbrachial arc. In small

specimens, they are often lacking altogether. The

inferomarginal plates, directly opposite the superomarginals and of about the same size, are similarly

clothed with granules, and most of them bear one

or two stout erect spines on the outer edge.

The actinal interradial area is large and reaches

to about the sixth or seventh inferomarginal. The

distinct oval plates are in regular rows and are

covered with granules; a few may bear a slender

peglike spine in the center. The adambulacral

plates bear 7-10 nearly equal, long furrow spines

in straight longi-series, only slightly curved. Behind

them, on the actinal surface of the plate, is a loose

assemblage of much shorter spinelets or granules,

some of which may be grouped to form a crude

sort of pedicellaria (as are some of the spinelets or

granules on the actinal interradial plates). Frequently there is a large spine in the center of the

actinal face of the adambulacral plate; some specimens lack these altogether, some have one on every

adambulacral plate, and some have them only on

the distal adambulacrals. The mouth plates are
NUMBER 126 35

large, tumid, and covered with short spines; at the

oral edge is a cluster of longer spines. The madreporite is large and covered, or partly covered, with

paxillae larger than those on the rest of the dorsum.

The species seems to be restricted in these collections to the northern Gulf of Mexico, in 350-1,000

fms.

MATERIAL EXAMINED.—Oregon Station 2820 (1)

[R=21 mm, r=8 mm, Rr=l:3]. Alaminos Stations:

13A/68-A-7 (16) [R=26 mm, r=8 mm, Rr=l:3];

15D/68-A-7 (5) [R=28 mm, r=9 mm, Rr=l:3];

13D/68-A-7 (5) [R=20 mm, r=7 mm, Rr=l:3];

17B/68-A-7 (3) [R=ll mm, r=4 mm, Rr=l:3];

15H/68-A-7 (15) [R=23 mm, r=9 mm, Rr= 1.2.5];

24/68-A-13 (2) [R=lll mm, r=28 mm, Rr=l:4];

27/68-A-13 (1) [R=80 mm, r=25 mm, Rr=l:3];

23/68-A-13 (2) [R=lll mm, r=29 mm, Rr=l:4].

Dytaster Sladen

Dytaster Sladen, 1885:608; 1889:60. [Type, by original designation, D. nobilis Sladen.]

Crenaster Perrier [non d'Orbigny], 1885:71; 1894:306. [Type

by original designation. C. mollis Perrier.]

Disc rather small, five arms long, thin; ambulacral

plates large, high; dorsal tegument thin, flexible

(as if it were several sizes too large for the starfish);

rays usually quite carinate; marginals rectangular,

opposite, straight-edged, usually bearing one prominent acute spine; superomarginal plates thin, confined to side of arm; dorsal paxillae tiny; actinal

interradial areas of moderate size; mouth plates

very prominent; adambulacral furrow spines in

straight, subequal series; madreporite large, covered

with paxillae.

Dytaster is a deepwater genus, specimens in this

collection reaching a greater depth than any other

seastars in the area. They are also, apparently, the

most abundant of the starfishes in depths below

500 fathoms (to 2,200 fathoms) in the Gulf of

Mexico. They are detritus feeders: bits of algae,

many forams, and worm tubes and small molluscs

have been found in the stomach.

With-additional material now available, it is

easy to see that the five species described from the

North Atlantic really constitute only one variable

species, for which the name D. insignis has priority. Sladen's Dytaster nobilis, from off Buenos

Aires, may be simply another variation of the same

species, but as I have not had an opportunity to

examine specimens, it should be maintained as a

separate species for the time being, on the basis

of Sladen's description of shorter, broader arms,

larger disc, and larger paxillae sheathed in membrane, as well as differences in adambulacral armature.

Dytaster insignis (Perrier)

PLATE 9: FIGURES C, D

Archaster insignis Perrier, 1884:253, pi. 9: fig. 5.

Archaster grandis Verrill, 1884: 218.

Dytaster exilis Sladen, 1889:65, pi. 2: figs. 3, 4, pi. 4: figs 9, 10.

Dytaster exilis gracilis Sladen, 1889:68, pi. 2: figs. 3, 4, pi. 4:

figs. 9, 10.

Dytastei exilis carinata Sladen, 1889:69.

Dytaster madreporifer Sladen, 1889:70, pi. 3: figs. 3, 4, pi. 32:

figs. 5. 6.

Dytaster biserialis Sladen, 1889:77, pi. 10: figs. 3, 4, pi. 13:

figs. 3, 4.

The five narrow, elongate arms, and small- to moderate-sized disc of this species are distinctive. The

dorsum, covered with small "fuzzy" paxillae, is

usually somewhat wrinkled and velvety looking.

The crowded, low paxillae are crowned with 5—10

long, slender, thorny spinules; on the midline of

the arms, they are fewer and may be reduced to a

few spinules without paxillar column. Occasionally

a specimen may have a few two-, three-, or fourvalved pedicellariae on the dorsal surface near the

base of the arms. The superomarginal plates are

prominent, mostly confined to the sides of the arms,

and each bears a single, large, erect spine near the

upper edge. The rest of the plate is covered with

fine spinules.

The inferomarginal plates are similar, except

that they may bear, in the interradius, more than

one spine. The small oval or polygonal plates of

the actinal interradial area are imbricate and covered with small spinules; there may be up to a

dozen large two-, three-, or four-valved pedicellariae in each interradial area (but in some specimens these may be altogether lacking). Actinal

interradial plates extend to about the fifth inferomarginal. The adambulacral plates, rectangular

and well separated from one another, bear 8-12

slender, subequal furrow spines in close longitudinal series. The actinal face of the plate bears a

random group of small spinelets. Distally there

may be a single large spine in the center of the
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

plate. The mouth plates are large, prominent, and

distinctive; they resemble somewhat the eletra of a

beetle and are covered with small spinelets. Their

armature is similar to that of the adambulacral

plates, with a cluster of heavier spines at the apex.

The madreporite is very large, near the margin,

and covered with paxillae which are larger than

the rest of the dorsal paxillae.

MATERIAL EXAMINED.—Oregon Stations: 2199 (3)

[R=54 mm, r=8 mm, Rr=l:5]; 2572 (1) [R=80

mm, r=9 mm, Rr=l:8]; 2821 (2) [R=57 mm, r =

10 mm, Rr=l:6]; 2813 (1) [R=20 mm, r=8 mm,

Rr=l:S]. Alaminos Stations: 5B/68-A-3 (1) [R=

54 mm, r=9 mm, Rr=l:5]; 7D/68-A-3 (2) [R=50

mm, r=12 mm, Rr=l:5]; 3B/68-A-3 (1) [R=40

mm, r=10 mm, Rr=l:5]; 3B/68-A-7 (1) [R=72

mm, r=13 mm, Rr=l:7]; 4A/68-A-7 (3) [R=8

mm, r=4 mm, Rr=1.2 (juveniles)]; 4E/68-A-7 (8)

[R=86 mm, r=12 mm, Rr=l:8]; 12B/68-A-7 (5)

[R=8 mm, r=5 mm, Rr=l:2 (juveniles)]; 9/68-

A-13 (1) [R=39 mm, r=9 mm, Rr=l:4]; 27/69-

A-ll (2).

Blakiaster Perrier

Blakiaster Perrier, 1881a:28; 1885:265. [Type, by monotypy,

B. conicus Perrier.]

Blakiaster conicus Perrier

PLATE 10: FIGURES A. B

This small species has a moderately broad disc; the

five arms are broader than high and taper evenly

to a subacute end. The disc dorsum is slightly

larger than the space confined by the marginals, so

that dorsal covering seems "too large," and the marginals are not visible from above, except on the

arms. The dorsal paxillae are large, low, and

rounded, with about 25 thorny, subclavate spinules,

thick and almost granuliform in the center of the

central paxillae, but fine and long around the periphery and on the outer paxillae. The superomarginal plates are ovate, slightly tumid, and have a

uniform covering of fine, thorny spinules, thicker

and clavate in the center of the plate and longer

and more acute around the edges. The inferomarginal plates are subprismatic, due to the presence

of a central vertical ridge of spine-bearing tubercles

near the upper edge of the plate, which carry three

strong, distally directed spines of moderate length.

The rest of the plate is covered with spinules similar to those of the superomarginals. In the interradii, the superomarginals extend considerably

beyond the inferomarginals. Actinal interradial

plates are rounded or subpolygonal and bear divergent spinules; there are a few unpaired median

plates, and a single row extends almost to the middle of the arm. The adambulacral plates bear a furrow series of 4 or 5 long, slender spines, a second

row of four slightly shorter spines, and about two

more irregular rows of much shorter spines. The

mouth plates are oval, convex, and covered with

small spinules distally, clavate spines proximally,

and terminate in a cluster of heavy subprismatic

apical spines. The madreporite is of moderate size,

covered with coarse gyri, and located a little more

than halfway between the center of the disc and

the margin. No pedicellariae were noted.

This species is apparently restricted to the Antilles, in 92-200 fathoms.

MATERIAL EXAMINED.—Oregon Station 6715 (1)

[R=26.5 mm, r=10 mm, Rr=l:2.5].

Tethyaster Sladen

Plutonaster (subgenus Tethyaster) Sladen, 1889:101. [Type,

by original designation, Asterias subinermis Philippi.]

Sideriaster Verrill, 1899:210. [Type, by original designation,

S. grandis Verrill.]

Moiraster Sladen, 1889:192. [Type, by monotypy, Archaster

magnificus Bell.]

Anthosticte Fisher, 1911:417. [Type, by monotypy, A. aulophora Fisher.]

The genus Tethyaster has been very thoroughly reviewed and revised by Clark and Clark (1954) and

is now so well defined as to need no further discussion. The only reservation I personally have

about their work as regards the Atlantic Tethyasters concerns the separation of T. vestitus and T.

magnificus, the former occurring from New Jersey

to the Orinoco River and the latter from St. Helena

and Ascension Islands. In view of the Caribbean

flavor of the St. Helena echinoderm fauna (see

Mortensen, 1933a; Madsen, 1950; and Downey,

1968), and the fact that specimens keying out to

T. magnificus have been taken in the Caribbean

(Leviton, personal communication), the single

character used by Clark and Clark, that is, when

during growth of the starfish the enlarged and flat-
NUMBER 126 37

tened spines of the actinal plates develop, seems

to me a matter of individual variation, not worthy

of even subspecific separation. This character is

further invalidated by the fact that a small T. magnificus and a large T. vestitus may be armed exactly

the same. Of the five species of Tethyaster from

the Atlantic, subinermis is confined to the eastern

Atlantic and Mediterranean, and T. pacei is known

only from South Africa. Only T. grandis and T.

vestitus come within the scope of this report.

Tethyaster grandis (Verrill)

PLATE 10: FIGURES C, D

Sideriaster grandis Verrill, 1899:220, pi. 30: figs. 8-8fc; 1914:

21; 1915:192, pi. 12: figs. 5, 5b.

Tethyaster grandis.-Clark and Clark, 1954:17, fig. \b, pis. 11,

12.

This stellate species has a large disc and five broadly

based arms which taper evenly to a moderately

acute point. Both actinal and abactinal surfaces

are extremely plane and even, the only height being

defined by the marginals. The dorsal paxillae are

less than twice as high as wide and arise from a

lobed base to a rounded top which bears 3-8

granuliform central spinules and about a dozen

longer peripheral spinules. The high, narrow superomarginals, confined mostly to the sides of the

rays, are densely covered with granules which become small and spiniform at the deep interplate

fascioles. The inferomarginals, of similar size and

shape, bear three or more large, flattened lanceolate spines and many small flattened spinules. The

actinal interradial areas are large, with large subpaxilliform plates in regular rows (there are frequently a few odd interradial actinals just below

the jaw). They are covered with large granules

and small peripheral spinules, and a single row

extends nearly to the end of the arm. The adambulacral plates project slightly into the furrow and

bear three flattened furrow spines, the central one

at right angles to the groove and the other two

nearly parallel to it. Behind, on the actinal face

of the plate, are 3 or 4 stouter truncate spines, and

behind them are a few much smaller spines and

frequently 1 or 2 stout, short, three-valved pedicellariae. The mouth plates are slightly raised

above the actinal surface and bear many stout

granuliform spinules, with a cluster of stout truncate spines at the apex. In many specimens, threeor four-valved pedicellariae are scattered all over

the actinal surface. The tube feet are small and

pointed. The madreporite is large and flat, with

shallow radiating gyri.

JUVENILES.—Young specimens of Tethyaster differ

from the adults in that the marginals extend more

onto the dorsal surface, the spines are not developed on the inferomarginals, the madreporite is

small and nearly concealed by paxillae, and the

adambulacral furrow spines are not flattened and

are more palmate.

Known previously only from the Gulf of Mexico,

this species has a depth range of 35—162 fathoms.

MATERIAL EXAMINED.^SIIver Bay Station 5106 (3)

[R=16 mm, r=7 mm, Rr=l:2]. Oregon Stations:

5913 (3) [R=il8 mm, r=6 mm, Rr=l:3]; 3621, 1.

Oregon II Station 10559 (1) [R=82 mm, r=21 mm,

Rr=l:4]. Alaminos Stations: 13/64-A-10 (1) [R=

105 mm. r=31 mm, Rr=l:3]; 5/68-A-13, 2.

Other Species of Tethyaster

Tethyaster vestitus, the other species falling within

the scope of this report, bears a large flat spine on

the actinal plates, and the inferomarginal spines

are broadly truncate, rather than sharp pointed. It

ranges from 08° to 40° north, and apparently does

not occur east of 75° west. Its depth range is 44-

146 fathoms.

Dipsacaster Alcock

Dipsacaster Alcock, 1893a:87; 1893b: 172. [Type, by original

designation, D. sladeni Alcock.]

Rays five; form depressed, disc broad, with welldeveloped actinal interradial areas; marginal plates

large; inferomarginals broader than superomarginals with tuft of spines at outer edge; superomarginals sometimes with 1-3 small tubercles; paxillae

with round or stellate base, tall pedicel crowned

with tuft of many small, crowded, erect spinelets;

papulae all over abactinal surface, 5 or 6 around

each plate; actinal intermediates bearing paxilliform group of spinules; fasciolar channels from

inferomarginals to adambulacrals; adambulacrals

with palmate or pectinate cylindrical or compressed

furrow spines, smaller spinelets on actinal face;

mouth plates prominent, broad, with numerous
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

spinules; madreporite large, hidden by paxillae;

tube feet pointed.

Dipsacaster antillensis Halpern

PLATE 11: FIGURES A, B

Dipsacaster antillensis Halpern, 1968:231-240.

Among the seastars collected by the Alaminos in

the middle of the Gulf of Mexico were two small

specimens of Dipsacaster antillensis, described from

the western Atlantic by Halpern (1968), from Little

Bahama Bank, in less than 900 meters. These two

small specimens are from the middle of the Gulf

of Mexico, in depth greater than 3,200 meters.

They are evidently juveniles of D. antillensis, and

quite apart from size (much smaller than Halpern's

specimens), have quite small dorsal paxillae, not at

all compact at least on the arms; the paxillar

spinules are longer, finer, and very thorny; the

inferomarginals are armed with a central, rather

than distomarginal, group of spines, at least one

of which is quite thick and thorny; massive pedicellariae are present on some of the inferomarginals, the actinal interradials, and many of the

adambulacral plates; the adambulacral furrow

spines are fewer in number and distinctly palmate; the madreporite is, as yet, rather small and

concealed by enlarged paxillae.

The disc is large, but the arms are moderately

long, narrow, and tapered, rather than being broad

and subpetaloid as in most Dipsacasters; in this

and in the armature of the inferomarginals, they

resemble D. nesiotes Fisher (1910b), but the dorsal

paxillae are decidedly different from those of Fisher's Hawaiian species. These specimens are quite

obviously juvenile, but Dr. Halpern has examined

these and other specimens and determined that

they are indeed young D. antillensis.

MATERIAL EXAMINED.—A laminos Stations: 9/69—

A-13 (1) [R=12 mm, r=4 mm, Rr=l:3]; 29/69-

A 13 (1) [R=12 mm, r=3.5 mm, Rr=l:4].

Family GONIOPECTINIDAE Verrill, 1899

This family is mainly distinguished by having cribriform organs between all the marginals; they are

continuous with channels on the oral surface covered with webbed spinelets, between transverse rows

of plates. The marginals are thin, high, and lamelliform, smooth and covered with membrane. The

two rows of tube feet are pointed and the ampullae

are single. Superambulacral ossicles are present.

The abactinal surface is paxillose.

Key to the Genera of Goniopectinidae

With odd interradial marginals and corresponding unpaired series of actinal interradial plates. .

Prionaster

Without odd interradial marginal and actinal plates Goniopecten

Goniopecten Perrier

Goniopecten Perrier, 1881a:24. [Type, by original designation,

G. demonstrans Perrier.]

There is only one Atlantic species.

Goniopecten demonstrans Perrier

PLATE 11: FIGURES C, D

Goniopecten demonstrans Perrier, 1881a:24 [without description]; 1894:295 [described]; Verrill, 1899:213, pi. 27.

This large five-armed species has a small disc, and

the moderately narrow arms taper to an acute

point. The paxillar area, broad on the disc, is quite

narrow on the arms. The paxillae are polygonal

(many rectangular), very crowded, with short paxillar columns, and crowned with an even coating of

small granules; they become progressively smaller

on the arms. The large superomarginals are confined entirely to the dorsal surface (in the adult);

there may be a small sharp spine on the distal edge

of some of the superomarginals, but in most specimens they are smooth and without ornamentation.

The long, broad inferomarginal plates form both

the sides of the arms and the ventral border; they

are similar to the superomarginals. Both series of

marginals are bordered by a row of small webbed

spinules, between which are deep fascioles that extend across the actinal surface to the mouth plates.
NUMBER 126 39

There are no odd interradial marginals. The actinal interradial plates, plane, polygonal, and

slightly imbricate, are in double rows between the

fascioles and extend only to the fifth or sixth inferomarginal; they may bear a few small pricklelike spinules. There is no odd interradial series.

The adambulacral plates are nearly square, slightly

concave on their actinal face, and the proximal inner corner projects slightly into the groove. They

bear a furrow series of about nine short, subequal

spines. The actinal face of the plate is bare, save

for a row of small spinules on the actinal edge.

The mouth plates are raised and prominent; they

stand nearly at right angles to the actinal surface.

They are covered with tiny prickles and bear, at

the apex, a cluster of short, stout, sharp spines. The

madreporite is very large, flat, and bare, and covered with fine radiating gyri. All plates except

those of the paxillar area are covered with a

smooth, slippery skin.

This species occurs in the Gulf and Caribbean, in

299-470 fathoms.

MATERIAL EXAMINED.—Oregon Stations: 6696 (6)

[R=104 mm, r=21 mm, Rr=l:5]; 6703 (10) [R=

114 mm, r=16 mm, Rr=l:7]; 5929 (1) [R=138

mm, r=28 mm, Rr=l:5]; 4702 (1) [R=125 mm,

r=25 mm, Rr=l:6]; 6697 (2) [R=108 mm, r=23

mm, Rr=l:5]; 4855 (1) [R=109 mm, r=21 mm,

Rr=l:5]; 4902 (1) [R=21 mm, r=6 mm, Rr=l:4;

juvenile, with superomarginal spines, small madreporite, large epiproctal cone, marginals more vertical than in adult]. Oregon II Station 10602 (1)

[R=109 mm, r=22 mm, Rr=l:6]. Alaminos Stations: 8/68-A-13 (1) [R=137 mm, r=22 mm, Rr

=1:5]; 27/69-A-ll (1) [R=133 mm, r=25 mm,

Rr=l:6; 15/68-A-13 (1) [R=28 mm, r=9 mm,

Rr=l:S]; 11A/68-A-7 (1) [R=8 mm. r=3 mm,

R r= 1:2.5].

Prionaster Verrill

Prionaster Verrill, 1899:205. [Type, by original designation.

P. elegans Verrill.]

There is only one species, Prionaster elegans Verrill, known from the Atlantic.

Prionaster elegans Verrill

PLATE 12: FIGURES A, B

Prionaster elegans Verrill, 1899:216, pi., 6gs. 4, Aa-c; 1915:197,

pi. 12: figs. 4-4<r.-Fisher, 1919:50.

The five arms of the species are narrow and taper

gradually to an acute point; they are slightly constricted near the base. The disc is of moderate

size, and the paxillar area on the arms is narrow.

The paxillae are in regular transverse rows on the

arms. They are small, rounded, moderately high,

not crowded, and crowned with 8—12 short subclavate spinules. The superomarginals are confined

to the sides of the rays, making the sides nearly

vertical. They are smooth in the interradial arc

but ornamented with small, irregular, glassy lowrelief tubercles on their upper surface on the arms.

Most bear a short, stout spine with a sharp hyaline

tip. The inferomarginals are bare on their lateral

surface; a few may bear a small spine near the

distal edge. The broad, deep fascioles between the

marginals extend to the mouth and are bordered

with short, webbed spinules. There is an odd interradial marginal, and two rows of actinal plates

extend from this to the base of the mouth plates.

Two rows of plates extend from the two marginals

on either side, but there are no actinal interradial

plates beyond the third marginal. The adambulacral plates are semicircular and project slightly

into the furrow; they bear about 10—14 spines

around the free edge, of which approximately eight

belong to the furrow series. The actinal face of

the adambulacral plate is bare. The mouth plates

are oval, slightly tumid, and bear scattered small

spinules; at the apex are two strong, stout spines;

on either side, within the mouth, is a row of spines

like those of the adambulacrals. The madreporite

is small and covered with small spinules rather than

gyri-

This species is known only from the Gulf of

Mexico, in 150-200 fathoms.

MATERIAL EXAMINED.—Oregon II Station 10858

(1) [R=37 mm, r=7 mm, Rr=l:5].

Family BENTHOPECTINIDAE Verrill, 1894

This family is characterized by a unique pair of

dorsal muscle bands on the flexible arms; it is

thought they may be used for swimming. The marginals are alternate and imbricate and bear long

spines. Pedicellariae are pectinate. The tube feet,

in two rows, are suckered and the ampullae are
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

double. There are no superambulacral ossicles. It

is possible that some members of this family are

luminescent, but reports to this effect are unconfirmed.

Key to the Genera of Benthopectinidae

1. Papular pores confined to specialized areas on the disc (papularia) 2

Papular pores not confined to papularia 3

2. Papulae in U- or V-shaped papularia, not swollen Cheiraster

Papularia not U- or V-shaped, more or less swollen Pectinaster

3. Odd interradial marginals present Benthopecten

No odd interradial marginals Luidiaster1

1

 Although no species referrable to Luidiaster have been taken in these collections, the genus

is included in this key because some of the species here included have been referred to it from

time to time in the past.

Benthopecten Verrill

Benthopecten Verrill, 1884:218. [Type, by original designation, B. spinosus Verrill.]

Pararchaster Sladen, 1885:610. [Type, by original designation,

P. pedicifer Sladen.]

There is only one species of Benthopecten known

from the area covered by this report.

Benthopecten simplex (Perrier)

PLATE 12: FIGURES C, D

Archaster simplex Perrier, 1884:264, pi. 1: fig. 8.

Pararchaster simplex.—Perrier, 1894:254.

Benthopecten simplex.—Fisher, 1911:143.

This fragile seastar has five long, slender arms and

a small disc. The thin dorsal membrane is set with

small, rounded plates bearing 1—3 tiny thorny

spinules. These plates are crowded on the disc,

some even fused together, but are widely scattered

on the arms. There are 1-10 large spines in the

center of the disc. The anus is large and surrounded by a cluster of small spinules. The long,

oval superomarginal plates, confined mostly to the

side of the arms, have a central mammelon bearing a long, stout, thorny spine; 1 or 2 small spinules

may occur near the base of this spine. The distal

edge of the plate overlaps the proximal edge of the

plate ahead. There is a larger odd interradial marginal, rather triangular in shape, confined wholly

to the dorsal surface and bearing a single very large,

acute spine. The smaller inferomarginal plates are

similar to the superomarginals and alternate with

them. There is almost always a large pectinate

pedicellaria between all the inferomarginals. In

the actinal interradial area, there is a single row of

2—6 small, rounded plates, each bearing a small

acute spinule; some or all of these plates may be

replaced by 1 or 2 large pectinate pedicellariae.

The adambulacral plates bear six long, somewhat

flattened, and blunt spines, parallel to the groove

and fused at the base; on the actinal face is a single

large spine and 1 or 2 small, slender spines. The

mouth plates are rounded and tumid; there are a

pair of stout spines at the apex, one to each jaw

half, and three smaller spines on each side. There

are 2-4 smaller spines on the actinal face of each

jaw half. The madreporite is raised, irregular, and

deeply channeled; it forms part of the base of one

of the odd interradial superomarginals.

This is a deepwater species, occurring from

1,100-1,500 fathoms from Nova Scotia to the central Gulf of Mexico.

MATERIAL EXAMINED.—Alaminos Station 4/69—A—

13 (7) [R=48 mm, r=6 mm, Rr=l:8]. Oregon

Stations: 2572 (1) [r=9 mm (all arms broken)];

2574 (6) [r=9 mm (all arms broken)]; 2571 (1)

[R=:48 mm, r=6 mm, Rr=l:8].

Cheiraster Studer

Cheiraster Studer, 1884:129. [Type, by original designation,

C. gazellae Studer.]

No odd interradial marginal plate; papulae form

bilobed U- or V-shaped group on disc dorsum near

base of rays; dorsal plates paxillate; marginals covered with small spinules or granules; furrow spines

continuous with a series of smaller spines around

outer edge of adambulacral plate.

Five species of Cheiraster have been reported
NUMBER 126 41

from the West Indies: C. mirabilis (Perrier), C.

coronatus (Perrier), C. echinulatus Perrier, C. planus Verrill, and C. enoplus Verrill. Perrier (1894),

Ludwig (1910), and Verrill (1915) all considered

C. coronatus no more than a subspecies of C. mirabilis at most, and I am inclined to view it as well

within the range of variation of C. mirabilis; the

name coronatus should be considered a junior synonym of mirabilis. Based on Verrill's (1915) descriptions, I am also inclined to believe that his

C. planus is simply a large mirabilis. None of the

differences he notes between the two species seem

to be anything more than normal growth factors,

or characters well within the range of variation.

Cheiraster enoplus is readily distinguished from

the other species by its numerous papular pores in

paired, elongate, leg-of-mutton-shaped papularia,

and by its truncate, grooved-tipped preoral spines.

It usually has a dense cluster of fairly long, acute

spines in the center of the disc (these may also be

present in C. mirabilis). C. echinulatus seems a

perfectly good and distinct species; it is quite small,

with a more broadly stellate shape than the other

species (Rr=l:4). It has fewer marginal plates

(less than 20, as opposed to 35-60 in other species).

The dorsal plates are also more distinctly paxillate.

Key to the Species of Cheiraster

1. About 200 papular pores per papularium; preoral spines truncate C. enoplus

Less than 50 pores per papularium; preoral spines not truncate 2

2. Rr=l: 4 (approximately) C. echinulatus

Rr—1:8 (approximately) C. mirabilis

Cheiraster mirabilis (Perrier)

PLATE 13: FIGURES A, B

Archaster mirabilis Perrier, 1881a:27.—Perrier, 1884 (part);

256, pi. 8: figs. 7, 8, pi. 9: fig. 4, pi. 10: fig. 3.

Cheiraster coronatus Perrier, 1894:271.-Ludwig, 1910:455.

Archaster coronatus.—Perrier, 1884:262; 1894:271.

Cheiraster mirabilis.—Verrill, 1915:124, pi. 14: figs. 5, 5a.

Cheiraster mirabilis coronatus.—Verrill, 1915:127.

Cheiraster planus Verrill, 1915:133, pi. 18: fig. 2.

The moderate-sized disc and five long, flexible arms

of this species are flat and plane. The dorsal plates

are small, hemispherical, and crowded and bear, on

their upper surface, a ring of up to a dozen small

thorny clavate spinules around the base of a large,

central, acute spine. The papular pores are in two

U-shaped rows of approximately 7 or 8 each, in

papularia between the bases of the arms and the

center of the disc. The superomarginals are oval,

set diagonally on the arms, and confined to the

sides of the arms, encroaching very little on the

dorsum. They bear a single large, acute spine on

a tubercle in the center of the plate, and the rest

of the plate is covered with fine spinules carried

on glassy bosses. They number about 40-50 on

adult specimens. The inferomarginal plates are

similar and alternate with the superomarginals.

Their central spine is slightly smaller, and they

may bear 1—5 supplementary smaller spines, especially proximally. They are almost wholly confined

to the ventral surface. The actinal interradial areas

have 1-3 rows of flat, rounded plates, none extending beyond the third marginal. They usually bear

a central spine and a number of small, blunt spinules. A pectinate pedicellaria may occur between

two adjacent plates. Pectinate pedicellariae may

also occur between adambulacral and inferomarginal plates.

The adambulacral plates project into the furrow

and bear a furrow series of about ten slightly divergent, flattened spines. Other smaller spinelets

continue around the side margins of the plate, and

the actinal face bears a single large spine. The

mouth plates are broad, and each half bears two

large acute preoral spines and 7-9 lateral spines,

as well as numerous small epioral spinules. The

madreporite is irregular in shape, may be small or

large, and is deeply channeled with coarse gyri.

This species occurs from George's Bank to the

West Indies to the western Gulf of Mexico, in

200-800 fathoms.

MATERIAL EXAMINED.—Oregon Stations: 1909 (1)

[R=120 mm, r=15 mm, Rr=l:8]; 1906 (4) [R=

128 mm, r=13 mm, Rr=l:9]; 1507 (2) [R=60 mm,

r=10 mm, Rr=l:6]; 1917 (1); 4702 (1) [R=50

mm, r=8 mm, Rr=l:6 (specimen abnormal in having very narrow arms and scattered papulae on disc
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and arms outside the papularia)]; 6696 (1) [R=82

mm, r=ll mm, Rr= 1:7.5]. Alaminos Station 8/68-

A-13 (1) [R=ll mm, r=4 mm, Rr=1.3].

Cheiraster echinulatus (Perrier)

PLATE 13: FIGURES C, D

Archaster echinulatus Perrier, 1876a:268; 1884:263, pi. 10: fig.

4.

Cheiraster echinulatus.—Perrier, 1894:278.-Verrill, 1915:129,

pi. 14: figs. 2, 3, pi. 19: fig. 2, pi. 25: fig. 1.

Pectinaster echinulatus.—Ludwig, 1910:449.

This is the smallest of the Atlantic Cheirasters. Its

form is more stellate, and the arms more rigid. The

dorsal plates are distinctly paxillate, and each bears

8—25 small spinous peripheral granules and a small

acute central spine. Each papular area has 3—8

pores at the base of the arms, often not in any particular arrangement, but sometimes, especially in

larger specimens, distinctly bilobed. The superomarginal plates (15—26) are large, rectangular, and

encroach considerably on the disc, so that the paxillar areas on the arms are quite narrow. Except

for the first two interradial plates, they bear a single

erect, acute spine, and there may be other small

spines around it, especially on the proximal superomarginals. The rest of the plate is clothed in

small spinous granules. The inferomarginal plates

bear one large spine and up to eight smaller spines

on the upper (or lateral) half of the plate, and the

lower (or ventral) half is covered with small spinules.

The marginals may be opposite or alternate. The

interradial actinal areas have a single row of 6 or 7

round plates, not extending beyond the second inferomarginal. They are covered with small acute

spines. The adambulacral plates project strongly

into the furrow and bear a palmate series of about

ten long furrow spines. On the actinal face of the

plate there are usually two large subequal spines

and a few small spinules. On each half of the

broadly rounded mouth plate are two stout preoral

spines and about eight small lateral spines; the

actinal face bears one or two large epioral spines

and a row of 2-5 smaller spines. The madreporite

is of modest size, and deeply grooved with coarse

gyri.

This species occurs principally in the Gulf of

Mexico, although the Albatross took it also in the

West Indies, according to Verrill. Its depth range

is from 65-400 fathoms. I do not believe this species gets much over 35 mm (R), and specimens of

less than half this size are fully mature.

MATERIAL EXAMINED.—Oregon Station 5913 (1)

[R—35 mm, r=8 mm, Rr=l:4]. Alaminos Stations:

11A/68-A-7 (1) [R=10 mm, r=3 mm, Rr=l:S];

20/65-A-9 (9) [R=16 mm, r=3 mm, Rr=l:4];

15/65-A-9 (1) [R=25 mm. r=6 mm. Rr=l:4].

Cheiraster enoplus Verrill

PLATE 14: FIGURES A, B

Cheiraster enoplus Verrill, 1915:135, pi. 18: fig. 1.

This is a more massive-appearing species than the

previous two, with long narrow arms and a rather

small disc. The dorsal plates are small, crowded,

slightly rounded, and not at all paxillate. They

bear 6—8 tiny spinules on top, and a few of the

central disc plates are enlarged and bear, on a

mammelon, a large central spine. The papularia

are distinctive: they are shaped like a pair of rabbit

ears and extend from near the disc center to beyond

the fourth superomarginal plate on the arms. Each

contains about 200 papular pores. The superomarginals are nearly square, set obliquely on the arms,

and do not encroach on the dorsal surface. They

bear a large, stout, erect spine in the center of the

plate and are sparsely clothed with fine spinules.

The inferomarginal plates are larger, rectangular,

and bear 3 or 4 large spines, several smaller spines,

and a number of spinules. Many also bear a pectinate pedicellaria near the actinal margin of the

plate. The actinal interradial area has 2-4 rows

of rounded plates, none of which extend beyond

the third inferomarginal. Each plate bears one

large slender spine and several spinules, and between many adjacent plates is a pectinate pedicellaria. The adambulacral plates project into the

furrow, and there are 6 or 7 long, slender, subequal

furrow spines, not acute.

A large stout spine and a few spinules are on the

actinal face. The mouth plates are not very large,

are broadly rounded on the oral margin, and bear

one or two large, truncate preoral spines with tips

grooved or bifurcate, and 6 or 7 smaller, slender

lateral spines. There is a single pair of epioral

spines and a row of small spines on either side of
NUMBER 126 43

the suture between the mouth plates. The raised

madreporite is of moderate size and covered with

coarse gyri.

This species has been recorded only from east of

Cuba, in 400 fathoms, and this record, from off

Guadeloupe, in 410—460 fathoms.

MATERIAL EXAMINED.—Oregon Station 6703 (1)

[R=104 mm, r=14 mm, Rr=l:8].

Pectinaster Perrier

Pectinaster Perrier, 1885b:7O. [Type, by original designation,

P. filholi Perrier.]

Papularia of enlarged dorsal plates raised above

dorsal surface, papular pores not in bilobed arrangement; triangular adambulacral plates projecting so strongly across furrow as to meet, or nearly

meet, in middle of groove.

Fisher (1911) considered the mode of attachment

of dorsal muscle bands in the arms at the proximal

terminus a useful and important character for separating these genera, but such an internal character

cannot be examined in unique specimens, dry specimens, or very small specimens, and as Pectinaster

and Cheiraster present a clear and distinct external

character in the nature of the papularia, dissection

is unnecessary anyway.

Verrill (1915) described three new species of

Pectinaster from the West Indies: P. mixtus, P.

gracilis, and P. dispar. He also reported on two

previously known species, P. vincenti and P. oligoporus. All of these seem to be perfectly good species, but only two, P. mixtus and P. gracilis, were

taken in these collections.

Key to the Species of Pectinaster

Dorsal plates tessellate. crowded, many enlarged P. mixtus

Dorsal plates rounded, subpaxillate, not crowded P. gracilis

Pectinaster mixtus Verrill

PLATE 14: FIGURES C, D

Pectinaster mixtus Verrill, 1915:140, pi. 5: fig. 2, pi. 15: fig.

2, pi. 17: fig. 1.

The disc is small and the five arms slender and

moderately long. The dorsal plates are subtessellate

and bear very small thorny peripheral spinules

which might almost be called granules. A larger

erect spinule or small spine occurs in the center

of most plates, some of which are enlarged. The

3-7 papular pores in a median cluster between

the base of each arm and the center of the disc

are associated with several enlarged plates (surrounding a central papular pore), but not necessarily four, as Verrill states. Verrill's type of this

species has four enlarged plates in two of the

papularia, 5-7 in two others, and eleven in the

fifth. The superomarginal plates are large and encroach quite far onto the dorsal surface; they are

set obliquely on the arms, giving the starfish the

appearance of having its ambitus defined by braid

trim. Each plate bears an erect acute spine near the

outer margin, whose length is less than the width

of the plate. The two interradial superomarginals

usually lack a spine. The superomarginals are

densely covered with minute granule-like spinules.

The inferomarginals alternate with the superomarginals and bear one large and 2-7 smaller

spines; the rest of the plate is covered with spinules

larger than those of the superomarginals.

The actinal interradial areas bear one or two

series of rounded spinule-covered plates, none extending beyond the second inferomarginal. A large

pectinate pedicellaria usually occurs between each

two. The adambulacral plates project strongly

across the furrow and bear a marginal series of

7-9 long, slender furrow spines and two large,

subequal, acute, sometimes slightly curved spines

on the actinal face. Each mouth plate bears four

very large, stout preoral spines and about nine

small lateral spines on each side. The actinal surface bears several pairs of smaller spines. The

madreporite is small, irregular, and covered with

coarse gyri; it almost or quite touches the superomarginals.

This species has a wide distribution in the West

Indies and the eastern Gulf of Mexico, although

it apparently does not get into the western Gulf.

It has a considerable depth range, from about 70

fathoms to about 1,000 fathoms.
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

MATERIAL EXAMINED.—Oregon Stations: 3637 (1)

[R=41 mm, r=8 mm, Rr=l:5]; 4398 (6) [R=29

mm, r=5 mm, Rr=l:6]; 33 (2) [R=18 mm, r=4

mm, Rr= 1:4.5.]

Pectinaster gracilis Verrill

PLATE 15: FIGURES A, B

Pectinaster gracilis Verrill, 1915:145, pi. 6: 6g. 1, pi. 14: fig.

4, pi. 15: figs. 1-lfe.

This small species has slender, evenly tapered arms,

and the regularly stellate form is depressed. The

dorsal plates are hemispherical, not at all crowded

or tessellate, and bear a tuft of small spinules. The

plates are of all sizes, and some of the larger ones

bear a small central spine. A few thickened and

enlarged plates, raised slightly above the general

disc surface, surround the 3—7 papular pores in each

papularium. The superomarginals encroach less

on the dorsal surface than in P. mixtus, and they

are set at a less oblique angle on the arms. They

are covered with small, rough spinules and each

bears a central erect spine a little longer than the

width of the plate. The inferomarginal plates are

nearly square, alternate with the superomarginals,

and bear one (interradially sometimes two) spine

like that of the superomarginals. They are sparsely

covered with small, rough spinules. The single

series of rounded actinal interradial plates does not

extend beyond the first pair of inferomarginals;

they bear a few short spinules, not sharp. No

pedicellariae were observed.

The projecting adambulacral plates bear a circlet of spines and spinules, a few small spinules

on the actinal edge and 7 or 8 slender spines on the

furrow edge. There is one (rarely two) large spine

on the actinal face. The mouth plates are wide,

short, and rounded. Each jaw half bears one large

preoral spine and 6 or 7 long, slender, divergent

lateral spines. The actinal surface bears several

pairs of short, acute spines. The small, slightly

sunken madreporite is very deeply channeled and

touches or nearly touches the adjacent superomarginals.

Verrill reports this species as very common in

the West Indies, in 70-300 fathoms.

MATERIAL EXAMINED.—Oregon station unknown

(6) off the Florida Keys, 200 fathoms [R=13 mm,

r=S mm, Rr=l:4].

Order VALVATIDA Perrier, 1884

The mouth plates in this order are inconspicuous.

The inferomarginal and superomarginal plates correspond with one another, and are without intermarginal channels. The tube feet are in two rows.

Pedicellariae, when present, are valvate and sunken

into the ossicles.

Key to the Families of Valvatida

1. Disc broad, arms not cylindrical; actinal interradial area generally large; marginals large

and conspicuous 2

Disc small, arms usually long and cylindrical; marginals small, inconspicuous

Ophidiasteridae

2. Mouth plates with one or two conspicuous recurved hyaline spines Odontasteridae

No recurved spines on mouth plates 3

3. Body thick, cushion-like; interbrachial septa calcareous Oreasteridae

Body not thick and cushion-like; interbrachial septa membranous Goniasteridae

Family ODONTASTERIDAE Verrill, 1899

In this family, the form is pentagonal or broadly

stellate; the abactinal surface is more or less paxillate. The tube feet are suckered, in two rows. The

marginal plates correspond and are not conspicuously spiny; there is an odd interradial marginal

in both series. The mouth plates bear one or two

conspicuous recurved hyaline spines per plate.

There are no superambulacral ossicles.

Only one genus in this family is represented in

western North Atlantic.

Odontaster Verrill

Odontaster Verrill, 1880:402. [Type, by original designation,

O. hispidus Verrill.]

Gnathaster Sladen, 1889;285. [Type, by subsequent designa-
NUMBER 126 45

tion, Astrogonium meridonale Smith (Verrill, 1899; 204).]

Gnathodon Verrill, 1899;204. [Erroneous spelling of Gnathaster Sladen.]

Peridontaster Koehler, 1920:190. [Type, by original designation, Calliderma grayi Bell.]

Epidontaster Koehler, 1921b: 190-195. [Type, by original designation, E. pentagonalis Koehler.]

Gymnognathaster Doderlein, 1927:297. [Type, by original

designation, G. gaussae Doderlein.]

Disc broad, arms short; form generally stellate; a

single heavy recurved spine with hyaline tip at

apex of mouth plates; entire body densely covered

with short spines; surface, particularly on ventral

side, looks furry.

Verrill has described three species of Odontaster

from the western North Atlantic. One, O. robustus,

is limited to the vicinity of Martha's Vineyard, and

is anyway easily distinguished by its broad heavy

marginals and by having the odd interradial marginals of both series triangular and exsert (not

touching the intermarginal suture). It otherwise

resembles O. setosus. O. setosus and O. hispidus are

limited to the northern part of the range covered by

this report, in 40-400 fathoms.

Key to the Species of Odontaster

Papular pores over entire dorsum except for narrow interradial area on disc; spines of ventral

plates heavy, peglike O. hispidus

Papular pores in definite papularia; ventral spines fine, setose O. setosus

Odontaster hispidus Verrill

PLATE 15: FIGURES C, D

Odontaster hispidus Verrill, 1880:402; 1894:263; 1895:136;

1899:205, pi. 29: fig. 8, 8a; 1915:119, pi. 13: fig. 6.-Fisher,

1928a:489.-Gray, Downey, and Cerame-Vivas, 1968:148,

fig. 22.

The disc is broad and the five (not uncommonly

four or six) arms are short. The form is generally

stellate, but may vary from nearly pentagonal to

stellate with relatively long arms. The dorsal plates

have stellate, imbricating bases and a rounded,

elevated, subpaxillate top bearing a bundle of

6—25 spinules. Papular pores occur singly between

the plates except in a narrow interradial area and

near the extreme tip of the arm. The superomarginal plates are nearly square, confined to the dorsal

surface, and covered with dense spinules. The sides

of the plates are broadly bevelled and the sutures

between the plates are wide, deep, and bare. The

odd interradial superomarginal is wedge shaped.

The inferomarginal plates extend beyond the

superomarginals, defining the ambitus. They are

more rectangular than the superomarginals and

bear a dense coat of thicker, heavier spines.

The actinal interradial plates are round, compact, and in regular rows parallel to the groove;

one series extends nearly to the tip of the arm.

These plates each bear several heavy spines, larger

than those of the inferomarginals. The adambulacral plates are small, short, and crowded. They

bear 2 or 3 stout, heavy furrow spines; the actinal

face bears 2 or 3 rows of one or two similar spines.

AH the adambulacral spines are subequal and

thicker than those of the actinal interradial plates.

Contrary to Verrill's description, the mouth plates

are small, being no larger than the next adjoining

adambulacral plates (Verrill was probably misled

by the very dense spination). The two halves are

joined by a membranous suture and each bears

5 or 6 heavy spines like those of the adambulacrals.

There is an odd, central, recurved spine at the

apex. It is not much longer than the other spines,

but is very much heavier, and the tip is hyaline.

The madreporite, halfway between the disc center

and the margin, is of moderate size, hemispherical,

and covered with fine, deep gyri.

This species ranges from New England to Florida,

in 40-400 fathoms.

MATERIAL EXAMINED.—Oregon II Stations: 10728

(2) [R=35 mm, r=14 mm, Rr=l:2.5]; 10667 (1)

[R=37 mm, r=16 mm, Rr=l:2.5]; 10715 (1)

[R=32 mm, r=15 mm, Rr=l:2].

Odontaster setosus Verrill

PLATE 16: FIGURES A, B

Odontaster setosus Verrill, 1899:207, pi. 29: figs. 1-lc, 2; 1915:

120.

This five-armed stellate or pentagono-stellate species

is generally smaller than O. hispidus. The round-
46 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

topped dorsal plates bear 10-25 long, fine spinules.

The papular pores are in six well-defined areas, a

small circular area in the center of the disc, and

five large papularia extending from near the central area to the base of the arms. The dorsal plates

in these areas are larger and higher than the other

dorsal plates. The superomarginal plates encroach

quite far onto the dorsal surface but are not confined to it. They are rectangular, broader than long,

and the odd interradial marginal is not differentiated from the others. The edges of the plates are

not beveled and the sutures between them are

shallow. Several pairs of superomarginals near the

tip of the arm may be in contact dorsally. The

inferomarginal plates are similar and both series

are clothed densely with fine spinules. The large

actinal interradial areas are filled with rounded

plates in regular rows, at least one of which extends more than two thirds the length of the arm.

They are clothed with moderately long setose

spinules.

The adambulacral plates are square and bear a

furrow series of 5 or 4 stout, blunt, divergent spines;

the actinal edge of the plate bears 5 or 6 long,

acute spines, and between the two series are 3-5

spines similar to those of the furrow series. The

small mouth plates bear a cluster of small preoral

spines, about five larger lateral spines, two pairs of

stout epioral spines, and a very large, heavy, hyalinetipped, unpaired, recurved spine. The madreporite

is small, round, channeled, closer to the center of

the disc than the margin, and surrounded by a few

enlarged dorsal plates.

Verrill states that this species is often taken with

O. hispidus. I am inclined to the view that,

although their ranges overlap, essentially O. hispidus is a northern species, while O. setosus is more

southern.

MATERIAL EXAMINED.—Oregon Station 272 (1)

[R=28 mm, r=12 mm, Rr= 1:2.3]. Combat Station

185 (1) [R=23 mm. r=14 mm, Rr=l:1.9].

Family GONIASTERIDAE Forbes, 1841

The Goniasteridae have a large disc and large

marginals. The actinal and abactinal plates are

usually tessellate. Papulae are confined to the dorsal

surface, usually single, and in a circumscribed area.

The tube feet, in two rows, have large suckers.

There is no large recurved spine on the mouth

plates. The actinal interradial areas are extensive.

The interbrachial septa are membranous.

The Goniasteridae of the North Atlantic have

undergone a recent extensive revision by Dr. Jerald

A. Halpern1. The section here on this family is

mainly a paraphrase of parts of this very excellent

monograph, done with the kind permission and

assistance of Dr. Halpern.

Goniaster Agassiz

Goniaster L. Agassiz, 1835b:441. [Type, by original designation, Asterias tessellatus Lamarck.]

Phaneraster Perrier, 1894:388. [Type, by original designation, Pentagonaster semilunatus Linck.]

Only one species, Goniaster tessellatus (Lamarck),

is known from the western Atlantic.

Goniaster tessellatus (Lamarck)

PLATE 16: FIGURES C, D

Pentagonaster semilunatus Linck, 1733:21, pi. 23: no. 37, pi.

24: no. 39, pi. 27: no. 45.-Perrier, 1876a:24-28; 1876:64,

67-Rathbun, 1879:148-149.-Sladen, 1889:265, 266, 267-

268, 655, 694, 748-749.-Ludwig, 1899:539.

Artocreatis tertia species Seba, 1761:11, pi. 6: figs. 9, 10.

Asterias granularis [part] Gmelin, 1791:3164.

Asterias tessellata [var. C and D] Lamarck, 1816:552.—de

Blainville, 1834; pi. 23: fig. 4.

Goniaster tessellatus.-L. Agassiz, 1835a: 143; 1835b: 191.-H. L.

Clark, 1909:110; 1946:81.-Fisher, 19l0a:172; 1911a:167-

168-Verrill, 1914:286.-Halpern, 1970a:256-265, figs. 23-25.

Goniaster cuspidatus Gray, 1840:280; 1866:10.-Tortonese,

1937:31-34, pi. 2: fig. 6, pi. 6: figs. 25-31.-Madsen, 1950:

209-211.-John and Clark, 1954:139.-A. H. Clark, 1954:

375.-A. M. Clark, 1955:18, 22.-Engel et al., 1960; p. 12.

Astrogonium cuspidatum.—Muller and Troschel, 1842:56.—

Dujardin and Hupe, 1862:394.-Perrier, 1869:85, 1876b:

p. 67.

Goniaster semilunatus.-\on Martens, 1866:86.-Koehler, 1909b:

87, pi. 5: fig. 5.

Astrogonium dubium Perrier, 1869:85.

Goniaster americanus Verrill, 1871:130-131; 1899:151-156, pi.

24a: figs. 1-2, pi. 26: figs. 1-6; 1915:104-107, pi. 13:5, 5a.-

Perrier, 1875a:1273; 1876a:261; 1876b:67. - Bernasconi,

1958a:13-15, pi. 1: figs. 1, 2.-Cherbonnier, 1959:107-108-

Gray, Downey, and Cerame-Vivas, 1968:148-150, fig. 23.

Goniaster africanus Verrill, 1871:131-132; 1899: 156-157, pi.

25: figs. l-2.-Perrier, 1875a: 1273; 1876a:261; 1876b:67.

Astrogonium semilunatum.—Ytrxiex, 1885:37.

Phaneraster semilunatus—Perrier, 1894:388-390.

*PhD dissertation, 1970, University of Miami, published

in part.
NUMBER 126 47

Key to the Genera of Goniasteridae *

1. Large prominent spines on abactinal surface Goniaster

No spines on abactinal surface 2

2. Entire body covered by skin obscuring plates Anthenoides

Body not covered by skin S

3. Superomarginal plates in contact medially throughout length of arm; form never pentagonal 4

Superomarginals in contact medially for less than half the length of arm; form pentagonal

or not 5

4. Furrow margins not strongly angular; internal radiating connecting ossicles present

Rosaster

Furrow margins strongly angular; no internal connecting ossicles Nymphaster

5. Abactinal plates of arms larger than those of disc Circeaster

Abactinal plates of arms not larger than those of disc 6

6. Unpaired median spine common to each pair of mouth plates 7

Two median spines to each pair of mouth plates 8

7. Single row of plates separating superomarginals along arm Paragonaster

Several rows of plates separating superomarginals Pseudatchaster

8. Abactinal plates not completely covered by granules 9

Abactinal plates completely covered by granules 11

9. Central naked area on some actinal plates Tosia

Actinal plates completely covered by granules 10

10. Abactinals surrounded by single row of granules PlinthasUr

Abactinals with more than single row of granules Litonotaster

11. Adambulacral granulation very crowded Peltaster

Adambulacral granules spaced 12

12. Rr about 4; adambulacral furrow margin strongly angular TesseUaster

Rr less than 2; adambulacral furrow margin not strongly angular Ceramaster

• Key adapted from Halpern to acommodate only those genera included in these collections.

This species has a broad, convex disc and five

short, broad arms. The abactinal surface is covered

with large conical spines on the primary plates.

Secondary plates are present only in the papular

areas, which form broad bands from the center of

the disc down each arm. There are six papular

pores around each primary plate. Abactinal plates

are covered with flat granules and each has a border

of larger flattened granules. The marginals are

massive and bare, each bordered by a single row of

flattened granules. The superomarginals are larger

than the inferomarginals, and the center of the

plate is drawn up to form a large, blunt, conical

spine. The last two pairs of superomarginals are

contiguous medially. The inferomarginals are

tumid, smaller, and more numerous than the

superomarginals. A few granule-covered triangular

plates occur intermarginally. The actinal interradial areas are large; the actinal plates are large,

flat, and polygonal, covered with coarse, rounded

granules; a few plates near the mouth may also

bear a large conical spine. The straight margin of

the adambulacral plates bears a furrow series of

4-6 long, flattened, truncate, subequal spines.

There are three rows of subambulacral spines, of

2-4 large, blunt spines, those of the second row

thicker and heavier than those of the first and

third rows. The mouth plates are small, narrow,

and armed like the adambulacral plates. The

madreporite, nearer the center of the disc than the

margin, is moderate to large in size, flat, roundly

triangular, and covered with fine gyri.

Small sugar-tongs pedicellariae are numerous on

both surfaces, with broad-lipped alveoli and long

spoon-shaped valves. The oculars are small and

truncate-conical. This is quite a variable species in

size and distribution of spines, but surprisingly

uniform in shape. Small specimens have a plane,

rather than inflated, abactinal surface, and the

superomarginals are less tubercle-like. They may

also lack the spines on the abactinal primary plates.

The distribution in the western Atlantic is from

Cape Hatteras to Brazil; elsewhere, it is known from
48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the tropical eastern Atlantic and the Indo-West

Pacific. The depth range is 22—430 meters.

MATERIAL EXAMINED.—Oregon Stations: 4392 (1)

[R=79 mm, r=43 mm, Rr=l:1.85]; 4086 (6)

[R=71 mm, r=41 mm, Rr= 1:1.8]; 2245 (2)

[R=27 mm, r=15 mm, Rr=l:1.9]. Oregon II Stations: 10518 (1) [R=59 mm, r=31 mm, Rr=

1:1.9]; 10533 (1) [R=65 mm, r=29 mm, Rr=

1:1.8]. Bowers Station 43 (5) [R=45 mm, r=25

mm, Rr=l:1.9]. Silver Bay Stations: 41/43 (1)

[R=52 mm, r=27 mm, Rr=l:1.9]; 1710 (2)

[R=45 mm, r=25 mm, Rr= 1:1.8]; 2926 (1) [R

=21 mm, r=14 mm, Rr=s 1:1.6]; 2009 (1) [R=16

mm, r=9 mm, Rr= 1:1.8]; 3284 (1) [R=9 mm,

r=6 mm, Rr= 1:1,5]; 2390 (2) [R=33 mm, r=20

mm, Rr= 1:1.6]; 2010 (1) [R=5 mm, r=3 mm,

Rr=l:1.7]. Combat Stations: 339/340 (1) [R=25

mm, r=14 mm, Rr= 1:1.8]; 98 (1) [R=10 mm,

r=6 mm, Rr= 1:1.7].

Anthenoides Perrier

Anthenoides Perrier, 1881a:23. [Type, by original designation,

A. piercei Perrier.]

Leptogonaster Sladen, 1889:326. [Type, by original designation, L. cristatus Sladen.]

Antheniaster Verrill, 1899:173. [Type, by original designation,

Anthenoides sarissa Alcock.]

Entire body covered by skin of moderate thickness

which obscures plates below it; granules, when

present, beneath skin; numerous secondary abactinal plates, extending along arms; secondary plates

increasing in number with age.

Of the two species occurring within the area

covered by this report, only A. piercei is represented

in the present collection. A. brasiliensis, described

by Bernasconi (1956), is known only from a few

specimens from south of 07°N; Halpern (1970, unpublished) points out that it may be an infraspecific

geographic variety of A. piercei, from which it

differs principally in lacking secondary abactinal

plates, large central granules and bivalved pedicellariae on most actinal plates, and well-developed

inferomarginal spinules.

Key to the Species of Anthenoides

Two rows of secondary abactinal plates along each arm A. piercei

No secondary abactinal plates A. brasiliensis

Anthenoides piercei Perrier

PLATE 17: FIGURES A, B

Anthenoides piercei Perrier, 1881a:23; 1884:168, 170, 184,

247-248, pi. 8: fig. 1; 1894:38.-A. Agassiz, 1888; fig. 379.-

Sladen, 1889:326, pi. 756-Verrill, 1915:113, pi. 3: fig. 2, pi.

10: figs. 1-16, 2-2/.-Fisher, 1919:328, 331, 332.-H. L.

Clark, 1941:49.-John and Clark, 1954:139.-A. H. Clark,

1954:375.

The general form is stellate, with a large disc and

five short, narrow arms. The entire animal is

covered by a thick membrane, full of small scattered granules, which obscures the plates. The

plates are in regular rows, and small secondary

plates separate the carinal and adradial rows on

the disc and for a short distance out on the arm.

The large single papular pores extend in regular

rows from the disc center to two-thirds the length

of the arm. A broad, triangular interradial area on

the disc is without papulae; the plates in this area

are conspicuously larger than other abactinal plates

near the center of the disc. The disc dorsum is

decidedly inflated.

The superomarginal plates are of moderate size,

conspicuous, and square to wider than long, and

unarmed. The two series of marginals correspond,

more or less, and form together a vertical ambitus.

Eight to ten pairs of distal superomarginals are

medially contiguous. The lateral and actinal faces

of the inferomarginals form an abrupt angle, the

actinal face being quite plane and flat. They are

covered with granules and bear, on the angled

edge, one to several short flattened truncate spines

or a group of enlarged granules. The actinal intermediate areas are large. Actinal plates are large,

polygonal, and tumid; they are covered with

rounded granules, those in the center of some plates

being quite coarse. The adambulacral plates bear a

furrow series of 6—8 long, slender spines; there is a

naked area between these and the subambulacral
NUMBER 126 49

series, which consists of two large and one or two

small, thick, rather conical or truncate spines. The

mouth plates are of moderate size and are swollen;

the two halves are well separated. Each half bears

a large apical spine and 8—10 marginal spines

similar to those of the furrow series. Spines on the

actinal face are similar to the subambulacrals. The

madreporite is large, plane, and nearer the center

than the margin; it is covered with coarse radiating

gyri. Large bivalved pedicellariae occur on many

of the actinal plates, and also frequently on some

inferomarginals; smaller ones may occur on the

abactinal surface. The oculars are small and inconspicuous. The color in life is orange to deep

red abactinally, cream white actinally.

This species is known from Florida to Guyana,

in 20-844 fathoms.

MATERIAL EXAMINED.—Oregon II Stations: 10853

(2) [R=82 mm, r=33 mm, Rr=l:2.5]; 10184 (2)

[R=58 mm, r=24 mm, Rr= 1:2.5]. Oregon Stations: 3608 (1) [R=82 mm, r=32 mm, Rr = 1:2.5];

4421 (1) [R=51 mm, r=22 mm, Rr = 1:2.5];

station unknown (3) [R=83 mm, r=30 mm,

Rr=l:2.5]; 6699 (1) [R=49 mm, r=16 mm,

Rr=l:3 (abberrant specimen with unusually long

arms and very few secondary abactinal plate)];

station unknown (1) [R=23 mm, r=10 mm,

R r= 1:2.3]. 2027 (1) [R=27 mm, r=ll mm,

R r= 1:2.5]; 6403 (1) [R=21 mm, r=8 mm, Rr=

1:2.4]. Alaminos Station 20/65-A-9 (1) [R=57 mm,

r=27 mm, Rr=l:2]. Silver Bay Stations: 281 (1)

[R=51 mm, r=18 mm, Rr=l:3 (like the specimen

from Oregon Station 6699)]; 2453 (1) [R=22

mm. r=9 mm, Rr=l:2.5].

Ceramaster Verrill

Ceremaster Verrill [Section C of Tosia], 1899:161. [Type, by

subsequent designation, Asterias granularis Retzius (Fisher.

1906:1054).]

Philonaster Koehler [subgenus], 1909b:74. [Type, by monotypy, Pentagonaster (Philonaster) mortenseni Koehler.]

Abactinal plates completely covered by granules;

radial abactinal plates tabulate; peripheral granules

distinct from central; usually no secondary abactinal plates (when present, confined to disc); no

superambulacral ossicles; no internal radiating

ossicles.

Only one species is known from the tropical and

subtropical western Atlantic.

Ceramaster grenadensis (Perrier)

PLATE 17: FIGURES C, D

Pentagonaster grenadensis Perrier, 1881a:19-20; 1884:168, 181,

186, 232-233, pi. 8: fig. 2; 1894:39, 390.-Sladen, 1889:265,

266, 744.

Pentagonaster affinis Perrier, 1884:168, 183, 186, 243, pi. 8:

fig. 4; 1894:40, 39O.-Sladen, 1889:265, 267, 744.

Tosia affinis.-Terrier, 1884:183.

Pentagonaster gosselini Perrier, 1884:35; 1885b:886; 1891a:

1226; 1894:399-401, pi. 26: fig. 4; 1896:45.-Koehler, 1909b:

84, pi. 1: fig. 9.

Pentagonaster haesitans Perrier, 1885a:36; 1894:397-399, pi.

23: fig. 7, pi. 25: fig. 2.

Pentagonaster deplasi Perrier, 1885a:34; 1885b:886.

Pentagonaster crassus Perrier, 1885a:34; 1885b:886; 1891a:

1226; 1896: p. 45.

Astrogonium greeni Bell, 1889:433^34, pi. 19: fig. 4.

Pentagonaster balteatus Sladen, 1891:688-690, pi. 25.—Farran,

1913:9-10.

Pentagonaster hystricis Marenzeller, 1891:439, 442, 445; 1893:

4-5, pi. 1: figs. 2-2a, pi. 2: figs. 26-2c.

Pentagonaster greeni— Bell, 1892:74-75, text-fig.—Nichols,

1903:249.

Pentagonaster granularis var. deplasi.—Perrier, 1894:401-402,

pi. 26: fig. 3.

Pentagonaster kergroheni Koehler, 1895:453-454, fig. 2.

Tosia grenadensis.~\err\\\, 1899:162.

Pyrenaster affinis.-Verrill, 1899:168; 1915:222.

Ceramaster affinis.—Fisher, 1911:165.

Ceramaster grenadensis.—Verrill, 1915:222.-H. L. Clark, 1941:

38-39.-A. H. Clark, 1954:375.-Halpern, 1970a:213-218, figs.

8-9.

Ceramaster hystricis.—Koehler, 1924:176-178. — Tortonese,

1938:272.-Cherbonnier, 1956:4.-Dieuzeide, 1960: pp. 69-71,

fig. 5.

Ceramaster balteatus.—Mortemen, 1927:82-83, fig. 45.—Tortonese. 1965:156.

The form is stellate to pentagonal, with five short

arms. The abactinal plates are low-tabulate, and

secondary plates, if present, are restricted to the

center of the disc. The plates of the conspicuous

radial areas are slightly enlarged and are in 3-5

regular rows from near the disc center to the marginals. The papulae are single, between the plates

of the radial areas. All plates are covered with

rounded granules; the peripheral series on each

plate is flattened and rectangular. The superomarginal plates are rectangular, about twice as

wide as long, and 10-17 in number (usually 12).

They may be completely covered with granules,

but usually there is a more or less extensive naked

area. The inferomarginal plates correspond to the

superomarginals and are covered with granules.

The interradial actinal area is large and the flat,
50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

polygonal, actinal plates are in regular rows and

covered with granules coarser than those of the

abactinal surface. The adambulacral plates bear

5 or 6 subequal compressed furrow spines with

rounded tips. The armature is in 3 or 4 irregular

rows of large rounded granules. Each mouth plate

half bears a large median spine and about eight

spines like the adambulacral furrow spines; the

face of the half-plate bears three large, compressed

spines along the suture, and the rest of the plate

is covered by a few granules. The madreporite is

small and rhombic. The oculars are small, bare,

and cordiform.

Small excavate sugar-tongs pedicellariae are

usually present both actinally and abactinally.

According to Halpern (1970, unpublished), this

species is known in the western Atlantic from

Florida to Brazil, in 200 to 2,500 meters. It also

occurs in the eastern Atlantic from Ireland to the

Equator, in the Mediterranean, and in the Azores.

MATERIAL EXAMINED.—Oregon Station 6696 (5)

[R=21 mm, r=13 mm, Rr = 1:1.8]. Oregon II Stations: 10841 (1) [R=22 mm, r=14 mm, Rr=l:1.7];

10842 (1) [R=27 mm, r=16 mm, Rr=l:1.7]; 10844

(2) [R=33 mm, r=18 mm, Rr=l:1.8].

Peltaster Verrill

Peltaster Verrill, 1899:168-169. [Type, by original designation, P. hebes Verrill (=Goniaster nidarosiensis Storm).]

Form pentagonal to arcuate pentagonal; abactinal

plates flat, covered by granules; secondary plates

present throughout radial areas; adambulacral furrow spines stout, moderately short; subambulacral

series graduated in size, grading from furrow spines

to granules of actinal intermediate plates; adambulacral granulation crowded.

Key to the Species of Peltaster

Many actinal plates adjacent to adambulacrals bearing short bivalved pedicellariae

P. nidarosiensis

Pedicellariae, when present, only of small excavate sugar-tongs type P. placenta

Peltaster nidarosiensis (Storm)

PLATE 18: FIGURES A, B

Coniaster nidarosiensis Storm, 1881:90-91.

Pentagonaster vincenti Perrier, 1885c:34-35; 1885a:886; 1894:

396-397, pi. 26: figs. 2a-2b.

Pentagonaster nidarosiensis.-Storm, 1888:61, 1901:12.

Coniaster acutus Lutken, 1890:359.

Non Goniaster acutus.—Heller, 1863:419, pi. 1: figs. 1-4.

Pentagonaster placenta [part] Ludwig, 1897:157.

Peltaster hebes Verrill, 1899:169-170, pi. 28: fig. 4.-Fisher,

1911a:162.-H. L. Clark, 1941:41-42.-A. H. Clark, 1954:375.

Hippasteria phrygiana [part] Ludwig, 1900:457-458.

Peltaster nidarosiensis-Grieg, 1905:3-13, figs. 1-2, 4-5; 1927:

I31.-Fisher, 1911a:162.-Verrill, 1915:28, figs. 4, 4a.-Mortensen, 1924:20.-Tortonese and Clark, 1956:348, 350, 351,

figs. 1A, 2A.-Halpern, 1970a:235-2S8, fig. 14.

The form of this species is pentagonal and the disc

is very large and flat. Both primary and secondary

abactinal plates are flat, irregularly round, and

covered with moderately large, rounded or slightly

angular, crowded granules. The arrangement of the

primary plates is regular, except in the center of

the disc. There are six papular pores around each

primary plate over a large area of the disc; only

small triangular interradial areas lack pores. The

18—20 massive superomarginal plates are about

twice as wide as long. They are covered with granules similar to those of the abactinal plates. The

inferomarginal plates correspond to the superomarginals except distally, where they become

smaller and more numerous; they bear granules

like those of the superomarginals. The large, rhombic, regularly arranged plates of the broad actinal

intermediate areas are closely covered with granules

larger than those of the marginals. Most of the

actinal plates in the row adjoining the adambulacral plates bear a small bivalved pedicellaria;

similar pedicellariae occur on some of the other

actinals. The adambulacral plates bear, on a slightly

curved furrow margin, 4-6 short, thick, compressed

furrow spines with blunt tips; a first subambulacral

row of three shorter, thicker, prismatic spines, and

about three irregular rows of 3 or 4 smaller spines,

becoming reduced to granules, occupy the actinal

face of the adambulacral plates.

The moderate-sized mouth plates bear, on each

half, 7—10 strongly compressed marginal spines
NUMBER 126 51

similar to the adambulacral furrow spines, and five

shorter, thicker spines behind them. The madreporite is moderate to large, flat, covered with deep

gyri, and surrounded by a row of slightly enlarged

granules. The oculars are small, triangular, and

bare. Tiny excavate sugar-tongs pedicellariae occur

on many of the abactinal plates, and small bivalved

pedicellariae on the actinal surface as noted. The

anus is surrounded by five large, triangular or

teardrop-shaped plates and a group of enlarged

flattened granules.

In the western Atlantic, this species occurs from

Georgia to Cuba, and in the eastern Atlantic from

Norway to the Bay of Biscay and the Canary

Islands; the depth range is 95-1,107 meters. This

specimen represents a range extension for the

species, to Martinique Passage.

MATERIAL EXAMINED.—Oregon II Station 10827

(1) [R=75 mm, r=54 mm. Rr=l:1.2].

Peltaster placenta (Muller and Troschel)

PLATE 18: FIGURES C, D

Goniodiscus placenta Muller and Troschel, 1842:59.—Dujardin

and Hupe, 1862:402.

Goniodiscus placentaeformis Heller, 1863:419-420, pi. 1: figs.

1-2; 1868:54.-Gasco, 1876;40-41, fig. 11.

Goniodiscus acutus Heller, 1863:420-421, pi. 1: figs. 3-4; 1868:

54.

Goniaster placenta.—\on Marenzeller, 1875:361.—Stossich,

1883:189.

Goniaster acutus.—von Marenzeller, 1875:362.—Stossich, 1883:

189.

Pentagonaster mirabilis Perrier, 1876a:40-41; 1878:21, 84;

1894:390.-Ludwig, 1879:540.-Carus, 1885:88.-Sladen, 1889:

265, 746.

Pentagonaster placenta.-Perrier, 1878:21, 84; 1894:390.-Ludwig, 1879:540; 1897:157-179, pi. 5: figs. 1, 2, 10, pi. 7:

figs. 24-42.-Carus, 1885:87.-Colombo, 1888:68.-Sladen,

1889:265, 266, 748.-von Marenzeller, 1895:11, 23.-Koehler,

1895:454-155.-Grieg, 1905:4.-Mangold, 1909:134.-Lo Bianco, 1909:561.

Pentagonaster acutus— Perrier, 1878:21, 84.-Carus, 1885:87.

Pentagonaster minor Koehler, 1895:451-453, fig. 1; 1896: pp.

61-62, pi. 2: figs. 5-7.

Pentagonaster planus Verrill, 1895:135-136.

Tosia placenta—Verrill, 1899:161.

Peltaster planus.-Verrill, 1899:170-171, pi. 28: figs. 3, 3a -

H. L. Clark, 1941:42.—Gray, Downey, and Cerame-Vivas.

1968:151-152, [not fig. 26].

Ceramaster placenta .—Koehler, 1921a:42, fig. 31; 1924:174;

1929: 2 [unnumbered], 1 pi.—Mortensen, 1927:80. 81.—

Tortonese, 1937:60; 1952:187, fig. 3.-Kolosvary, 1937:447.

pi. 32: fig. 19.-Cuenot, 1948:238.-Wilson, 1955; fig. 3-Forest, 1955: pi. 40.-Cherbonnier, 1956:10; 1958:15.-Vevers.

1956:485.—Buchsbaum and Milne, 1960; pi. 130.-Strenger.

1963:446, pi. 164.

Peltaster placenta—John and Clark, 1954:149. — Halpern,

1970a: 238-244, figs. 15-16.

Sphaerodiscus placenta.-Tortonese and Clark, 1956:347-352,

figs. 1B, 2B.-Madsen, 1958:90-94, fig. 2-Tortonese, 1965:

157-159. figs. 72-74.

In this species, the form is pentagonal and flat.

The disc is large. The abactinal plates are arranged

in regular rows from center to margin, and secondary plates are limited to the center of the disc

and the papular areas. The abactinal plates are

flat, irregularly polygonal, and covered with

crowded, rounded granules; the peripheral granules are flattened, movable, and slightly larger than

the other granules. Six papular pores surround each

primary plate except in a narrow interradial area.

The large square superomarginals, 12—20 in number, may be completely covered with coarse,

crowded, hemispherical granules or may have a

more or less extensive naked area in the center,

bearing a few scattered granules. The inferomarginal plates correspond generally to the superomarginals and are similarly granulated; there are

usually a few more of them.

The actinal intermediate areas are very large,

covered with small, flat, polygonal plates in regular

rows, which in turn are covered with closely

crowded granules larger than those of the abactinal

plates. The adambulacral plates bear a furrow series

of 4-6 thick, compressed, blunt-tipped, subequal

spines. The three prismatic spines of the first subambulacral row are thicker and shorter than the

furrow spines, and there are three additional

irregular rows of 3 or 4 progressively smaller

granuliform spines. The mouth plates are of moderate size and bear two parallel rows of large,

compressed, thick spines, 7-9 on the margin and

five on the actinal face; the rest of the mouth plate

is covered with granules. The madreporite is large,

round, and not surrounded by specialized granules.

The oculars are small, cordiform, and naked. Small

excavate sugar-tongs pedicellariae, with rounded,

spoon-shaped valves, are generally abundant on

both surfaces; those on the actinal surface are larger

than those on the abactinal surface.

Tn the western Atlantic, this species occurs from

Martha's Vineyard to Key West and throughout

the Caribbean. It ranges from the Bay of Biscay
52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and the Mediterranean to the Equator in the

eastern Atlantic. Its depth range is from 10 to 695

meters.

MATERIAL EXAMINED.—Oregon II Station 10628

(2) [R=81 mm, r=59 mm, Rr= 1:1.5]. Oregon

Stations: 6699 (1) [R=75 mm, r=50 mm, Rr=

1:1.5]; 6696 (1) [R=75 mm, r=50 mm, Rr= 1:1.5].

This slightly abberrant specimen has been the

source of confusion; placed variously at one time

or another in four different genera in the Goniasteridae, it was only by a careful feature by

feature (almost plate by plate) study that its true

position was recognized. The arms are longer and

more produced than is normal for a Peltaster. The

flat tessellate abactinal plates are irregularly arranged, and secondary plates in the radial areas

are numerous. The superomarginal plates are

smaller than is usual in P. placenta, and they are

slightly concave. They are also more numerous,

and the inferomarginals correspond interbrachially,

but are almost two to one on the arms. Pedicellariae

are very abundant on abactinal, marginal, and

actinal plates, and a few occurring on the adambulacrals near the mouth are very large. Many of

the adambulacral spines, particularly those near

the mouth, are very thick and their tips are grooved

and pitted. Every actinal plate in the row adjacent

to the adambulacrals bears a pedicellaria.

PHnthaster Verrill

PHnthaster Verrill [Section B of Tosia], 1899:161. [Type, by

original designation, Pentagonaster perrieri Sladen.]

Pyrenaster Verrill, 1899:166. [Type, by original designation,

Pentagonaster dentatus Perrier.]

Only one species, PHnthaster dentatus (Perrier), is

known from the Atlantic.

PHnthaster dentatus (Perrier)

PLATE 19: FIGURES A, B

Pentagonaster dentatus Perrier, 1884:168, 170, 179, 180, 185,

242-243, pi. 3: fig. 8; 1894:36, 39, 4O.-Sladen, 1889: 265^

267, 744.-Farran, 1913:10-11.-Grieg, 1921:21-23.

Pentagonaster grandis Perrier, 1885c:35; 1885a:886; 1894:32.

Pentagonaster perrieri Sladen, 1889:265, 267, 746.-Perrier,

1894:31, 34, 391, pi. 25: figs. \a, lb.-Koehler, 1895:15;

1896:60-61, 124-125; 1909b:85-86, pi. 2: fig. 7; 1924:182.

Pentagonaster concinnus Sladen, 1891:690, pi. 26: figs. 1-5.

Tosia (PHnthaster) perrieri.-V trriW, 1899:161-Fisher, 1906-

1054.

Tosia (PHnthaster) compta Verrill, 1899:161, 163-165, pi. 27:

fig. 2.-Fisher, 1906:1054.

Tosia (PHnthaster) nitida Verrill, 1899:151, 163-165, pi. 27:

figs. 1, la, l&.-Fisher, 1906:1054.

Pyrenaster dentatus.—Verrill, 1899:167, pi. 27: figs. 3-36.

PHnthaster dentatus-Fisher, 1910a: 172; 1911a:165.-Verrill,

1915:107.-H. L. Clark, 1941:42-43.-A. H. Clark, 1954:375.

—Gray, Downey, and Cerame-Vivas, 1968:151.—Halpern,

1970a:244-252, figs. 17-19.

PHnthaster perrieri—Fisher, 1910a: 172; 1911a:165.-Koehler,

1921a:2.-Mortensen, 1927:83-84, figs. 46, 47.

Goniaster africanus Koehler, 1914:169-170, pi. 7: figs. 1-4.

This extremely variable species varies in shape from

stellate to arcuate-pentagonal, to simple pentagonal, to pentagonal with very short arms produced

at the angles of the pentagon. The disc is large and

flat. The abactinal plates are thin, flat, polygonal,

and naked save for a single row of small granules

around each plate, which vary in number. Secondary plates are few and confined to the disc center

and proximal radial areas. Papular pores are large;

six surround each plate over the entire dorsum

except in a narrow triangular interradial area.

The superomarginal plates vary in shape, size,

and number; the number, from 10-22; the shape,

from longer than wide to square, to wider than

long; and the size, from forming a flat, narrow

border around the ambitus to forming a thick,

raised border of tumid plates. Frequently the distal

pair of marginals is enlarged, being twice as long

as the next adjacent pair; 1-4 distal pairs are in

contact medially. Superomarginal plates may be

completely covered with spaced granules, but

usually there is a bare area along each side and

across the abactinal edge. Inferomarginal plates

correspond to superomarginals except distally,

where they are smaller and more numerous; they

are covered with spaced granules. The actinal plates

are large, flat, and rhombic; they are covered with

coarse, evenly spaced granules larger than those of

the marginals. The adambulacral plates bear a

furrow series of 5-8 close, slightly compressed,

blunt spines, and the actinal face of the plate is

covered with spaced granules larger than those of

the adjoining plates. The mouth plates are large,

long, and triangular; they are conspicuously outlined with angular enlarged granules; the spines

of the furrow margin are like the adambulacral

furrow spines, but thicker and subprismatic. The

median spines are large and triangular in cross

section.
NUMBER 126 53

The madreporite is of moderate size, slightly

tumid, and irregularly square. The oculars are

small, cordiform, and naked. Small excavate sugartongs pedicellariae may occur on either surface, or

both, or neither; they are usually not abundant, or,

if so, are confined to the abactinal surface.

In the western Atlantic, this species is abundantly

distributed from North Carolina to Brazil and in

the Gulf of Mexico. In the eastern Atlantic, it is

known from Ireland to Liberia and the Canary,

Cape Verde, and Azores Islands. The depth range

is 229-2,117 meters.

MATERIAL EXAMINED.—Oregon Stations: 3654 (1)

[R=51 mm, r=30 mm, Rr= 1:1.7]; 2775 (1) [R=51

mm. r=37.5 mm, Rr=l:1.4]; 2824 (1) [R=44 mm,

r=31 mm, Rr=l:1.4]. 5784 (1) [R=36 mm, r=25

mm, Rr= 1:1.4]; 3601, (1) [R=40 mm, r=25 mm,

Rr=l:1.6]. 2776 (2) [R=46 mm, r=30 mm,

Rr=rl:1.5]; 3561 (1) [R=58 mm, r=36 mm,

Rr= 1:1.5]; 1508 (1) [R=40 mm, r=20 mm, Rr=

1:2]; 3573 (1) [R=43 mm, r=25 mm, Rr=l:1.7];

5929 (2) [R=47 mm, r=30 mm, Rr=l:1.5]; 2771

(2) [R=57.5 mm, r=35 mm, Rr=l:1.4]; 4855 (2)

[R=37 m, r=25 mm, Rr=l:1.5]; 4416 (4) [R=25

mm. r=:16 mm, Rr=l:1.9]; 1507 (2) [R=40 mm,

r=21 mm, Rr=l:1.9]; 489 (4) [R=31 mm, r=25

mm, Rr=l:1.2]. Oregon II Stations: 10491 (1)

[R=43 mm, r=30 mm, Rr=l:1.4]; 10636 (1) [R=

27 mm, r=20 mm, Rr=l:1.3]; 10802 (1) [R=67

mm, r=43 mm, Rr= 1:1.5]; 10876 (1) [R=48 mm,

r=30 mm, Rr=l:1.5]; 10633 (1) [R=32 mm, r=25

mm, Rr= 1:1.2]; 10827 (1) [R=37 mm, r=24 mm,

Rr=l:1.5]. Alaminos Stations: 3C/68-A-7 (1) [R=

32 mm, r=18 mm, Rr=l:1.9]; 12B/68-A-7 (1)

[R=32.5 mm, r=20 mm, Rr=l:1.6]; 15H/68-A-7

(2) [R=42 mm, r=30 mm, Rr= 1:1.4]. Combat

Stations: 364 (2) [R=26 mm, r=20 mm, Rr=

1:1.4]; 450 (1) [R=30 mm, r=20 mm, Rr=l:1.5].

Combat station unknown (1) east of St. Augustine

[R=53.5 mm. r=40 mm. Rr=l:1.3].

Tessellaster Clark

Tessellaster H. L. Clark, 1941:36. [Type, by monotypy, T.

notabilis Clark.]

Tessellaster notabilis H. L. Clark

PLATE 19: FIGURES C, D

Tessellaster notabilis H. L. Clark, 1941:36, pi. 5: fig. 1.-

A. H. Clark, 1954:375.-Halpern, 1970a:218, figs. 10-11.

This stellate species has a large disc and five long,

narrow arms. The low-tabulate, mostly hexagonal

abactinal plates are in regular series parallel to

the carinals and are covered with rounded granules;

the peripheral granules on each plate are slightly

flattened and rectangular. The papular pores are

restricted to moderately small, oval, radial areas. Six

pores surround each plate. The marginals correspond except on the distal half of the arms, where

they alternate. The superomarginal plates are approximately square, and 15—17 distal pairs are in

contact medially. A single row of small, rounded

granules surrounds each plate except the distal

ones. Many of the mostly naked superomarginals

have a central, irregularly shaped indentation filled

with granules. The inferomarginal plates are similar, except that the granule-filled indentations are

larger and more numerous. The large, flat, rhombic

actinal plates are closely covered with granules

slightly larger than those of the marginals.

The furrow margin of the adambulacral plates

is strongly angular and bears 7 or 8 short, stout,

compressed but slightly divergent spines with

rounded tips. There is a wide, bare area between

the furrow spines and the subambulacral spines,

which are in 2 or 3 irregular rows of 4-6 large,

coarse granules. The mouth plates bear a marginal

series of 8 or 9 stout, compressed spines and a

larger, rounded, compressed median spine; the

actinal face of the plate bears three regular rows

of granules similar to the subambulacral spines of

the adambulacral plates. The madreporite is of

moderate size, irregular shape, and deeply honeycombed. The oculars are small, oval, and bare.

There are no pedicellariae.

This species is known only from Florida to Cuba,

in 329-575 meters.

MATERIAL EXAMINED.—Oregon Station 6699 (1)

[R=76 mm, r=21 mm, Rr=l:3.5].

Tosia Gray

Tosia Gray, 1840:11. [Type, by original designation, T. amtralis Gray.]

In this genus only one species, Tosia parva, occurs

in the Caribbean and Gulf of Mexico.
54 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Tosia parva (Perrier)

PLATE 20: FIGURES A, B

Pentagonaster parvus Perrier, 1881a: 19; 1884:36, 37, 231, pi.

7: figs. 7-8.-Sladen:265, 267, 746-747.

Goniaster americanus (part) Verrill, 1899:154-156, pi. 26:

fig. 6.

Plinthaster dentatus (part).—Gray, Downey, and CerameVivas, 1968: fig. 25.

Tosia parva.-Halpern, 1969b:503-506, fig. 1.

The form is pentagonal. The abactinal plates are

polygonal to round, slightly convex, and surrounded by a single row of large, flat, square granules which, in the radial areas, are frequently fused

to form a continuous ring around the plate. Five

or more of the abactinal primary plates are larger

than the rest of the abactinal plates. The center

of each abactinal plate bears 0—6 round granules

embedded in deep pits. Papulae are confined to an

oval radial area. There are 6—8 massive, tumid

supermarginal plates, each surrounded by a single

row of small, rounded granules. Single, small, round

granules or clusters of up to twelve granules are

scattered over the surface of the plates.

There are 8-10 inferomarginal plates corresponding to the superomarginals and similar to them,

but usually bearing more than one row of peripheral granules. Between the two series of marginals

are several rows of enlarged granules, those at the

inframarginal angles being particularly large. The

large, flat, rhombic actinal plates are arranged in

five chevrons and are surrounded by two or three

rows of coarse, round granules; many of the plates

have a more or less extensive central naked area.

The adambulacral plates have a straight furrow

margin bearing five subequal, compressed, blunttipped spines. The subambulacral spines are in 3 or

4 irregular rows of 3-5 short, granuliform spinules.

The mouth plate margin bears spines similar to the

adambulacral furrow spines but more compressed;

the median spine is not conspicuously enlarged.

The rest of the mouth plate bears a few angular

granules. The madreporite is of moderate size and

roundly triangular. The oculars are small and

naked. Pedicellariae are rare; when present, they

are small, with two slender valves embedded in

deep pits, and confined to the actinal surface.

This species ranges from North Carolina to

Trinidad, in 30-597 meters.

MATERIAL EXAMINED.—Silver Bay Station 2020

(1) [R—18 mm, r=12 mm, Rr=l:1.5]. Oregon

Stations: 5915 (1) [R=12 mm, r=8 mm, Rr =

1:1.5]; station unknown (6) [R=22 mm, r=15 mm,

R r= 1:1.5].

Rosaster Perrier

Rosaster Perrier, 1894:36. [Type, by monotypy, Pentagonaster

alexandri Perrier.]

Rosaster alexandri (Perrier)

PLATE 20: FIGURES C, D

Pentagonaster alexandri Perrier, 1881a:22-23; 1884:168-172,

177-178, 183, 238-242, pi. 6: figs. 3-8; 1894: pp. 37, 335.-

Sladen, 1889:256, 267, 296, 748.

Rosaster alexandri.-Perrier, 1894:36, 38-40, 387.—Verrill,

1899:197; 1915:111-113, pi. 11: figs. S-5b, pi. 7: fig. 2 -

Fisher, 1919:240-243, 247, 255.-H. L. Clark, 1941:41.-Madsen, 1951:89.-A. H. Clark, 1954:375.-Halpern, 1970a:208-

212, figs. 6-7.

The five arms of this stellate species are short,

narrow, and strongly tapered. The abactinal surface

is flat and the plates tabulate and irregularly round.

The bases of the plates are lobed and connected by

internal radiating ossicles. Each plate bears a

central group of two or three short, clavate spinules

and 12-20 divergent, peripheral spinules more

pointed and quite thorny. The radial papular

areas are oval and conspicuous; there are six large

pores around each plate.

The 18-20 inferomarginal and superomarginal

plates correspond except on the distal half of the

arm, where they may alternate. The superomarginals are wider than long and contiguous throughout the length of the arm, about seven pairs in

contact medially. They are covered with spinules

like those of the abactinal plates, and the spinule^

along the edges of the superomarginals are like

the peripheral spinules of the abactinals. The inferomarginals are like the superomarginals. The

actinal intermediate area is of moderate size, and

the plates are large and convex. They are covered

with regularly spaced conical spinules. The rectangular adambulacral plates bear 4 or 5 long,

thin, blunt furrow spines. The two or three

divergent spines of the first subambulacral row are

longer and stouter than the furrow spines, and

behind them are one or two irregular rows of

conical spinules. The mouth plates bear about six
NUMBER 126 55

marginal spines similar to the adambulacral furrow

spines, with the median spine large, compressed,

and rounded at the tip. There are about four pairs

of long, stout, moderately acute spines on the

actinal face. The madreporite is small and irregularly square. The oculars are moderately small,

cordiform, and naked. Small excavate sugar-tongs

pedicellariae may occur on actinal, abactinal,

and/or marginal plates, or may be entirely absent.

This species is distributed from the Straits of

Florida and the Gulf of Mexico throughout the

West Indies to Brazil. The known depth range is

60-443 meters.

MATERIAL EXAMINED.—Oregon II Station 10513

(1) [R=24 mm, r=12 mm, Rr=l:2]. Oregon Stations: 4398 (1) [R=15 mm, r=6 mm, Rr= 1:2.5];

station unknown (4) [R=27 mm, r=11.5 mm, Rr=

1:2.3].

Circeaster Koehler

Circeaster Koehler, 1909a:83. [Type, by subsequent designation, C. marcelli Koehler (Spencer and Wright, 1966).]

In this genus only one species, Circeaster americanus (A. H. Clark), occurs in the area covered by

this report.

Circeaster americanus (A. H. Clark)

PLATE 21: FIGURES A, B

Lydiaster americanus A. H. Clark, 1916:141-144; 1954: 367.

Circeaster occidentalis H. L. Clark, 1941:46-49, pi. 4: fig. 2.—

A. H. Clark, 1954:376.

Circeaster americanus.—Halpern, 1970a:265-270, figs. 26-28.

The form is stellate, with a large inflated disc and

five broad, tapering arms. The abactinal plates are

round or irregularly polygonal and surrounded by

a single row of flat, square granules. The centers of

the plates are mostly naked, but some, especially

near the center of the disc, may bear up to ten

round granules in deep pits. The abactinal plates

of the arms are conspicuously larger than those of

the disc. Secondary abactinals are present only in

the papular areas, which are extensive, papulae

being absent only from a narrow interradial area.

The papular pores are single, and six surround each

primary plate. The superomarginal plates, about

forty-four in number, are square interradially, and

twice as wide as long distally. They are surrounded

by a single row of granules and bear 15—25 scattered

granules on the surface. Three to fourteen superomarginals are in contact medially.

The inferomarginal plates correspond to and

resemble the superomarginals except that the surface granules are more numerous (40—50). The

large actinal intermediate areas extend about halfway down the arm. The plates are surrounded by

large prismatic granules, and the center of each

plate is occupied by a large bivalved or spatulate

pedicellaria. The furrow margin of the adambulacral plates is slightly angular and bears 4—6 short,

stout, prismatic, truncate spines, somewhat compressed; behind is a row of 3 or 4 smaller, similar

spines and behind them a large, bivalved pedicellaria; the rest of the plate is surrounded by granules

similar to those of the actinals. The mouth plates

are long, narrow, and bear a furrow series of six

spines, longer and stouter than the adambulacral

furrow spines. The median spine is longest, and

very compressed. A row of stout prismatic granules

occurs on either side of the wide median suture.

The madreporite is irregularly round to somewhat

stellate, about three times the size of the adjacent

abactinals. The oculars are large, oval, and may

bear a few truncate tubercles. Small bivalved or

spatulate pedicellariae occur on many of the abactinals and a few of the marginals; similar but larger

pedicellariae are abundant actinally.

This species is known from the Gulf of Mexico,

the Straits of Florida, Cuba, Colombia, Trinidad,

and Guyana, in 503-1,446 meters.

MATERIAL EXAMINED.—Oregon Stations: 6702 (3)

[R=76 mm, r=33 mm, Rr= 1:2.3]; 6696 (1) [R=77

mm, r=28 mm, Rr= 1:2.9]; 6697 (2) [R=62 mm,

r=26 mm, Rr= 1:2.9]; 4416 (1) [R=62 mm, r=25'

mm, Rr= 1:2.4]; 5930 (1) [R=53 mm, r=21 mm,

Rr=l:2]. Oregon II Stations: 10636 (2) [R=98

mm, r=34 mm, Rr= 1:2.9]; 10633 (1) [R=90 mm,

r=39 mm, Rr = 1:2.3].

Litonotaster Verrill

Litonotaster Verrill, 1899:171-172. [Type, by original designation, Pentagonaster intermedius Perrier.]

Abactinal plates flat, thin; not completely covered

by granules, but bearing more than single marginal
56 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

row; no secondary abactinal plates; papular areas

narrow; no superambulacral ossicles; no internal

radiating ossicles.

There are two species in this genus which occur

in the area covered by this report, but only one,

L. intermedius, has been taken in these collections.

Key to the Species of Litonotaster

Retinal plates covered by conical spines L. intermedius

\ctinal plates covered by hemispherical granules L. rotundigranulum

Litonotaster intermedius (Perrier)

PLATE 21: FIGURES C, D

Pentagonaster intermedius Perrier, 1884:243, pi. 5: figs. 5-6.

Non Pentagonaster intermedius— Alcock, 1893a:90.

Litonotaster intermedius.—Verrill, 1899:172, pi. 28: figs. 5-5b,

- H . L. Clark, 1920:85; 1941:43.-Madsen, 1951:88.-A. H.

Clark, 1954:375.-Halpern, 1969a: 130, figs. 1B, 2, 3.

The form of this species is stellate and the disc

is large, with a thin, flexible, inflated dorsal tegument. The five arms taper smoothly from the

rounded interbrachial arcs to narrow blunt tips.

The abactinal plates are small, flat, and irregularly

round; they are covered with small, closely crowded

granules, save for a bare space in the center of some

of the plates. The small papular areas are restricted

to a narrow band from the base of the arms to the

center of the disc. The superomarginal plates,

about twenty in number, are square to slightly

longer than wide, and one or two pairs may be in

contact medially. They are surrounded by a row

of granules similar to those of the abactinal plates,

and in the interbrachial arc they are flattened in

the vertical plane and the lower two-thirds of the

plate is covered with coarse granules. Distally, the

plates become more angular and the surface granulation is reduced or absent.

The inferomarginal plates correspond to the

superomarginals and are covered with short conical

spinules. The actinal intermediate area is large, and

the flat polygonal plates do not extend down the

arms. They are covered with short conical granules.

The large rectangular adambulacral plates have

a straight furrow margin bearing 6-8 short, blunt,

compressed spines; the rest of the plate bears small

conical spinules similar to those of the actinals.

The large mouth plates bear 12 or 13 furrow spines

similar to those of the adambulacrals, and the

median spine is longer and strongly compressed.

The rest of the plate is covered by conical spinules

parallel to the suture, which is broad and prominent. The madreporite is irregularly round and

slightly tumid. The oculars are small, cordiform,

and frequently bear a few truncate tubercles. Twoor three-valved short excavate pedicellariae occur

on some inferomarginals and actinals and on most

of the adambulacral plates.

This species is known from the Gulf of Mexico,

the Windward Passage between Cuba and Hispaniola, and from Guyana, in 1,958-3,294 meters.

MATERIAL EXAMINED.—Alaminos Stations: 10/68-

A-13 (1) [R=16 mm, r=7 mm, Rr=l:2]; 9/68-A-13

(4) [R=17 mm, r=7 mm, Rr= 1:2.4]; 4A/68-A-7

(5) [R=25 mm, r=11.5 mm, Rr=l:2]; 3C/68-A7 (1) [R=18 mm, r=9 mm, Rr=l:2].

Paragonaster Sladen

Paragonaster, Sladen, 1889:617. [Type, by subsequent designation, P. ctenipes Sladen (Fisher, 1919:228).]

Single row of large, flat (not paxilliform) abactinal

plates along most of arm; always reaching terminal

plate. Unpaired (but not recurved) median spine at

apex of each mouth plate pair.

Of the two species found in western Atlantic

tropical and subtropical waters, only P. subtilis

was taken in these collections. P. grandis, from

northern Cuba and Brazil (275-540 meters), is

mainly distinguished by having much shorter arms

and laige, crowded spines on the actinal plates.

Key to the Species of Paragonaster

Rr greater than 4; small spaced spines on actinals P. subtilis

Rr less than 3; large crowded spines on actinals P. grandis
NUMBER 126 57

Paragonaster subtilis (Perrier)

PLATE 22: FIGURES A, B

Goniopecten subtilis Perrier, 1881a:26; 1884:168, 183, 187,

253-254, pi. 5: figs. 3-4; 1885c:41; 1885a:884.-Sladen, 1889:

726.

Archaster formosus Verrill, 1884:383-384; 1885:519, 543.

Pentagonaster elongatus Perrier, 1885c:38.

Paragonaster cylindratus Sladen, 1889:311, 314-318, 655, 693,

713, 752, pi. 51: figs. 3, 4, pi. 53: figs. 3, 4.-Perrier, 1894:

357.-Verrill, 1894:257; 1899:196.

Paragonaster formosus.-Verrill, 1894:257; 1895:137; 1899:196.

Paragonaster elongatus.—Perrier, 1894:35, 357, 362-363, pi. 21:

fig. 3, pi. 24: fig. 4-Verrill, 1899:196.

Paragonaster strictus Perrier, 1894:35, 357, 363-365, pi. 25:

fig. 5.-Verrill, 1899:196.

Paragonaster subtilis.-Perrier, 1894:31, 35, 40, 357, 358-362,

pi. 23: fig. 5, pi. 24: fig. 3.-Verrill, 1899: p. 196-Koehler,

1909b:86-87, pi. 4: fig. 2.-Grieg, 1921:20-21.-Mortensen,

1927:79.-Macan, 1938:361.-Madsen, 1951: p. 89.

This stellate species has a relatively small disc and

five long, narrow arms. The abactinal plates are

paxillate, with a short, stout pedicel and a rounded

top bearing large, round, flattened granules centrally and a single row of slender spinelets peripherally. A single row of large, flat, rectangular

plates, similarly ornamented, extends out the arms.

Papular areas are large, covering most of the disc

except a small central and interradial area. There

are about eighty-four nearly square superomarginal

plates that are mainly in the horizontal plane. They

are covered by large, round, flattened granules and

are separated medially on the arms by a single row

of plates. The inferomarginal plates correspond to

the superomarginals and are covered with short,

spaced spinules and 6—10 longer lanceolate spines.

The actinal areas are small, not extending beyond

the third marginal. The plates are covered with

short, spaced spinules and some also bear one or

two longer lanceolate spines. The pectinate adambulacral plates bear a curved furrow series of 5—8

blunt, cylindrical spines, 1-3 lanceolate spines on

the face of the plate, and subambulacral spinules

like those of the actinal plates.

The mouth plates are convex and prominent;

they are covered with long lanceolate spines, thickly

clustered at the apex of the plate and more widely

spaced on the actinal face of the plate; the spines ai

the apex are quite thorny. The suture between the

plates is narrow. The madreporite is small, irregular in shape, and mostly or entirely hidden by the

adjoining abactinal plates. It is covered with very

coarse deep gyri. The oculars are large, long, oval,

and covered with spaced, round granules. There

are no pedicellariae. In all of the specimens I have

seen, the first pair of tube feet is greatly enlarged

and extended toward the mouth. The significance

of this feature escapes me.

In the western Atlantic, this species occurs in

the Gulf of Mexico and from about 37° to 41° N;

it is also known from the Azores, the Cape Verde

Islands, and the Bay of Biscay. Its bathymetric

range is 2,455—4,375 meters.

MATERIAL EXAMINED.—Alaminos Stations: 29/69-

A-13 (5) [R=56 mm, r=12 mm, Rr=l:4.9]; 4A/68-

A-7 (1) [R=65 mm, r=13 mm, Rr=l:5].

Nymphaster Sladen

Nymphaster Sladen, 1885:612. [Nomen nudum.]

Nymphaster Sladen, 1889:294-295. [Type, by subsequent designation, Nymphaster protentus Sladen, 1889 (^Pentagonaster arenatus Perrier, 1881a.—objective synonym). (Fisher,

1917:90).]

Arms long; R/r greater than three; superomarginals

contiguous medially throughout length of arm;

abactinal plates low-tabulate; adambulacral plates

with strongly angular 'furrow margins, becoming

apophyses distally; gonads in single tuft; no internal

radiating ossicles.

Two species in this genus occur in the tropical

western Atlantic; a third, N. ternalis (Perrier), has

been placed in the synonymy of N. arenatus by

Halpern (1970a).

Key to the Species of Nymphaster

Actinal plates covered by rounded granules; no spines on inferomarginal plates JV. arenatus

Actinal plates covered by short conical spinules; prominent spines on some inferomarginals

N. subspinosus
58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Nymphaster arenatus (Perrier)

PLATE 22: FIGURES C, D

Pentagonaster arenatus Perrier, 1881a:21; 1884:168, 176, 178,

179, 181, 182, 186, 236-238, pi. 7: figs. 3, 4; 1894:39.

Pentagonaster ternalis Perrier, 1881a:20; 1884:168, 180, 181,

186. 233-234. pi. 1: fig. l.-A. Agassiz, 1888: fig. S77.

Dorigona arenata.—Pemer, 1885c:39, 40; 1885a:884; 1894:31,

82. 33, 34, 38. 39, 40. 379-382, pi 21: fig. 5, pi. 22: fig. 6,

pi. 24: figs. 5, 6-Koehler, 1895:15; 1896:57-58, 124-125;

1909b:83, pi. 3: fig. 7.-Grieg, 1921:19-20, pi. 4: figs. 5-8.

Dorigona ternalis— Perrier, 1885c:39, 40; 1894:39, 371-375.

Non Dorigona ternalis.—Koehler, 1909a: 54-58, pi. 8: figs. 5, 6.

Nymphaster basilicus Sladen, 1889:295, 296, 306, 308-310,

655. 693, 709, 752, pi. 57: figs. 8, 9.-Verrill, 1899:186.

Nymphaster arenatus.-Sladen, 1889:752.-Verrill, 1899:186 —

Farran. 1913:ll-12.-Fisher. 1919:277.-Koehler, 1924:182.-

Mortensen, 1927:84-85, fig. 48.-H. L. Clark, 1941:39-40.-

A. H. Clark, 1954:375.-John and Clark, 1954:139.-Halpern, 1970a:223-228, fig. 28B.

Nymphaster ternalis.-Sladen, 1889:752.-Verrill, 1899:185, pi.

26: fig. 7.-Fisher, 1919:264, 266, 268, 276, 278, 282.-A. H.

Clark. 1954:375.

The general form is stellate, with a broad disc and

five long, sharply tapered arms. The abactinal surface is pentagonal, and the plates are low-tabulate

and irregularly round. They are covered by regularly spaced, rounded granules. The bases of the

radial plates are prominently lobed. Single papular

pores cover most of the abactinal surface except

for a small, triangular, interradial area. The superomarginal plates, in contact medially throughout the

length of the arm, number 24-64, depending on

size. They are two and one-half times as wide as

long interbrachially, and approximately square

distally. The abactinal and lateral surfaces are at

right angles to each other. They are covered by

rounded, regularly spaced granules. The inferomarginal plates correspond to the superomarginals

and are similarly granulated.

The large rhombic actinal plates, which do not

extend beyond the third or fourth marginal, are

covered with coarse, rounded, regularly spaced

granules. The adambulacral plates are strongly

angular (becoming apophyses beyond the actinal

areas) and bear a furrow series of 6-11 moderately

long, compressed spines with blunt tips. The subambulacral spines are in three or four irregular

rows of 4 or 5 large granules. The mouth plates

are long, narrow, and triangular; they bear 10 or 11

furrow spines slightly longer than the adambulacral

furrow spines. The median spine is longest and

most compressed. A row of tall, rounded spines

lines the median suture; the rest of the plate is

covered with scattered granules. The madreporite

is tumid and covered with coarse, radiating gyri.

The ocular plates are small, oval, and covered with

granules. Small excavate pedicellariae, when present, occur on some of the abactinal, inferomarginal,

and/or actinal plates, and two- three- or four-valved

spatulate pedicellariae occur on some adambulacrals.

This species is abundant from the northern Gulf

of Mexico to Brazil, in 275-3,000 meters. In the

eastern Atlantic, it occurs from Ireland to Morocco

and the Canary Islands.

MATERIAL EXAMINED.—Oregon II Stations: 10206

(11) [R=60 mm, r=20 mm, Rr=l:3]; 10624 (2)

[R=84 mm, r=22 mm, Rr=l:4]; 10514 (1) [R=84

mm, r=22 mm, Rr=l:4]; 10844 (1) [R=75 mm,

r=19 mm, Rr=l:4]; 10876 (1) [R=? (all arms

broken), r=20 mm,]; 10877 (5) [R=62 mm, r=16

mm, Rr=l:4]; 10872 (1) [R=70 mm, r=15 mm,

Rr= 1:4.7]; 10900 (2) [R=74 mm, r=20 mm,

Rr=l:3.8]; 10491 (2) [R=18 mm, r=5 mm,

Rr=l:3.5]; 10602 (2); 10794 (3); 10843 (1) [R=43

mm, r=12.5 mm, Rr=l:4]. Oregon Stations: 2820

(10) [R=75 mm, r=17 mm, Rr=l:4.5]; 4855 (3)

[R=57 mm, r=14 mm, Rr=l:4]; 4413 (2); 1946

(1); 1885 (1); 2774 (1); 1920 (2); 6703 (3) [R=53

mm, r=13 mm, Rr=l:4]; 2005 (1) [R=74 mm,

r=21 mm, Rr= 1:3.6 (specimen has cluster of enlarged granules on most interbrachial marginals)];

4416 (9) [R=35 mm, r=12.5 mm, Rr = 1:2.5]; 3654

(2) [R=66 mm, r=17 mm, Rr=l:4]; 2353 (7)

[R—86 mm, r=21 mm, Rr=l:4 (largest specimen

and few others have small spinules on some marginals)]; 3560 (3) [R=47 mm, r=13 mm, Rr=l:3];

1505 (4) [R=69 mm, r=18 mm, Rr=l:4]; station

unknown (2); 6696 (4) [R=41 mm, r=ll mm,

Rr=l:4] 2814 (5) [R=63 mm, r=16 mm, Rr

= 1:4]; 3574 (1) [R=90 mm, r=21 mm, Rr=l:4.5];

489 (1) [R=19 mm, r=6 mm, Rr=l:3]; 5784 (1).

Alaminos Stations: 87/69-A-ll (2) [R=67 mm,

r=12 mm, Rr=l:5]; 78/69-A-ll (2) [R=70 mm,

r=15 mm, Rr=l:4.5]; 11/68-A-13 (1) [R=59 mm,

r=14 mm, Rr=l:4]; 2H/67-A-5 (1) [R=80 mm,

r=15 mm, Rr=l:5]; 1/68-A-13 (1) [R=61 mm,

r=13 mm, Rr=l:4.5]; 15H/68-A-7 (1) [R=70

mm, r=20 mm, Rr = 1:3.5]; 13A/68-A-7 (1);

4E/68-A-7 (1); 15/65-A-9 (3) [R=20 mm, r=6
NUMBER 126 59

mm, Rr=l:3]; 24/68-A-13 (1) [R=19 mm, r=4

mm, Rr=l:5]. Combat Station: 450 (3) (small

spinules on a few of the superomarginals).

Nymphaster subspinosus (Perrier)

PLATE 23: FIGURES A, B

Pentagonaster subspinosus Perrier, 1881a:21; 1884:168, 176—

178, 185, 234-236, pi. 6: fig. 1.

Dorigona subspinosa.—Perrier, 1885c:39, 40; 1894:38, 39, 375-

379.

Nymphaster subspinosus.-Skiden, 1889:752.-Verrill, 1899:185.

- H . L. Clark, 1941:40.-A. H. Clark, 1954:375.-Halpern,

1970a:228-232, figs. 12-13.

This species is very similar to N. arenatus, except

that most of the inferomarginal plates bear 1—3

short, conical spines, some of the superomarginal

plates may bear a similar spine, and one or two

small conical spines may occur on some of the

actinal plates. I do not consider these differences

as being of a specific nature, especially as many

specimens of N. arenatus bear more or less conical

granules on the actinal plates, and enlarged granules or incipient spines on the marginals. Two

specimens collected by the Oregon off the Florida

Keys in 200 fathoms could easily be assigned, one

to N. arenatus and one to N. subspinosus—in fact,

there is no other way one could interpret these

specimens if both species are to be maintained.

MATERIAL EXAMINED.—Combat Station 450 (1) [1

(with 1 specimen of N. arenatus)]. Oregon, no station, off Florida Keys, 200 fms. [R=70 mm, r=22

mm, Rr=l:3].

Pseudarchaster Sladen

Pseudarchaster Sladen, 1885:617. [Nomen nudura.]

Pseudarchaster Sladen, 1889:109. [Type, by subsequent designation, Pseudarchaster discus Sladen (Verrill, 1899:191).]

Aphroditaster Sladen, 1889:116. [Type, by original designation, A. gracilis Sladen.]

Several rows of abactinal plates extending far along

arm; usually more than one row reaching terminal

plate. Unpaired (but not recurved) median spine

at apex of each mouth plate pair.

Of the two species in this genus known from the

western North Atlantic, only P. gracilis was taken

in these collections.

Key to the Species of Pseudarchaster

Actinal plates bear conical spines 3-5 times longer than wide P» gracilis

Actinal plates usually bear lanceolate spinet P- pareiU

Pseudarchaster gracilis (Sladen)

PLATE 23: FIGURES C, D

Aphroditaster gracilis Sladen, 1889:117-120, pi. 17: figs. 1-2,

pi. 18: figs. 7-8.-Verrill. 1899:195.-Fisher, 1919:227-228.

Pseudarchaster concinnus Verrill, 1894:250-255; 1895:132;

1899:193, pi. 30: figs. 3-3&.-H. L. Clark, l941:31-32.-Madsen. 1951:89.-A. H. Clark, 1954:375.

Astrogonium gracile— Perrier, 1894:342, 354.

Astrogonium aphrodite Perrier, 1894:342, 354, pi. 21: fig. 2,

pi. 23: fig. 2.

Astrogonium necator Perrier, 1894:350-352, pi. 23: fig. 1.—

Koehler, 1909b:74-75.

Pseudarchaster aphrodite.—Verrill, 1899:195.

Pseudarchaster necator.—Verrill, 1899:195.

Pseudarchaster ordinatus Verrill, 1899:194-195, pi. 30: figs.

4-4&.-A. H. Clark. 1954:375.

Astrogonium eminens Koehler, 1907:34-37: 1909b:68-71, pi.

16: figs. 3-6.

Astrogonium aequabile Koehler, 1907:37-40; 1909b:66-68, pi.

11: figs. 1-4.

Astrogonium marginatus Koehler, 1909b:71-73, pi. 14: figs.

1-4.

Pseudarchaster gracilis— Mortensen, 1927:86.—Macan, 1938:p.

355.

The general form of this species is stellate and

plane, with a broad disc and five arms of moderate

length which taper to an acute point. The abactinal plates are paxillose, and more than one row

extends nearly or quite to the terminal plate. The

paxillae are short, with rounded tops covered with

large, round, flattened granules and a peripheral

row of short spinules. Papulae occur singly between plates over most of the disc, except for a

narrow, triangular, interradial area. The superomarginal plates are two to four times as wide as

long, and mainly in the vertical plane, forming a

broad border to the abactinal surface. They are

densely covered with large, round, flattened granules. The inferomarginal plates correspond to the

superomarginals and are covered with short, sharp
60 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

spinules and longer conical spines. The actinal

interradial areas are covered with irregularly

shaped plates in regular rows from adambulacrals

to marginals; the plates are covered with short,

terete spinules and a few longer lanceolate spines.

The plates of the row next to the adambulacral

plates are larger and more regular than the others,

and between many of them a pectinate pedicellaria

occurs.

The adambulacral plates are strongly angular

and bear a furrow series of 6—9 cylindrical spines.

The subambulacral spines are similar but more

widely spaced, and there are 1—3 longer conical

spines in the center of each plate. The mouth

plates are prominent and tumid, with a wide suture

between the halves. They bear a furrow series of

eleven spines, and an unpaired, large, thick, median

spine occurs at the apex of each jaw. The rest of

the plate is covered with spaced spinules. The

madreporite is small, rhombic, and covered with

coarse gyri. The ocular plates are small, more or

less oval, and usually naked. Pedicellariae are as

noted above.

In the western Atlantic, this species is distributed

from Cape Cod to Surinam, and in the eastern

Atlantic it is known from the Azores and the West

Coast of Africa to the Equator. Its depth range is

1,090-2,174 meters.

MATERIAL EXAMINED.—Oregon II Stations: 10795

(1) [R=190 mm, r=38 mm, Rr=l:5-by far the

largest specimen of Pseudarchaster I have seen, being nearly four times the size of an average adult

specimen]; 10624 (3) [R=85 mm, r=25 mm, Rr

= 1:3]; 10602 (1) [R=33 mm, r=13 mm, Rr=l:3].

Oregon Stations: 5929 (1) [R=58 mm, r=17 mm,

R=l:4]. 6696 (2) [R=51 mm, r=15 mm, Rr=l:3

(abactinal gonopores, two to each interradial area

of disc near margin, plainly visible in these specimens)]; 4421 (1) [R=30 mm, r=ll mm, Rr=l:3];

2771 (3) [R=51 mm, r=17 mm, Rr=l:3]; 4300 (6)

[R=46 mm, r=18 mm, Rr=l:3 (gonopores on

these specimens plainly visible)]; 4416 (1); 6703 (3)

[R=50 mm, r=I4 mm, Rr=l:4]. Alaminos Stations: 15/68-A-13 (1) [R=63 mm, r=20 mm,

Rr=l:3]; 10A/68-A-7 (1) [R=43 mm, r=13 mm.

Rr= 1:3.8 (four-armed specimen)].

Family OREASTERIDAE Fisher, 1911a

The body is thick, cushion-like, and covered with

granulose skin. The papulae are numerous and in

definite areas. The abactinal plates are stellatereticulate, granulose, and usually bear large, conical

spines or tubercles. The interbrachial septa are

calcareous. The tube feet are in two rows, with

large suckered discs.

Only one genus and species in this family is

known from the Caribbean and Gulf of Mexico.

Oreaster Muller and Troschel

Oreaster Muller and Troschel, 1842. [Type, by original designation, Asterias reliculatus Linnaeus, 1758.]

Pentaceros Schultze, 1760 [non binom.].

Pentaceros Gray [n°n Cuvier and Valenciennes], 1840:276.

[Type, by original designation, Asterias reticulatus Linnaeus, 1758.]

Oreaster reticulatus (Linnaeus)

PLATE 24: FIGURES A, B

Asterias gigas Linnaeus, 1753:114, pi. 9: fig. 1.

Asterias reticulata Linnaeus, 1758:661.—Retzius, 1783; 1805;

pi. 14.-Lamarck, 1816:556.

Pentaceros reticulatus.—Gray, 1840:276; 1866:6.—A. Agassiz,

1877:108-112, pi. 16: figs. 6-lL-Perrier, 1878:21, 52, 83.

-Viguier, 1879:193, pi. 11: figs. 4-6, pi. 12: figs. 3,4.-Sladen,

1889:344, 762-Ives, 1891:339.-Nutting, 1895:52, 187, 202,

212,-Leipoldt, 1895:634.—(incl. P. lapidarius Grube).-

Sluiter, 1895:56.-H. L. Clark, 1898a:5, 6; 1901:237.-Ihering,

1898:155.-Conant, 1900.—Duerden, 1900:613, 620.—Tennent

and Keiller, 1914.

Oreaster gigas.-Verrill, 1867:278-279; 1868:367.-Lutken, 1859:

64-75.-Rathbun, 1879:149.

Oreaster reticulatus.—Mutter and Troschel, 1842:45, pi. 3:

fig. 2.-Field, 1893:84.-Doderlein and Hartmeyer, 1910:151—

152.-H. L. Clark, 1919:53-55, 71; 1933:22-23.-Boone, 1933:

80-82, pis. 41-42.-Doderlein, 1936:319-320, pi. 31: figs. 3-

3«.-Engel, 1939:3, 7.-A. H. Clark, 1939:442.-Caso, 1944:

248-253, 2 figs.; 1961:59-62, figs. 20-21.-Fontaine, 1953:

182, fig.-Breder, 1955: pi. 1: fig. 4.-Bernasconi, 1958b:135-

136. pi. 4: figs. 1-2; 1960:25.-Tommasi, 1958:16-17, 32-33,

pi. 3: fig. 2.-Madsen, 1959:163, fig. l.-Brito, 1960:5-6, pi.

1: fig. 3.-Stanek, 1960:49, figs.-Thomas, 1960:167-168.-A.

M. Clark, 1962: pi. 2: figs. n-d.-Ummels, 1963:73-81, pis.

3-6.—Gray, Downey, and Cerame-Vivas, 1968:146, figs.

I9a-b.

Oreaster lepidosus Grube, 1857.

Asterias sebae Blainville, 1834:240.

Oreaster aculeatus Gray, 1866.

Oreaster reticulata.-Duerden, 1896:285.

Oreaster reticulatus var. bermudensis H. L. Clark, 1944:372-

374, figs. 1-2.

This very massive starfish normally has five arms,

but may have from four to seven. Linck's De Stellis
NUMBER 126 61

Marinis (1733) has a woodcut of Linck in his laboratory examining a four-armed specimen; as he

based most of his classification on the number of

arms, he described this specimen as belonging to a

separate group. The disc is broad and high. The

arms are short to moderately long, becoming proportionately shorter with increased size. The primary dorsal plates are thick and irregularly shaped;

there is a conspicuous circle of plates in the center

of the disc and three irregular rows of plates on

each arm. The numerous other primary plates are

not. in any particular pattern. All the primary

plates are connected in a reticulate pattern by

small, irregular, or rod-shaped secondary plates.

Most primaries bear a large, heavy tubercle or

stout spine. Spaces between the reticulations are

papular areas, with numerous tiny papular pores.

The entire surface, dorsal and ventral, with the

exception of the spines themselves, is covered with

a moderately thick membrane closely set with small

granules. The large tumid superomarginal plates

define the ambitus and each bears a very stout,

blunt spine. The inferomarginal plates are confined to the ventral surface; they are large and

rounded and sometimes bear a short, stout, central

spine, but usually the center of the plate bears

instead a few enlarged granules. The actinal interradial areas are large, the plates (concealed by

granules somewhat larger than those of the dorsum) are tessellate, and many of the proximal ones

bear one or more short, stout, central tubercles or

spines. One series of actinolaterals continues nearly

to the end of the arm. Valvate pedicellariae, set

into small cup-shaped plates or granules, are numerous, especially proximally, and small pedicellariae

also occur on the dorsal surface. The small, square

adambulacral plates bear a furrow series of about

five short, flat, subequal spines and the actinal face

of the plate bears a large, flat lanceolate spine with,

usually, a small spine beside it. The mouth plates

are small and armed with heavy blunt spines. The

madreporite is of moderate size, flat, smooth, plane,

and covered with fine shallow gyri. The young

Oreaster reticulatus is not inflated, and the marginals are relatively larger and more conspicuous

than in the adult. The variation in shape, from

pentagonal to stellate, the differing degrees of inflation of the disc, and the variety of colors, from

dark green, brown, dark red, orange and red, and

yellow to light tan, make this an extremely variable

species.

The species is common in shallow-water grassand-sand flats from Florida to Brazil; it has also

been taken in the Cape Verde Islands, Bermuda,

and occasionally occurs as far north as Cape Hatteras.

MATERIAL EXAMINED.—Oregon Stations: 5390 (2)

[R=91 mm, r=40 mm, Rr=l:2.3 (height of disc

= 34 mm)]; 1937 (1) [R=71 mm, r=34 mm, Rr

= 1:2.2 (height of disc=34 mm)]; 1934 (1) [R=47

mm, r=22 mm, Rr=l:2 (height of disc=22 mm)];

1938 (1) [R=69 mm, r=30 mm, Rr= 1:2.3 (height

of disc=:29 mm)]; 3603 (1) [R=15 mm, r=7 mm,

Rr=l:2 (height of disc=5 mm)]; 5456 (1) [R=15

mm, r=8 mm, Rr=l:2 (height of disc=5 mm)].

Note that all of the above specimens are rather

small; this species commonly reaches a size of R =

200 mm. r=100 mm, and height of disc=80 mm.

Family OPHIDIASTERIDAE Verrill, 1867

In this family, the disc is small and the arms are

usually long and cylindrical. The actinal interradial areas are generally small, as are the marginal

plates. The skeleton is tessellate and covered with

a granular skin (smooth in Leiaster). The tube

feet are in two rows, with well-developed suckers.

The ambulacral furrow is narrow.

Among the best understood, taxonomically, of

the families of starfishes are the Ophidiasteridae.

The genera are clearly defined, and the characteristics used to separate species within the genera are

stable and valid. The present comfortable status

of this family is due largely to H. L. Clark (1921),

whose careful analysis of the group was done with

such simple clarity as to make his keys intelligible

and useful to all.

It is, therefore, surprising to find A. H. Clark in

1954 confusing Tamaria floridae (he called it Ophidiaster floridae), Ophidiaster alexandri, Ophidiaster

pinguis, and Hacelia superba, which he apparently

thought all one species, and synonymized under the

name Hacelia floridae! I have examined the types

of all of these species, and there is no doubt in my

mind that they are all valid and distinct.

H. L. Clark himself did not, apparently, examine

in any comparative way Linckia bouvieri and

Linckia nodosa; he considered L. nodosa a junior
62 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to the Genera of Ophidiasteridae

1. With internal connecting plates 2

Without internal connecting plates S

2. Covered with glassy spinules Chaetaster

Without glassy spinules Drachmaster

S. Abactinal plates in regular longi-series 4

Abactinal plates not in regular longi-series 7

4. Papular areas occurring below inferomarginals 5

No papular areas below inferomarginals Tamaria

5. Eight series of papular areas 6

Ten series of papular areas Hacelia

6. Alternating large and small adambulacrals Leiaster

Adambulacral plates all alike Ophidiaster

7. Adambulacral armature spiniform Narcissi*

Adambulacral armature granuliform Linckia

synonym of L. bouvieri. The two species are, of

course, quite distinct, as I pointed out in 1968.

Four new genera, Neoferdina Livingston (1931),

Celerina A. M. Clark (1967), Drachmaster Downey

(1970), and Copidaster A. H. Clark (1948), have

been added to the family since Clark's revision, and

the genus Chaetaster, long considered the type of

a monogeneric family, is here restored to the Ophidiasteridae (see discussion under that genus).

Although Leiaster has a smooth, thin tegument

completely lacking in granules or scales and Copidaster has a granulose or scaly covering, the affinities between these genera are unmistakable. In

addition to having the same number of rows of

plates, arranged exactly the same and of a similar

shape, both of these genera have a subambulacral

spine, not on every adambulacral plate, but on alternate plates. The adambulacral plates themselves are alternately broad and narrow. Copidaster

and Leiaster also share the same peculiar pedicellariae (few in Leiaster, abundant in Copidaster)

with very long, thin, straight valves in deep alveoli.

I do not believe Copidaster lymani can be maintained in a separate genus; it rightly belongs in

the genus Leiaster.

The character of alternating large and small

adambulacral plates in the genus Leiaster is, of

course, suggestive of the Zoroasteridae, as is the

shape and arrangement of the dorsal plates. These

characters hardly seem likely ones for parallel evolution as an adaptation to the environment, especially as the Ophidiasteridae are mainly restricted

to shallow water and the Zoroasteridae to deep

water.

Ocular plates, seldom mentioned in asteroid taxonomic literature, and of little importance in most

families, seem to be quite a reliable specific character in the Ophidiasteridae. Even with growth

changes, the basic shape, ornamentation, and relative size of the oculars is usually consistent.

Chaetaster Muller and Troschel

Chaetaster Muller and Troschel, 1840:321. [Type, by original

designation, Asterias subulata Lamarck, 1816 (—Astcrias

longipes Retzius, 1805).]

Disc small, arms five, long, rounded; all plates

above adambulacrals similar, in definite rows, marginals difficult to distinguish; paxilliform plates

raised in center, with expanded base connected to

adjoining plates by small internal radiating ossicles; plates bear numerous very sharp, slender,

glassy spinules; single rather large papular pores

confined to dorsal surface (none below marginals);

adambulacral plates bear furrow series of straight,

subequal spines parallel to groove; on actinal face,

a dense bundle of spines longer and larger than

those of adjoining plates; odd interradial marginal

plate frequently present; ampullae single; interradial septa calcified; pedicellariae unknown.

This is apparently a tropical-subtropical genus,

with a depth range of 30—1,000 fathoms.

This is undoubtedly one of the most puzzling of

the starfish genera; it has features suggesting at

least five other widely separated genera. The presence of internal connecting plates in a recently

described ophidiasterid {Drachmaster bullisi Dow-
NUMBER 126 63

ney, 1970) would seem to justify the restoration of

Chaetaster to the family Ophidiasteridae, from

which it was removed on the basis of this single

character by Sladen in 1889 and Ludwig in 1897.

The lack of a thick smooth or granulose tegument

and the presence of bundles of glassy spinules are

thus the only important features left separating

Chaetaster from the other Ophidiasteridae. I do

not believe these characters to be of familial significance. At the time I published my description

of Drachmaster bullisi, I failed to notice that the

internal connecting plates of this species invalidated the separation of the family Chaetasteridae

from the Ophidiasteridae.

Certain features of this genus also seem to point

to the Asterinidae. Although the Asterinidae commonly have a larger actinal interradial area, the

elliptical plates, their arrangement and ornamentation, the type of adambulacral armature, and the

small marginals seem to draw the Ophidiasteridae

and the Asterinidae closer together; certainly I do

not believe they can possibly belong in separate

orders, as they are now classified.

Chaetaster nodosus Perrier

PLATE 24: FIGURES C, D

Chaetaster nodosus Perrier, 1876b:250.—Sladen, 1889:398, 399,

778.-Verrill, 1915:116, pi. 8: 6gs. 1, 2, pi. 13: figs. 4, 4a.-

A. M. Clark, 1951b: 1260.

Chaetaster longipes Sladen [part], 1889:399.—Verrill [part],

1915:117.

This stellate species has a small disc and five long,

well-rounded arms. The well-spaced plates are in

regular rows; carinal, 4 or 5 adradials, superomarginal and inferomarginal. They are all similar (elliptical, center raised and flat-topped, base slightly

expanded, top covered with numerous small, sharp

glassy spinules) save for a few scattered plates here

and there on the arms which are larger, raised

above the general surface, and distinctly nodose.

The plates above the superomarginals are connected to one another by small internal plates and

large papular pores occur singly between each plate.

No pores occur below the superomarginals, and

superomarginal and inferomarginal plates are alike

and indistinguishable from the dorsal plates. A

single row of similar interradial actinal plates does

not extend much beyond the disc. The small adambulacral plates are nearly square and bear a furrow

series of six spines, the proximal one small and the

other five subequal and in a straight row parallel

to the groove; unlike all the other spines, the furrow series of spines are not hyaline. The actinal

face of the adambulacral plate bears a dense bundle of larger, stouter spines. The two halves of the

mouth plate are long, narrow, and well separated.

They are densely covered with long fine spines,

those at the proximal end longest and extending

across the mouth. The madreporite is small, flat,

and deeply channeled.

This species is found in 30-200 fathoms, in the

West Indies and the Gulf of Mexico; it is not common. A specimen was recently collected off Cape

Hatteras.

MATERIAL EXAMINED.—Oregon Stations: 4459 (2)

[R=60 mm, r=8 mm, Rr=l:7]; 1416 (1) [R=55

mm, r=7 mm, Rr=l:8]; station unknown (1) [R

=53 mm, r=6 mm, Rr=l:9].

Drachmaster Downey

Drachmaster Downey, 1970a:77. [Type, by monotypy, D.

bullisi Downey.]

Drachmaster bullisi Downey

PLATE 25: FIGURES A, B

Drachmaster bullisi Downey, 1970a:77, figs. 1-6.

The disc is small and rather flat, with a central

anal pore surrounded by a number of enlarged

spiniform granules. The madreporite is irregularly

triangular or elliptical, sunken below the general

disc surface, and covered with deep gyri. The arms

are long, cylindrical, not much tapered, and terminate bluntly. All plates are covered with moderately thick skin, closely beset with small squami

form or minutely spinous granules of uniform size.

There are eleven very regular longitudinal rows

of smooth, slightly tumid plates on the arms, connected internally in transverse series by small tumid

plates. There are eight longitudinal rows of conspicuous papular areas, with from one to three

pores per area. There are two rows of actinolateral

plates, the row next to the adambulacrals containing twice the number of plates as the row above.

The adambulacral armature consists of two short,
64 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

blunt, somewhat flattened furrow spines, connected

near their bases by skin, and a much longer, flat,

acute subambulacral spine. The spines of the margin of the mouth plates are similar to and continuous with the furrow spines; on the actinal face

of the mouth plates are two pairs of large spines,

each spine of the distalmost pair accompanied by a

small spinelet.

Small, typically ophidiasterid pedicellariae occur

either on each superomarginal or between the

adambulacrals and the inferomarginals; they also

occur on the abactinal surface. The oculars are

large and tumid, and the terminal portion bears

large, low, rounded tubercles.

MATERIAL EXAMINED.—Oregon Stations: 5970 (1

holotype) [Rr=33 mm, r=4.5 mm, Rr=l:6]; 5923

(1 paratype) [R=14 mm, r=4.5 mm, Rr=l:4].

Tamaria Gray

Tamaria Gray, 1840:283. [Type, by original designation, T.

fusca Gray.]

Body wall rigid; abactinal plates well developed;

in regular longitudinal series; test granulose; no

spines on abactinal surface of arms; madreporite

small, simple; papular areas in 4-6 series, none

below inferomarginals.

Key to the Species of Tamaria

1. Four series of papular areas; arms short, subpetaloid T. passiflora

Six series of papular areas; arms moderately long, tapering 2

2. Oculars small, concealed by granules; average 3 or 4 pores per papular area T. floridae

Oculars large, bare; average 7 or 8 pores per papular area T. halperni

Tamaria floridae (Perrier)

PLATE 25; FIGURES C, D

Ophidiaster floridae Perrier, 1881a:9; 1884:221, pi. 4: fig. 1.—

Verrill, 1915:90.

Tamaria floridae.-H. L. Clark,1921:91.-Downey, 1971b:44.

Hacelia floridae [part].-A. H. Clark, 1954:376.

Although no specimens of this species were taken

in the present collections, a brief description of

the holotype is included here.

There are six longi-series of papular areas, 1—7

pores per area. There are two equal adambulacral

furrow spines, and a single, nearly round, very flat

and appressed subambulacral spine, with a small

spinelet or granule on each side of the base. Spines

similar to the subambulacral spines occur on many

proximal interradial actinal plates. There seem to

be about three regular rows of actinal interradials.

The few pedicellariae, similar to those of T. passiflora, are confined to the actinal surface. The oculars are small and concealed by granules. The

madreporite is small to moderate in size, triangular,

flat, and inconspicuous. The carinal, adradial, superomarginal, and inferomarginal plates are equal

and similar; there are no supplementary plates.

The anus is inconspicuous, not much larger than

a papular pore, but with about three enlarged

granules surrounding it.

This small species has caused considerable confusion. Perrier gave the locality for the type as

"' . . . ramene par la drague de 123 pieds de profondeur dans le detroit de Floride Communique

par M. Alex. Agassiz avant les dragues du Blake."

Verrill (1915) suggested that Perrier meant 123

fathoms, not 123 feet. H. L. Clark (1921) assumed

that 123 fathoms was meant, and from this deduced

that the locality given was wrong, as none of the

Blake stations listed by Agassiz were at 123 fathoms.

He concluded that one of two Bibb stations off

Florida was meant. The label accompanying the

holotype gives no data but "Blake coll." Dr. William N. Eschmeyer (1965) corrected station data

errors for the fishes collected by the Blake, but was

then of the opinion that no errors had been made

in data for the invertebrates. Perrier did give some

depths in feet and some in fathoms, and I can see

no reason for the assumptions made by Verrill and

Clark. Similar confusion exists about the dimensions of the holotype; here Perrier did make a

mistake, but Verrill and Clark did not improve

matters. I carefully measured the holotype, and the

correct dimensions are given below.

Finally, A. H. Clark (1954) apparently did not

examine the type at all—he called it Hacelia flo-
NUMBER 126 65

ridae, and included in its synonymy three other

quite distinct species.

MATERIAL EXAMINED.—Holotype (MCZ No. 757),

Blake, Straits of Florida, 123 ft [R=31 mm, r=7.5

mm, Rrz=l:4 (arm base=9 mm, midarm=7.5 mm,

arm tip—3 mm)]. Albatross Station 2672, 31°31'N,

79°05'W, 227 fms, May 1886 (1) [R=25 mm, r=5.7

mm, Rr=l:4 (arm base:=7 mm, midarm=4.9 mm,

arm tip=2.1 mm)].

Tamaria halperni Downey

PLATE 26: FIGURES A, B

Tamaria halperni Downey, 1971b, p. 46, fig. 1.

The disc of this species is of moderate size, and

the five arms taper evenly to rounded points. There

is a conspicuous central anus surrounded by enlarged granules. The carinal, adradial, superomarginal and inferomarginal plates are slightly tumid,

equal, and similar; there are no secondary plates.

There are six series of papular areas, with 7-10

pores per area (fewer on small specimens). In most

specimens, a few of the distal marginals, which

are slightly enlarged, have a central naked area

devoid of granules; otherwise, all plates bear an

even coating of small granules. There are 4 or 5

rows of actinolateral plates, one row extending twothirds the length of the arm. The two thin, flat

furrow spines of the adambulacral plates are nearly

square, even, regular, and equal. The subambulacral spine is oval, longer than broad, flat, and

appressed. Small specimens frequently have a small

spinule or enlarged granule between the subambulacral spines. There are two subambulacral

spines on the first adambulacral plate. There is a

pair of similar spines on the actinal face of the

mouth plates, while the spines of the mouth margin

are like the furrow spines. The pedicellariae are

numerous, one on each adambulacral plate and

others scattered on all surfaces; their valves are

short and triangular, in deep, well-defined alveoli.

The madreporite is of moderate size, plane, with

radiating gyri. The oculars are large, smooth, oval,

and raised above the general surface.

This species is apparently widespread in the

Caribbean and would undoubtedly have been reported on before had it not been confused with

T. floridae. Unfortunately, when this species was

described, I failed to give the derivation of the

specific name; it is named for Dr. Jerald A. Halpern, to whom I am very grateful for his many

helpful comments during the preparation of this

study.

MATERIAL EXAMINED.—Oregon Stations: 4928 (1)

[R=22 mm, r=5 mm, Rr= 1:4.5]; 2643 (1) [R=26

mm, rz=7 mm, Rr=l:4]; station unknown (1) [R

=20 mm, r=6 mm, Rr=l:3.5]; 6715 (3) [R=28

mm, r=6 mm, Rr=l:4.5]. Oregon II Stations:

10513 (1) [R=36 mm, r=10 mm, Rr=l:3.5]; 10849

(1 holotype) [R=53 mm, r=12 mm, Rr=l:4];

10850 (1) [R=50 mm, r=ll mm, Rr=l;4.5]. Also,

1 (MCZ No. 3999, as Ophidiaster alexandri), Atlantis Station 3480, Cuba, off Bahia de Matanzas,

200 fms [R=52 mm. r=12 mm, Rr= 1:4.5].

Tamaria passiflora1 Downey

PLATE 26: FIGURES C, D

Tamaria passiflora Downey, 1971b:49, fig. 3.

This heavy, solid species has a small disc and five

short, stubby, massive arms, slightly constricted at

the base, high on the sides, and plane on the actinal surface. The central anus is tiny, inconspicuous, and not marked by specialized granules. The

carinal, superomarginal, and inferomarginal plates

are large and separated by two series of papular

areas (none below the inferomarginals). There

are 3-15 pores per papular area. Between the carinals and the superomarginals, and between the

proximal marginals, a few small irregular secondary

plates occur. The granules covering the surface of

the starfish are squamiform and larger than those

of T. floridae. The actinal surface is plane, with

numerous small irregular plates, not in regular

rows except for the slightly larger series next to the

adambulacral plates, which, although irregular in

size and shape, are more or less in series; they continue to the end of the arm and most bear a pedicellaria. The two short, broad adambulacral furrow

spines are thick, flattened, and granuliform; there

is a shorter, smaller spine behind them. The subambulacral spine is large, flat, leafshaped, and appressed; a similar spine occurs on a few of the

actinal interradial plates.

1

 Etymology (not given in original description): L. passiflora, passionflower, for the five rather petaloid arms.
66 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The jaws are concealed by large, flattened granules and leaflike spines; they are bordered by spines

similar to those of the adambulacral furrow series.

The sugar-tongs pedicellariae are confined to the

actinal surface; the valves are short and sunken in

deep alveoli with high, raised, thickened lips. The

madreporite is small, flat, and deeply channeled.

The oculars are minute.

MATERIALS EXAMINED.—Oregon II Station 10858

(1 holotype) [R=24 mm, r=8.5 mm. Rr= 1:2.8].

Linckia Nardo

Linckia Nardo, 1834:717. [Type, by original designation, L.

typus (—Asterias laevigatus Linnaeus, 1758).]

Acalia Gray, 1840:285. [Type, by original designation, Linckia

pulchella Gray.]

Disc small, arms long, cylindrical; papulae in irregularly disposed papular areas, confined to abactinal surface; abactinal plates not in regular

longi-series; no pedicellariae; granuliform adambulacral armature in two series.

There are three species known from the Atlantic,

one of which (L. guildingii) is circumtropical, except for the eastern Pacific.

Key to the Species of Linckia

1. Large tumid plates occurring randomly on disc and arms 2

No large tumid plates L. guildingii

2. Minute spines on some lateral plates; ocular without granules L. nodosa

No spines on lateral plates; oculars covered with granules L. bouvieri

Linckia guildingii Gray

PLATE 27: FIGURES A, B

Linckia guildingii Gray, 1840:285; 1866:14.-Perrier, 1875b:

4O8.-A. Agassiz, 1877:105, pi. 14: figs. l-6.-Rathbun, 1879:

148.-Sladen, 1889:410.-Ives, 1891:339.-Russo, 1894:163.-

Leipoldt, 1895:634.-H. L. Clark, 1898b:412; 1898a:6; 1899:

130; 1901:340; 1921:67.-Verrill, 1901:36; 1907:281, pi. 34:

fig. c; 1915:96, pi. 28: fig. 3; 1902:36.-Richters, 1912:121,

figs. l-42.-Livingstone, 1932:254.-Boone, r936:241.-Tortonese, 1937:395, pi. 7: fig. 77; 1937a:75, pi. 4: figs. 10-11.—

Domantay and Roxas, 1938:222. pi. 14: figs. 80-81.-H. L.

Clark, 1938:133; 1944:374.-Hayashi, 1939:284, fig. 4, pi. 5:

figs. 8-10, pi. 6: fig. 2.-Caso, 1941:155, figs. 1^5.-Ely. 1945:

18, pi. l.-Habe, 1952:80.-Anon., 1953:188, figs. l-5.-Fontaine, 1953:183. fig.-Tommasi, 1959:17, pi. 3: fig. 3.-Utinomi, 1959:108, pi. 54: fig. 6.-Brito, 1960:5, pi. 1: fig. 2.-

Caso, 1962:70.-Ummels, 1963:81, pi. 7.-Clark in Clark and

Davies, 1966:608, pi. 18.-Pope. 1967:311, 314, fig.-Roa,

1967:279.-Downey, 1968:41; 1970a:81.

Ophidiaster omithopus Muller and Troschel, 1842:31.—Lutken, !859:80.-Dudjardin and Hupe, 1862:361.

Linckia ornithoptn.-Verrill, 1867:344; 1868:367.-Lutken,

1859:80; 1871:266.

Scytaster stella Duchaissang, 1850:4.

Linckia padfica Gray, 1840:285.-Studer. 1884:24.-Sladen,

1885:36.-Russo, 1894:162.-Sluiter, 1895:60. - Whitelegge,

1897:157.-Kellogg, 1904:354.-Ludwig, 1905:158.

Ophidiaster ehrenbergi Muller and Troschel, 1842:31.

Linckia nicobarica Lutken, 1871:265.

Linckia ehrenbergi.-de Loriol, 1885:31, pi. 10: figs. l-7a.—

Sladen, 1885:31.-Leipoldt, 1895:649.-Sluiter, 1895:60.-Ludwig, 1899:542.-Bell, 1903:245.-Tortonese, 1937:74, pi. 9:

fig. 58.

This versatile starfish has a small disc and, normally, five long, cylindrical, blunt-tipped arms. The

arms readily autotomize, and 2-7 arms are not uncommon; a specimen in the National Museum of

Natural History has only one arm, with no trace

of a disc or regenerating arms at all. The arms

may also vary in length on a single specimen, due

to regeneration after injury or spontaneous autotomy. The plates of the dorsum are not arranged

in regular rows. The papular areas between the

plates increase in size and in number of pores per

area with growth; small specimens have small papular areas, with 1-5 pores per area, while in large

specimens, the areas are as large as the plates, with

up to forty pores per area. Superomarginal and

inferomarginal plates are of a uniform size, and

separated by a series of papular areas. Two regular

rows of actinal plates extend nearly to the end of

the arm; there are about twice as many actinal

plates per row as there are inferomarginals. The

entire surface is covered with a dense coating of

small, flat-topped granules. There are two adambulacral furrow spines, side by side, the proximal

one longer and broader than the distal one; behind

and between them, the single subambulacral spine
NUMBER 126 67

is heavier. These adambulacral spines are so close,

and so crowded by adjacent granules, that they are

difficult to distinguish. The mouth plates are similarly armed.

The madreporites are of moderate size, flat, and

covered with deep gyri; they may be multiple, but

I have seen specimens with none. Two is the usual

number for normal, five-armed adults. They are

usually interradial, as in other starfishes, but one

specimen in this collection has, in addition to the

normal interradial madreporite, another one halfway out on an arm. The oculars are usually large

and covered with granules; they may be compound

(made up of two or three plates).

This species is distributed throughout the tropical waters of the world, except for the eastern

Pacific. It is a shallow-water starfish, mostly reefdwelling.

MATERIAL EXAMINED.—Silver Bay Station 1972 (4)

[R=38 mm, r=5 mm. Rr=l:7.5].

Linckia nodosa Perrier

PLATE 27; FIGURES C, D

Linckia nodosa Perrier, 1875c:417; 1884:226.-Sladen, 1889:409,

786.-Fisher, 1906:1088.-Bell, 1884:509.-Caso, 1962:72, fig.

26.-Downey, 1968:42.

Linckia formosa Mortensen, 1933a:4SO, fig. 10b, pi. 22: figs

2-4.

Linckia bullisi Moore, 1960:414, fig. 1.

Non Linckia nodosa.—Verrill, 1915:93 [see L. bouvieri].

This is a much less variable species than L. guildingii; I have never seen specimens with other than

five arms, of approximately equal length. The disc

is small and the arms moderately long, slender,

cylindrical, tapering, and blunt tipped. The dorsal

plates are irregularly arranged, and many are enlarged and tumid. The papular areas are inconspicuous, none below the inferomarginals, a series

between the inferomarginals and the superomarginals, and about four irregular series on the abactinal surface. There are 2-8 pores per area. The

superomarginal and inferomarginal plates are in

regular, nearly equal rows, with, occasionally, a few

of the superomarginals enlarged and tumid. The

marginals may or may not bear a central small tubercle or enlarged granule. The actinal surface is

plane, with 2 or 3 rows of regular, rectangular

actinal plates, only one row extending to the end

of the arm.

All plates are covered with a fine, dense coat of

small granules, those of the actinal surface slightly

larger than those of the abactinal surface. The

adambulacral furrow spines are two, one broad,

blunt, and flat, and the other slender and less blunt;

behind is a single thick, heavy, granule-like subambulacral spine. The mouth plates are armed

with two slender and two broad, square-tipped, flattened spines at the apex, and behind them are two

broader, square, flat spines. The madreporite

(when present) is flat and inconspicuous, with gyri

few and deep. The oculars are prominent, raised,

rounded, and bare. There are no pedicellariae.

This species has undergone all kinds of taxonomic vicissitudes; Verrill (1915) confused it with

L. bouvieri, H. L. Clark (1921) synonymized it

with L. bouvieri, and Mortensen (1933a) and

Moore (1960) overlooked it entirely. Perrier

(1875c) based his description of this species on

"Deux individus de la collection Michelin. Localite inconnu." The Michelin collection was a

wide-ranging one, and contained many specimens

from the western Atlantic, as well as from other

parts of the world. It was acquired by the Paris

Museum in 1868. Perrier indicates that quite a few

specimens were without locality labels. Fortunately,

specimens identified as L. nodosa by Perrier himself are in the collection of the Museum of Comparative Zoology at Harvard.

This species ranges, apparently, from Florida to

Brazil and St. Helena, in 62-475 fathoms.

MATERIAL EXAMINED.—NO data, probably collected by the Oregon (1) [R=36 mm, r=7 mm,

Rr=l:5]. Silver Bay Station 2010 (2) [R=27 mm,

r =5 mm, Rr=l:5].

Linckia bouvieri Perrier

PLATE 28; FIGURES A. B

Linckia bouvieri Perrier, 1875c:414.—Studer, 1884:24; Koehler,

19O9b:632.-Madsen, 1951:214, pi. 15: fig. 2.-Downey, 1968:

42.-Gray, Downey, and Cerame-Vivas, 1968:148, fig. 21.

Linckia nodosa.-Verrill, 1915:93, pi. 13: figs. 2-2a, pi. 29:

figs, la, lb.

This species has a small disc and five long, thick,

and flabby cylindrical arms, slightly constricted at

the base and tapering to an acute terminus. Owing

to the size of the papular areas, the skeleton is

looser than in the two previous species, and the
68 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

animal has a flaccid appearance. It is the largest

of the western Atlantic Ophidiasteridae. The disc

dorsum has a large, tumid, hemispherical central

plate, with a ring of 5-7 similar plates around it;

these primary plates are separated by small tumid

and nontumid plates. The inconspicuous anus is

at the base of the centrodorsal plate, surrounded by

slightly enlarged granules. The abactinal surface

of the arms bears about five irregular rows of plates

(some enlarged like the primary plates of the disc)

connected by small, narrow ossicles.

The large papular areas between the marginals

and on the abactinal surface are separated by small,

narrow, elongate ossicles; there are 10—40 pores per

papular area. The superomarginal plates are similar to the inferomarginals—subtriangular, somewhat

imbricate, and connected to them by narrow slender

ossicles. There are 3—5 rows of crowded actinolateral plates, irregular except for the row next to

the adambulacrals. The actinal plates overlap from

inferomarginals to adambulacrals. The plates of

the row overlapping the adambulacral plates are

larger and regular. The entire surface is densely

covered with small granules; those on the marginals

and major actinal plates are noticeably larger. The

adambulacral furrow spines consist of one broad,

flat spine and one small, slender spine per plate,

with a broad, flat, circular subambulacral spine on

the actinal face. The mouth spines are similar.

The madreporite is large, raised, and covered with

very fine gyri. The oculars are of moderate size,

raised, and usually covered with granules (the granules may be worn off in large specimens). There

are no pedicellariae.

The known distribution of this species is North

Carolina to Florida, Yucatan, and the Cape Verde

Islands, in 35-300 fathoms.

MATERIAL EXAMINED.—Oregon Stations: 4938 (1)

[R=134 mm, r=14 mm, Rr=l:9]; 4939 (1) [R=

144 mm. r=16 mm. Rr=l:9].

Ophidiaster L. Agassiz

Ophidiaster L. Agassiz, 1835b: 191. [Type, by original designation, Asterias ophidiana Lamarck, 1816.]

Body wall rigid, abactinal plates well developed, in

regular longitudinal series; no spines on abactinals;

skin uniformly granulose; papular areas in eight

series, only one below inferomarginals in each side;

rays cylindrical; madreporite small, simple.

Of the known species of Ophidiaster reported

from the area covered here, only O. guildingii is

represented in this collection. O. alexandri is

known only from a few specimens, and its distribution is limited to the northernmost part of the area

under study (Florida and Georgia), in 200—276

fathoms. O. pinguis is known only from the type,

which was from off southwest Cuba, in 320 fathoms.

It is superficially very like Hacelia superba. A

fourth species, O. bayeri, from Florida, was described by A. H. Clark (1948) from a single immature specimen in very poor condition. I have

been unable to distinguish any characters on which

this species might have been based.

Key to the Species of Ophidiaster

1. No pedicellariae; no supplementary subambulacral spinelet O. guildingii

Pedicellariae present; small supplementary spinelet at base of at least some

subambulacral spines .'. 2

2. Base of arms distinctly swollen; five rows of actinal intermediate plates O. pinguis

Base of arms not swollen; three rows of actinal intermediate plates O. alexandri

Ophidiaster guildingii Gray

PLATE 28: FIGURES C, D

Ophidiaster guildingii Gray, 1840:284; 1866:13.-Lutken, 1871:

23O.-Perrier, 1875c:387; 1878:16, 47, 80, 96.-Sladen, 1889:

402, 782.-Doderlein and Hartmeyer, 1910:152.-Verrill, 1915:

90.-H. L. Clark, 1921:79; 1901:237; 1933:23, pi. 2; 1898a:6.

-Fisher, 1928:490.-Tortonese, 1937:79, pi. 8: fig. 48.-Fontaine, 1953:182.-Ummels, 1963:86, pi. 8.

Scytaster mulleri Duchassaing, 1850 [not seen].

Ophidiaster flaccidus Lutken, 1859:86.—Verrill, 1867:344.

This five-armed species, usually of small to moderate size, has a small disc and long, cylindrical

arms, slightly constricted at the base. The plates

are in regular rows—carinal, one adradial, superomarginal, inferomarginal, and one actinolateral, all

reaching the end of the arm. The papular areas
NUMBER 126 69

occur between each row; they are large and contain 5-12 pores each.

The plates are roughly triangular and imbricate.

The entire surface is covered with fine, even granulation. The adambulacral plates each bear one

large and one small, granuliform, flattened, furrow

spine and a larger flat, blunt, appressed, subambulacral spine, separated from the furrow series by

an area of granules. The mouth plates are armed

like the adambulacrals. The madreporite is small,

plane, and slightly sunken. The oculars are large,

rounded, raised, and bear a number of small tubercles. There are no pedicellariae.

This species is distributed throughout the West

Indies, in shallow water.

MATERIAL EXAMINED.—Oregon Station 5438 (1)

[R=33 mm, r=4 mm, Rr=l:8].

Narcissia Gray

Narcissia Gray, 1840:287. [Type, by original designation, N.

teneriflae Gray (—Asterias canariensis d'Orbigny, 1839).]

Only one species in this genus occurs in the western

Atlantic.

Narcissia trigonaria Sladen

Narcissia trigonaria Sladen, 1889:414, pi. 65: figs. 5-8.—Verrill,

1915:97.-H. L. Clark, 1921:58-Brito, 1962:3, pi. 1: fig. 9.-

Tommasi, 1966:244.—Gray, Downey, and Cerame-Vivas,

1968:147, fig. 20.

Narcissia trigonaria var. helenae Mortensen, 1933a:429, fig.

10a, pi. 20: figs. 4-6.

This distinctive species is well represented in the

collection by a good growth series showing the

progressive reduction in relative size of the disc,

and the development of the arms from flat and

plane in small specimens to high and triangular in

cross section, with a marked carinal ridge, in adult

specimens. In small specimens, the carinal plates

are large and in a regular row; they are irregular

and obscure in larger specimens. Papular pores

occur singly, none below the inferomarginals, and

none in a narrow interradial band on the disc. Both

marginal series are regular, even, and large, and are

mostly confined to the lateral arm surface.

A single regular row of actinal intermediate

plates extends to the end of the arm; in most specimens, each bears one or two small sugar-tongs pedicellariae between flat-sided granules. Similar

pedicellariae occur also on many of the marginal

plates and on most of the larger abactinal plates

as well. The two valves of the pedicellariae are

long, delicate, and slender, with 2 or 3 small terminal teeth. All plates are covered with flattened

polygonal granules, those of the actinal surface

coarser than those of the abactinal surface. The

adambulacral plates bear a furrow series of 4 or 5

thick, blunt, somewhat flattened spines; the five

subambulacral spines are thicker and broader, subprismatic, but shorter than the furrow spines; behind them is a row of 3-5 prismatic spinules or

enlarged granules. These three rows of spines are

on distinct "terraces" of the adambulacral plates.

The mouth plates bear a crowded marginal row of

huge, blunt, triangular spines, constricted at the

base; behind them is a similar shorter row of spines,

and behind them are three short, prismatic spines

on each jaw half. The madreporite is small and

inconspicuous, near the apex of one interradius.

The oculars are nearly round, rather large, and

bare (save for a few terminal tubercles) in the

adult, but covered with fine granules in small specimens.

This species is distributed from North Carolina

to Brazil, in 20-50 fathoms. Although most of the

specimens in this collection are small, the species

reaches a size of more than 200 mm (R).

MATERIAL EXAMINED.—Oregon II Station 10517

(4) [R=70 mm, r=8 mm, Rr=l:9 (height of

disc=16.5 mm)]. Oregon Stations: 5739 (1 juvenile) [R=24 mm, r=6 mm, Rr=l:4 (height of

disc=4 mm)]; 2086 (1) [R=69 mm, r=ll mm,

Rr=l:6.5 (height of disc=14 mm)]; 2250 (4)

[R=74 mm, r=9 mm, Rr=l:8 (height of disc=

16 mm)]; 5023 (2 juvenile) [R=20 mm, r=6 mm,

Rr=l:3 (height of disc=4 mm)]; 2248 (1) [R=51

mm, r=9 mm, Rr=l:6 (height of disc=12 mm)];

5700 (1) [R=88 mm, r=21 mm, Rr=l:4 (height

of disc=25 mm)].

Two other Ophidiasteridae, Leiaster lymani

(A. H. Clark, 1948) and Hacelia superba H. L.

Clark (1921), have been described from the area

covered by this report (Key Largo, Florida, and

Barbados, respectively), but were not taken in

these collections.
70 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Order SPINULOSIDA Perrier, 1884

This order is characterized by having prominent,

unkeeled mouth plates; there is no definite and

distinct marginal frame; pedicellariae are usually

absent, but if present, they consist merely of

grouped spines. The aboral skeleton may be reticulate, imbricate, or absent, and in many forms the

plates are pseudopaxillate; the principal plates are

often cruciform. Ambulacral plates are not crowded.

The tube feet, generally in two rows, have welldeveloped sucking discs.

The undefinable family Poraniidae, represented

in this collection by the genera Poraniella and

Marginaster, is not included in the key that follows.

Key to the Families of Spinulosida

1. Dorsal membrane above abactinal surface; no marginal or actinal plates; mouth plates broad,

plowshare shaped Pterasteridae

No supradorsal membrane; marginal and, usually, actinal plates present; mouth plates not

broad and plowshare shaped 2

2. Mouth plates prominent, unkeeled; marginal plates paxillate Solasteridae

Mouth plates small; marginal plates not paxillate 3

3. Marginal plates large, blocklike; actinal plates in regular transverse rows Ganeriidae

Marginal plates inconspicuous; actinal plates absent, or, if present, not in regular transverse

rows Echinasteridae

Family SOLASTERIDAE Perrier, 1884

The family Solasteridae is characterized by having

a large disc and five to many long arms; the aboral

skeleton is open and irregularly reticulate. Marginal paxillae and oral intermediate plates are

present. There are no conspicuous marginal plates,

and the mouth plates are prominent and unkeeled.

There are no pedicellariae, and the suckered tube

feet are in two rows. Bundles of spinelets usually

ornament the plates.

Two genera in this family have been reported

from the tropical-subtropical western Atlantic.

Key to the Genera of Solasteridae

Superomarginal plates as large as inferomarginals Lophaster

Superomarginal plates much smaller than inferomarginals Solaster

Solaster Forbes

Solaster Forbes, 1839:129. [Type, by original designation,

Asterias endeca Linnaeus, 1771.]

Endeca Gray, 1840:183. [Type, by original designation, Solaster endeca Forbes (=Asterias endeca Linnaeus; Asterias

aspera Muller).]

Polyaster Gray, 1840:183. [Type, by original designation, Solaster papposus Forbes (=Asterias papposus Linnaeus;

Asterias stellata Retzius).]

One series of marginals well developed, superomarginals always smaller than inferomarginals; abactinal pseudopaxillae with stellate or cruciform base;

actinal intermediate plates extending more or less

along arm.

Two species of Solaster are represented in this

collection.

Key to the Species of Solaster

Rr 2 or 3; dorsutn very inflated; pseudopaxillae low, minute; subambulacral spines 6-8

S. notophrynus

Rr 4; dorsum not much inflated; pseudopaxillae high, small; subambulacral spines 3 or 4

S. caribbaeus
NUMBER 126 71

Solaster notophrynus Downey

PLATE 29: FIGURES C, D

Solaster notophrynus Downey, 1971a:39, fig. 1.

This large, handsome species has a broad disc and

seven short, broadly based arms tapering to a small

point. The dorsum is inflated and moderately thin.

The papular pores are single and numerous, and

the papulae are unusually large. The abactinal

plates are Y-shaped or cruciform, with long imbricating lobes, irregular on the central disc and

midarms, but in regular, oblique-transverse rows

elsewhere. Small, low, well-spaced pseudopaxillae

bear 10-40 minute thorny spinules. Narrow interradial bands of fused plates have few or no papulae.

The superomarginal plates are indistinguishable

from the adjacent abactinal plates. The highcrescentic inferomarginal plates are confined to the

actinal surface; they are about three times as wide

as long, and bear numerous small spinules on the

crest. Bare spaces between the inferomarginals are

about twice the length of the plate. The wellspaced actinal plates are in irregular chevrons, one

row extending nearly three-quarters the length of

the arm. They may be small and round, or larger

and elongate, and bear 4-14 small spinules. The

curved adambulacral furrow margin bears 4 or 5

setose spines, and a transverse ridge on the actinal

face of the plate bears 6-8 longer, stouter, acute

spines.

The mouth plates are wide and prominent, with

8—10 long, tapering, webbed oral spines, the central

pair longest, and a pair of fans of 4 or 5 smaller

setose spines, one above the other, at the sides; the

face of the plate is covered with small setose

spinules, and the suture is wide and bare. The

madreporite is small and nearly hidden by enlarged

pseudopaxillae; it is located nearer the margin

than the center, at the top of a patch of fused interradial abactinal plates. Internally, an interradial

strut between the mouth and the body wall is a

single column of plates embedded in a strand of

tissue supporting the abactinal roof.

MATERIAL EXAMINED.—Oregon Station 5929 (1

holotype) [R=114 mm, r=49 mm, Rr = 1:2.3].

Solaster caribbaeus Verrill

PLATE 30: FIGURES A. B

Solaster caribbaeus Verrill, 1915:54, pi. 28: figs. 1-la.—Gray,

Downey, and Ceraxne-Vivas, 1968:152, fig. 27.

The disc is moderately broad, and the eight arms

are short, rounded, and tapering. The dorsum is

slightly inflated, and the cruciform dorsal plates

form a close reticulum. The pseudopaxillae are

about as high as the width of a plate and bear

6-10 upright spinules about as long as the pedicel.

Solid areas of fused plates, mostly without paxillae,

occur in the interradii. Papular pores are single

and occur over the entire surface. The superomarginal plates are indistinguishable from the adjacent

abactinal plates. The inferomarginal plates are

raised, broader than long, and bear 10—15 spinules.

They define the actinal ambitus. The actinal interradial areas are narrow and the plates extend far

out on the arm. They are spaced, not much raised

from the surface, and bear 1—6 short, sharp, stout

spines larger than the spinules of the abactinal

surface. The adambulacral plates bear, on a curved

furrow margin, four moderately long, subequal

spines, and on the actinal face of the plate, a transverse series of 3 or 4 longer, stouter, acute spines.

The large mouth plates have a rounded apical

margin bearing 6-8 long, stout spines, evenly

graded from two large apical spines to small spinules at the side.

Behind these, on the side of the plate, is a fan

of three small appressed spines. There are two

stout suboral spines. The rest of the plate is bare

or has a few small setose spinules. The madreporite

is small, round, raised, and surrounded by enlarged

paxillae. Internally, the interradial strut of the

supporting plates from mouth to body wall is a

solid triangular composite of plates hanging like

an apron nearly to the actinal floor.

This species is known from North Carolina to

the West Indies in about 35—355 fathoms.

MATERIAL EXAMINED.—Oregon II Station 10825

(1) [R=58 mm, r=15 mm, Rr=l:4].

Lophaster Verrill

Lophastcr Verrill, 1878:214. [Type, by original designation,

Solaster furcifer Duben and Korcn.]

Sarkaster Ludwig, 1905:185. [Type, by original designation,

5. validus Ludwig.]

Only one species is known from the tropical western

Atlantic.
72 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Lophaster verrilli A. H. Clark

PLATE 30: FIGURES C, D

Lophaster verrilli A. H. Clark, 1938:6.

Lophaster radians.-Verrill, 1915:51, pi. 5: fig. 2, pi. 7: fig. 3,

pi. 11: figs. 1-lc [non Lophaster radians Perrier, 1884].

This species is stellate, and the disc is moderately

broad. The five arms are of moderate length and

subacute. The abactinal surface is convex, regularly

and openly reticulate.. The dorsal plates are cruciform or stellate, with large, high, flat-topped

pseudopaxillae bearing numerous setose spinules.

Large papulae occur singly or in small groups over

the entire abactinal surface. The marginals are

equal and opposite or alternate, similar to but

much larger than the abactinal plates. The small

actinal intermediate areas have few plates, one row

extending nearly to the end of the arm. The actinal

plates are small, round, and bear a central group of

small spinules. The adambulacral plates are wider

than long and spaces between them are twice the

width of the plate. They bear a webbed furrow

series of 4-7 long, slender, subequal spines parallel

to the groove and, on the actinal face, an oblique

series of 2—5 shorter, stouter spines.

There is a bare space between the two series of

spines, and behind the oblique subambulacral

series nearly half the plate is bare. The mouth

plates are broad, roundly triangular, and prominent. A raised ridge borders the broad suture on

each side. There are 20-22 webbed spines around

the margin of each pair of mouth plates, the two

apical pairs being longer and stouter than the

others. A few small spinules occur on the rest of

the plate, mainly along the ridges. The small

madreporite is round and raised.

This species is known from Florida and the West

Indies, in 56-200 fathoms.

MATERIAL EXAMINED.—Oregon Station 4912 (1)

[R=38 mm, r=13 mm, Rr=l:3]. Combat Stations:

171 (1) [R=15 mm, r=6 mm, Rr=l:3]; 185 (1)

[R=20 mm, r=6 mm, Rr= 1:3.5].

Family PTERASTERIDAE Perrier, 1875c

The family Pterasteridae is characterized by having

a dorsal membrane distinct from the abactinal

surface and supported above it by bundles of spines

arising from the cruciform abactinal plates. A central osculum in this membrane opens into the

so-called nidamental chamber, the space between

the abactinal surface proper and the upper dorsal

membrane. Elongate lateral spines on the adambulacral plates serve the same function as that of

the actinal plates in other asteroid families. The

mouth plates are broad and plowshare shaped.

Calcareous parts in these starfishes are reduced

and usually fragile. The tissue is frequently flabby

or gelatinous. This is principally a deepwater

family, and its distribution is worldwide.

Key to the Genera of Pterasteridae

1. Adambulacral spines not webbed, not in transverse series Hymenaster

Adambulacral spines partially or completely webbed, in transverse or oblique series 2

2. Adambulacral furrow spines webbed to actinolateral spines; paxillar spines long, setose

Pteraster

Adambulacral furrow spines only partially webbed, not webbed to actinolateral spine;

paxillar spines short, robust Calyptraster

Hymenaster Thomson

Hymenaster Thomson, 1873:120. [Type, by original designation, H. pellucidus Thomson.]

Armature of adambulacral plates not in transverse

webbed combs; spinelets of paxillae not protruding

through membrane; membrane contains many

muscle fibers and few to many spiraculae.

At the time Sladen published his report on the

Challenger asteroids, 24 species of Hymenaster had

already been described, a remarkable advance in

the knowledge of a genus which was first described

from a single specimen discovered by Wyville

Thomson a mere 25 years before, especially for a

group confined (with the exception of a single

species) to the abyssal depths. Seven of these species

had been described from the Atlantic, and Verrill

subsequently added another species from off the

east coast of the United States. As H. L. Clark
NUMBER 126 73

(1920) pointed out, the species of Hymenaster are

difficult to determine; a deep-sea group, they usually

arrive at the surface in a damaged condition,

further complicated by the fact that they are soft

bodied and gelatinous, with hard parts few and

delicate. Clark quite rightly based the classification

of Hymenaster on hard-part morphology, as, indeed,

is the case for all asteroids. However, hard parts

in Hymenaster are usually deeply embedded in

thick flesh, and nearly impossible to examine critically without drying the specimen. Clark considered

the oral and suboral spines and the adambulacral

spines as the most important and dependable

features for separating species within the genus;

unfortunately, these spines are loosely attached and

frequently lost in collecting and preservation. Even

the surfaces to which they attach are often badly

abraded, due to the generally soft and delicate

calcite of all hard structures within the group. Another factor hampering the determination of species

is the rarity of specimens; most of the 49 species

presently included in the genus are based on single

specimens. Because of the difficulty of preserving

these animals, many of the types have been lost,

and the genus is in dire need of a thorough revision

on a worldwide basis.

Because of the paucity of material and the poor

state of preservation, the identifications of species of

Hymenaster herein should probably be considered

as only tentative.

Key to the Species of Hymenaster

1. One suboral spine on each jaw plate H. modestus

More than one suboral spine 2

2. Two suboral spines on each jaw plate; disc round to pentagonal H. rex

Three suboral spines on each jaw plate; disc substellate H. anomalus

Hymenaster rex Perrier

PLATE 31; FIGURES A, B

Hymenaster rex Perrier, 1894:186, pi. 13: fig. 2.

The form of this species is more round than pentagonal, and inflated in life. It is of the consistency

of a dense jellyfish; the hard parts are few and

delicate. The five radial areas are well defined,

and the broad interradial areas are without plates,

spines, or calcareous deposits. Over the ambulacral

areas, there are about 5—7 rows of delicate cruciform

plates with long, thin, flattened lobes and a moderately high columnar pedicel. On top of the

pedicel are 3 or 4 long, fine hyaline spines with

expanded bases. On the carinals, these spines are

about twice the length of the short pedicel, but

both pedicel and spines increase in length from

carinal to outer row. The lobes are in contact or

slightly overlap those of the adjacent plates, and

all abactinal plates are also in contact with the

ambulacral and/or adambulacral plates. The spines

support the dorsal membrane but do not protrude

through it. I could find no spiraculae. The central

osculum is large, and the webbed valves of 12—18

long, delicate glassy spines, as long as the longest

abactinal paxillar spines, have an expanded base,

each mounted on a small, triangular, flattened

plate which in turn is borne on the upper ridge

of the large, heavy, crescentic, oscular plate.

There are five of these plates forming a ring in

the center of the abactinal surface proper, inside

the nidamental chamber. The interradial madreporite, just outside this ring, is hemispherical and

covered with deep gyri. The central stomach contains a few large, round, flat, delicate sieve plates.

The gonads lie in clusters on either side of the

umbulacrals, separated by a membranous interbrachial septum containing a few large, flattened,

elongate or cruciform plates. The upper part of the

ambulacral plates is about three times as long as

wide and quite thin; they overlap. Below this

broad, winglike flange, the plate narrows to a strut,

connecting it with the flared outer or lower part

of the plate that rests on the inner adambulacral

plate. There are two sets of plates in the adambulacral position; the inner plate is by far the largest,

cruciform to nearly square, with the lobes quite

short except for the outer lobe (away from the

groove) on a few of the proximal plates, which may

be long, broad, and strong. The outer adambulacral plate rests on the actinal face of the inner

adambulacral and bears three short, acute, conical
74 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

spines on the furrow margin, and a small leaflike

opercular plate on the actinal margin covering the

small segmental pore. The inner adambulacral

plate bears a single, long, slender, lateral spine

which supports the actinal interbrachial membrane.

This spine has an expanded base and the tip

is thicker than the midsection of the spine. There

are 28-30 of these spines, the first 5 and the last

6 or 7 comparatively short; the rest are subequal.

The broad mouth plates are nearly straight along

the front, or apical, margin; each pair bears a stout

pair of apical spines and a similar pair of epioral

spines; they are thick, slightly flattened, and may

be bifurcate at the tip. The margin also bears, to

the side, 3 or 4 smaller acute spines. Interradially,

between the mouth plates and resting on them, is

a large round plate (odontophore) with an upwardextending flange. The ambulacral grooves are wide

and there are two rows of large tube feet. The

peristomial membrane is broad.

This specimen was badly damaged and poorly

preserved; it was necessary to dry it to prevent

further deterioration and to examine plates. During

drying, large fatty globules were observed over the

entire surface of the specimen.

While this specimen closely resembles others in

the genus Hymenaster, it differs from the description of that genus in several important respects;

there are no muscle fiber bands, no spiraculae, and

the two series of adambulacral plates have not,

to my knowledge, been observed in any other

asteroid. I suspect that this may be the same species

as that described by Perrier as Hymenaster rex; it

would be impossible for him to have seen the

double row of adambulacral plates without drying

the specimen. The muscle fiber bands he describes

may have been merely stretched strands of tissue.

Although no spiraculae are present on the specimen

before me, the very poor state of preservation may

account for this; the dorsal membrane at the center

of the disc had entirely disintegrated by the time

the specimen was received.

This species has previously been reported only

from the eastern Atlantic, in 400-800 fathoms.

MATERIAL EXAMINED.—Alaminos Station 14C/68-

A-7 [R=81 mm, r=73 mm, Rr=l:l.l].

Hymenaster modestus Verrill

PLATE 31: FIGURES C, D

Hymenaster modestus Verrill, 1885:151; 1894:277; 1895:203.

This substellate form has a broad disc and five

short, broad, petaloid arms. The dorsal membrane

is thin and fragile and contains a few minute granular deposits. The paxillar abactinal plates are

cruciform, with a moderately high pedicel that

bears 3 or 4 long, slender spines, about twice the

length of the pedicel. There are roughly five rows

of plates on each arm. The spiraculae are numerous

and small; larger holes in the membrane, which

might be mistaken for large spiraculae, are tears

in the dorsal membrane. These are present in every

specimen I have seen, including the type. The

osculum is large and has five valves of 6—14 long

spines, evenly graded from the longest in the center

of the valve to the shortest at the ends. These are

webbed together throughout their length, and the

valve is longer than the diameter of the osculum,

so there is considerable overlap when the osculum

is closed. The five ossicles supporting the valves are

the most massive ossicles present, crescentic, with a

short, sturdy pedicel broadly expanded at the top

into a tumid, crescentic knob. The actinal surface

is without deposits of any kind; the actinal tissue is

supported by actinolateral spines of moderate

length, and there is a naked interradial area.

The lateral fringe is wide and unsupported; in

contracted specimens, however, the actinolateral

spines may extend beyond the fringe. The ambulacral grooves are wide, with two rows of massive

tube feet. The adambulacral plates bear three

slender, subequal furrow spines. There are 15—25

actinolateral spines, of which the fifth to the seventh

are the longest. The mouth plates are longer than

wide, and each half bears a long, stout preoral

spine, a similar suboral spine, and 3 or 4 smaller,

more acute marginal spines, the outermost one of

which might be considered a second suboral, as

it sits slightly behind the line of the other 2 or 3

marginal spines. The aperture papillae are broad,

flat, leaf shaped, and bear longitudinal ridges. They

are borne on the adambulacral plates between the

furrow spines and the actinolateral spine.

I have examined the type specimen of this species,

and it is a juvenile, as indicated by the small size

and the presence of several long spines at the

terminus of the arms, in the position of the oculars,

a juvenile character in most asteroids. Nevertheless,

I have no hesitation in assigning the larger sped-
NUMBER 126 75

mens below to the species H. modestus, thus extending the range of this species, previously

reported as occurring from Nantucket to Cape

Hatteras, in 1,098-1,451 fathoms.

MATERIAL EXAMINED.—Oregon Stations: 2821 (2)

[R=30 mm, r=19mm, Rr= 1:1.1]; 2820 (3) [R=38

mm, r=23 mm, Rr= 1:1.3].

Hytnenaster anomalus Sladen

PLATE 32: FIGURES A, B

Hymenaster anomalus Sladen. 1882:228; 1889:512, pi. 89: figs

3, 4, pi. 91: figs. 4-6.

This substellate species has a broad plane disc

and five broad-based arms tapering to an acute

point. The dorsal membrane consists of a dense

network of thick fibrous muscle bands, forming a

firmer dorsum than is usual in Hymenaster. The

membrane is supported in regular rounded elevations over the whole abactinal surface; these

elevations are produced by the paxillar spines, of

which there are three on a short pedicel to each

H-shaped dorsal plate; the three spines are not

divergent, but are carried erect as a single unit.

The osculum is enclosed by a ring or coronet of

low papillose elevations, within which the reticulate

muscle bands continue partway up the five triangular valves. The valves consist of about 5 or 6 moderately long spines; because of the fleshy covering

over the spines, the exact number is difficult to

determine. The spiraculae are minute and occur

in the meshes of the reticulum. The actinal surface

is covered with fine muscle fibers. The ambulacral

grooves are wide and lanceolate, with two rows of

stout, straight, untapered tube feet. The adambulacral plates bear 3 or 4 glassy furrow spines, usually

webbed, either all together or in 2-2 or 2—3 groupings, and not in transverse series.

The aperture papillae are large, flat, and nearly

circular; Sladen described them as small and elongate-oval, but I believe this may be variable. The

actinolateral spines number about 13—17, with the

sixth the longest; there is a narrow interradial area

unsupported by spines, but the sixth spine on each

side touches the sixth spine of the adjacent arm at

the ambitus. The narrow, lateral fringe is scalloped

between the actinolateral spines which protrude

slightly through it. The mouth plates are small,

and each bears three large, stout suboral spines.

Because of the position in which this specimen is

preserved, it is impossible to see the marginal edge

without destroying the specimen; Sladen says this

species has two small marginal spines on the oral

edge of each plate.

With its relatively sturdy dorsal membrane and

its partially webbed adambulacral spines, this

species is almost as close to Pteraster as to Hymenaster; a new genus should probably be erected for

it when additional material becomes available. For

the time being, it is best left in the genus Hymenaster.

This species was known previously only from

Tristan da Cunha, in 1,425 fathoms.

MATERIAL EXAMINED.—Alaminos Station 4/69-A13. (1) [R=14 mm, r=8 mm, Rr=l:1.7].

Pteraster Muller and Troschel

Pteraster Muller and Troschel, 1842:128. [Type, by original

designation, Asterias militaris O. F. Muller.]

Pterasterides Verrill, 1909:547. [Type, by original designation,

Pteraster aporus Ludwig.]

Lophopteraster Verrill, 1895:203. [Type, by original designation, L. abyssorum Verrill.]

Hexaster Perrier, 1891b: 1227. [Type, by original designation,

H. obscurus Perrier.]

Temnaster Verrill, 1894 (as subgenus):275; 1905 (as genus):

202. [Type, by original designation, Pteraster (Temnaster)

hexactis Verrill (=:Hexaster obscurus Perrier).]

Muscle bands present in supradorsal membrane, but

not usually reticulate; adambulacral armature in

transverse webbed combs; actinolateral spines forming free, independent lateral fringe, not merged in

actinal floor; small spiraculae sometimes present.

Some confusion exists about the western Atlantic

tropical and subtropical species of Pteraster. Perrier's original description of P. caribbaeus (1881b)

is vague, and it is impossible to determine on which

specimen he based his description. In a later (1884),

more complete description, one of the two specimens from Nevis (Blake Station 151) evidently

formed the basis for the description. Perrier here

stated that one of the specimens was in bad shape,

and at least his measurements of major and minor

radii are based on the best of the two specimens. He

also had three smaller specimens, badly deteriorated, from two other stations, and a sixth specimen,

from Sand Key, Florida (Blake collection), which

differed in important respects from the good specimen from Nevis. Obviously, this latter specimen
76 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

must be considered the type of P. caribbaeus; Perrier did not designate a holotype, so the specimen

now in the collection of the Museum of Comparative Zoology at Harvard bearing catalog number

957 (the "good" specimen from Nevis) is hereby

designated the lectotype of Pteraster caribbaeus

Perrier. The two small specimens marked paratypes of P. caribbaeus in the Museum of Comparative Zoology (MCZ 958) from Frederikstadt (Blake

Station 130), along with the above-mentioned

specimen from Sand Key, correspond more nearly

to the type of P. militaroides H. L. Clark; it is unfortunate that Clark (1941) mentions, as the distinctive feature of P. caribbaeus, the heavy, white

epioral spines. He apparently examined the Sand

Key specimen, rather than the Nevis specimen.

This makes all the more peculiar the fact that he

described a new species in the same paper, P. militaroides, which is in fact identical with Perrier's

Sand Key specimen of "P. caribbaeus." His specimen of P. caribbaeus, from Station 2978C, is the

same species as his specimens from Cuba which he

called P. militaroides. To compound the confusion,

Verrill (1915) redescribed P. caribbaeus on the

basis of a specimen from Albatross Station 2667,

which is in fact the species later described by Clark

as P. militaroides. Clark's description of this species

is wholly inadequate, and it is only by comparison

of type material that the two species can be distinguished. And contrary to Clark's statement, there

is a much greater contrast between P. militaris and

P. militaroides than between P. militaris and P.

caribbaeus. Heavy white or hyaline suboral spines

of very dense, solid calcite are not present on the

lectotype of P. caribbaeus (which has large, rather

blunt, and sometimes slightly curved suborals of a

echinoid-like lattice-structured calcite), but are

present on the holotype of P. militaroides. Other

differences are discussed in the descriptions of the

two species.

Key to the Species of Pteraster

1. Suboral spines present 2

Suboral spines absent P. personatus

2. Suboral spines hyaline 3

Suboral spines not hyaline P. caribbaeus

3. Broad lateral fringe present P. rugosus

Lateral fringe narrow or absent 4

4. Supradorsal membrane with numerous calcareous deposits 5

Supradorsal membrane without deposits, spongoid, transparent P. acicula

5. Spiraculae small, numerous P. militaroides

Spiraculae large, not numerous Pteraster species

Pteraster personatus Sladen

PLATE 32: FIGURES C, D

Pteraster personatus Sladen, 1891:694, pi. 27: figs. 1-5.—Bell,

1893:94.-Koehler, 1896a:447; 1896b:49; 1907:24.-Nichols,

190S:251.-H. L. Clark, 1908:285 [in key].-Fisher, 1911a:369

[in key].-Farran, 1913:23.

Pteraster reductus Koehler, 1907:25; 1909b:96, pi. 3: figs. 8, 9,

pi. 20: fig. lO.-Grieg, 1932:28, pi. 5: figs. 6, 7.

This substellate species has a broad disc and five

broad-based arms which taper to an acute point.

The dorsal membrane is fine and lacy, with many

small spiraculae. The abactinal plates are cruciform, with the lateral extensions flattened and

broadly truncate; they are not as long as the tall

pedicel, which is expanded at the top, like a tall,

attenuate, inverted cone. The pedicels bear 10—25

fine, hairlike, glassy spines which are not quite

as long as the pedicel. These spines cross over and

mingle with the spines of adjacent plates, so the

whole abactinal surface appears to be covered with

dense, matted fur.

The valves of the moderate-sized osculum are

supported on five large fan-shaped pedicels and

consist of a great number of fine long spines. There

is a narrow but distinct lateral fringe visible from

the abactinal side. The actinal membrane is supported by 18-25 long actinolateral spines which

meet in midinterradius. There is frequently a

smaller, finer supplementary spine embedded in the

membrane beside the actinolateral spine. The

adambulacral furrow spines are 3—7 (usually five),

webbed together in a curved series; these spines are
NUMBER 126 77

long, acute, and subequal. The high, plowshareshaped mouth plates bear a marginal series of 5 or

6 long webbed spines similar to but longer than

the adambulacral furrow spines; there are no suboral spines, the face of the mouth plate being quite

bare.

Sladen and others have reported this species off

the coast of Ireland; I am convinced, although I

have not examined his types, that Koehler's Pteraster reductus, from the Azores, is a juvenile of

this species; he based his species on the smaller

size and longer spines (correlative characters in

asteroids), and on the presence of six, instead of

five, adambulacral spines. As noted above, the

number of adambulacral spines is variable.

MATERIAL EXAMINED.—Alaminos Station 87/69-

A-ll (1) [R=70 mm, r=28 mm, Rr=l:2.5].

Pteraster caribbaeus Perrier

PLATE 33: FIGURES A, B

Pteraster militaris A. Agassiz [non Muller], 1877.

Pteraster caribbaeus Perrier, 1881a: 13; 1884 [part]:216.

Non Pteraster caribbaeus-Verrill, 1915:82, pi. 7: fig. 4.—

H. L. Clark, 1941:60.

This form is broadly stellate, with a broad disc and

five acutely pointed arms. The dorsal surface is

moderately inflated. The dorsal membrane is crisscrossed with tough transparent irregular muscle

bands which connect the underlying paxillar spines.

There are a few calcareous deposits in the membrane, and spiraculae are small to moderate in size,

and numerous. The tall slender pedicels of the

abactinal plates arise from flattened, broadly lobed

bases and support up to a dozen long, fine, glassy

spines which protrude a short distance through the

membrane. The central osculum is surrounded by

five ill-defined valves of 10-15 long, subequal, blunt

spines, all webbed together and forming a continuous ring of closely set spines. The actinal surface is relatively small; the membrane is without

deposits and is supported by 12-16 long, sturdy,

ridged actinolateral spines, the sixth or seventh

being the longest and meeting at the ambitus.

Occasionally, there are two spines, side by side.

The tips of the spines support a broad lateral

fringe. There are 5 or 6 long slender adambulacral

furrow spines, webbed together in a curved series,

and grading from shortest inner spine to longest

outer spine. The mouth plates bear, on the side,

five small, equal spines, and apically, two longer,

stouter spines. There may be up to three long,

sturdy, nonhyaline suboral spines, or there may be

none. None of the mouth spines are webbed.

This species seems to be confined to the Leeward

Islands, in 350-360 fathoms.

MATERIAL EXAMINED.—Oregon Station 6696 (1)

[R=36 mm. r=16 mm, Rr=l:2].

Pteraster rugosus Clark

PLATE 33: FIGURES C, D

Pteraster rugosus H. L. Clark, 1941:61, pi. 6: fig. 1.

This nearly pentagonal species has a highly inflated

disc and five short, broad arms which curve upward

at the tip, so the end of the ambulacral groove is

actually on the abactinal surface; the interradial

arcs are wide and shallow. The dorsal membrane

is thin, with numerous small spiraculae, and contains many small calcareous deposits. It is produced

in a regular pattern of peaks by the underlying

spines. The cruciform abactinal plates have long,

Hat lobes and a tall pedicel, narrow at the base

but expanded to a stout, ridged or grooved top

which bears 5-8 spines, about as long as the pedicel

and not particularly slender or acute. The central

osculum is small and surrounded by a series of

continuous spines about the same length as the

paxillar spines. The actinal surface is plane and

distinctive by reason of its 20-30 flat, closely ser

actinolateral spines whose square tips support a

broad lateral fringe. The adambulacral furrow

spines are erect and webbed, in a curved series;

there are 4—6 spines (Clark says 5 or 6) and the

inner one or two are small, but the others are

long and not particularly slender or acute.

The mouth plates bear seven marginal spines,

webbed together, on each half; the two at the apex

of the plate are long and slender, and the others

decrease gradually in length to a small spine at

the outer edge of the margin. On the actinal face

of the mouth plate is a single large, stout suboral

spine, white at the base but hyaline at the tip.

The tube feet are notably smaller and more delicate than those of most species of Pteraster. This

species is distinctive because of its generally neat

and regular appearance compared to other Pterasteridae. The largest specimen in the collection has an
78 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

average of four adambulacral furrow spines, as

compared to the 5 or 6 noted by Clark, but it is

larger than Clark's largest specimen, and is undoubtedly the same species; this may be a function

of growth. The white color noted by Clark in live

material is retained in preserved specimens.

The distribution presently known is from northern Cuba to the Leeward Islands, in 50-255

fathoms.

MATERIAL EXAMINED.—Oregon Stations: 4994 (1)

[R=24 mm, r=17 mm, Rr=l:1.5]; 5937 (1) [R=

12 mm, r=7 mm, Rr= 1:1.8].

Pteraster militaroides Clark

PLATE 34: FIGURES A, B

Pteraster caribbaeus [part] Perrier, 1884:2I6.-Verrill, 1915:

82. pi. 7: fig. 4.-H. L. Clark, 1941:60.

Non Pteraster caribbaeus.—Perrier, 1881a.

Pteraster militaroides H. L. Clark, 1941:61.

The form of this species is plump and stellatopentagonal, with the interradii slightly indented.

The five arms are recurved at the tip, so the end

of the ambulacral groove is on the abactinal surface.

The supradorsal membrane is heavily encrusted

with numerous small calcareous deposits, and has

a lacy appearance due to the many small spiraculae.

The membrane rests loosely on the underlying

paxillar spines, so the dorsal surface is rugose, but

the spines do not protrude through the membrane

nor lift it in tent-like peaks above the general

surface. The lobes of the abactinal plates are short,

broad, and flat, and the pedicel is thick, columnar,

and short; it bears 6-12 long, slender, glassy spines,

not acute. Usually, there is one central spine and

about nine peripheral spines, all about of equal

length. The central osculum is of moderate size and

surrounded by a continuous ring of numerous,

close-set spines about the same length as the paxillar

spines.

The actinal membrane is mostly without deposits; it is supported by 20-25 short, ridged, blunttipped actinolateral spines which neither reach the

ambitus nor support a marginal fringe; the supradorsal membrane extends well onto the actinal

surface interradially. There are a few small, supplementary actinolateral spines, either near the

base of the large actinolaterals or between a pair

of them. There are seven adambulacral furrow

spines (six distally; on the smallest specimens,

there are 6 or 5 spines) webbed together in a curved

series and graded from shortest at the inner (or

groove) end of the series to longest at the outer

end. The mouth plates have a prominent central

ridge along the suture. Each plate bears a long,

stout, hyaline suboral spine and a marginal series

of about eight spines (6—10), four at the side of the

plate and, at right angles to these, four along the

front margin. The four spines on the side are short

and subequal; the four on the front margin are

longer and grade upward to longest at the apex. All

are slender and blunt tipped, and none are nearly

as long as the suborals.

Clark reported this species as occurring only in

northwestern Cuba; this collection contains specimens from off the Leeward Islands and from the

western Caribbean, between the Caymen Islands

and Panama. The depth range is 250—355 fathoms.

MATERIAL EXAMINED.—Oregon Stations: 1917 (2)

[R=23 mm, r=ll mm, Rr=l:2]; 1885 (1) [R=17

mm, r=9 mm, Rr=sl:9]; 1929 (1) [R=22 mm, r =

13 mm, Rr=l:1.8]; 5929 (1) [R=19 mm, r=ll

mm, Rr= 1:1.7].

Since the publication of my brief paper on Marsipaster acicula Downey, 1970c, I have had the opportunity to examine many more specimens of

Pterasteridae and have concluded that Marsipaster

Sladen (1882) is not a good genus; it is practically

indistinguishable from Pteraster. The characters

on which Sladen based Marsipaster are minor and

subject to variation even within a species. The

nature of the supradorsal membrane in Marsipaster

is certainly distinctive, but scarcely of generic

significance when one considers how this feature

may vary between species of Pteraster. The web

connecting the suboral spine to the other mouth

plate spines is certainly not of sufficient importance

to be a generic character, while the placement of

the adambulacral spines (Sladen says high in the

furrow and horizontal in position) does not hold

true for both of the species described by Sladen in

his paper (true for M. spinissimus, not true for

M. hirsutus). In light of these conclusions, it is

thought appropriate to include here a more

thorough description of the species I described as

Marsipaster acicula.
NUMBER 126 79

Pteraster acicula (Downey)

PLATE 34- FIGURES C, D

Marsipaster acicula Downey, 197Oc:3O9, fig. 1.

This small, plump form is pentagonal, with an

inflated disc and five short arms, hardly produced

beyond the disc. The supradorsal membrane is thin,

fragile, and transparent, and strongly reminiscent

of spongin. Spiraculae are small and numerous.

The pedicels of the abactinal plates are of moderate

length, sturdy and columnar. They bear a crown of

8—25 long, fine, glassy, acute spines which are

webbed together at their bases, below the supradorsal membrane, like an inverted umbrella. These

spines protrude through the supradorsal membrane

for as much as half their length. The central osculum is inconspicuous and is surrounded by numerous fine spines, exactly like the paxillar spines,

webbed together in a continuous ring around the

osculum. The actinal interradial area is small, the

supradorsal membrane continuing onto the actinal

surface in this region. The actinolateral spines are

large, but not much longer than the longest adambulacral spine. A few spines extend beyond the

actinal membrane and may be webbed together,

but there is no continuous lateral fringe. The six

erect adambulacral furrow spines are webbed

together for their entire length; the inner two are

small and parallel to the groove, while the outer

ones are longer and set at right angles to the groove.

The first adambulacrals are webbed continuously

across the base of the mouth plates. The mouth

plates are broad and bear five free marginal spines

and one suboral spine, the suboral and apical

marginal spines subequal, long, stout, acute, and

glassy-tipped.

This species seems to be confined to the northern

Gulf of Mexico, in 142-720 fathoms.

MATERIAL EXAMINED.—A laminos Station 12A/68-

A-13 (1). Specimen too damaged for meaningful

measurements, but the species is very small (holorype: R=8 mm).

Pteraster species

PLATE 35: FIGURES A, B

The form of this specimen is distinctly stellate, with

a moderately broad disc and five long attenuate

arms. The dorsal membrane is thin. Spiraculae are

moderately large and few. Large rounded granules

of calcite are present in the membrane. The pedicels of the cruciform abactinal plates are columnar,

stout, and about as long as the lobes of the plate;

they bear 6-15 longer, clear spines, ridged and

lattice structured like the spines of many echinoids.

The five plates surrounding the osculum have a

thick pedicel, only slightly expanded at the top and

bearing about twenty long, slender, webbed spines.

On the actinal surface, 25—30 long, sturdy, blunttipped spines, of the same lattice structure as the

dorsal spines, support a narrow marginal fringe.

These spines meet at the interradius. The adambulacral plates bear a curved series of six webbed,

subequal spines, fine and acute. The broad mouth

plates bear a single long, blunt suboral spine,

slightly curved; the rest of the mouth-plate armature is, unfortunately, badly damaged. It seems to

consist of a thick preoral spine, webbed to an adjacent, slightly smaller spine, and, at the side of the

plate, three small unwebbed spines. The oral

armature and the structure of the spines in general,

as well as the height and thickness of the pedicels

and the presence of round granular deposits in the

supradorsal membrane and the attenuate arms,

serve to distinguish this specimen from Pteraster

caribbaeus, which appears to be its nearest relative.

Although I believe this to be a hitherto undescribed

species, formal description will have to await more

and better-preserved material.

MATERIAL EXAMINED.—Oregon II Station 10827

(1) [R=26 mm. r=10 mm, Rr=l:2.5].

Calyptraster Sladen

Calyptraster Sladen, 1882:207. [Type, by original designation,

C. coa Sladen.]

This genus, because of the partly webbed furrow

spines, not webbed to the actinolateral, and

oblique, rather than transverse, in position, seems

to fall exactly between Pteraster and Hymenaster.

The fact that one species, C. coa, has a delicate

lacy supradorsal membrane like many Pterasters,

and the other, C. personatus, a thick, robust, somewhat gelatinous membrane, like most Hymenasters,

further supports this view, which Sladen pointed

out in his original diagnosis of the genus.

Only two species of this genus are known; Calyptraster coa Sladen is known only from off Brazil.
80 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Calyptraster personatus (Perrier)

PLATE 35: FIGURES C, D

Cryptaster personatus Perrier, 1894:191, pi. 14: fig. 3.

Calyptraster personatus.—Madsen, 1947:3, figs. 1-2.

The depressed form of this species is broadly stellate and the abactinal surface is plane. There is

a sharp ambital demarcation between the actinal

and abactinal surfaces. The supradorsal membrane

is strong, muscular, and gelatinous, and spiraculae

are of moderate size and numerous. The dorsal

paxillar columns are short and robust and bear

a crown of 4-6 short, blunt, divergent spines which

support the membrane but neither protrude

through it nor raise it in peaks. The central osculum is surrounded by a ring of short spines, and

the five large triangular valves consist each of 11

or 12 spines webbed together, the shortest at either

end and the longest in the middle. The actinal

membrane is supported by eighteen or more sturdy,

slightly flattened actinolateral spines with blunt

(or even expanded) tips, the sixth or seventh being

the longest. The actinolateral spines do not meet

in the interradius. There are four short, slender,

acute adambulacral furrow spines in oblique series,

partially webbed by a delicate membrane; the webbing does not extend to the actinolateral spine, and

there is a distinct hiatus between the furrow spines

and the actinolateral spine.

The mouth plates are small and nearly concealed

by 3 or 4 pairs of huge, clavate, thorny-tipped sub

oral spines (Sladen and Madsen were inclined to

consider the anterior pair as epioral, but I believe

that, because their character and position are the

same as the two or three pairs behind them, they

must be considered suborals). Well within the

mouth and on the side margin of each mouth plate

is a single small acute spine (sometimes two).

This specimen establishes beyond a doubt that,

despite Madsen's speculations, C. personatus and

C. coa are separate and distinct species. Madsen

calls C. coa the "western Atlantic species" and C.

personatus the "eastern Atlantic species"; this species has hitherto been reported only from the

Azores and the Madeira Islands, from 2,150-2,995

meters.

MATERIAL EXAMINED.—Oregon Station 2567 (1)

[R=20 mm, r=10 mm, Rr=l:2].

Family GANERIIDAE Sladen, 1889

This family is characterized as having small mouth

plates, not spade shaped or plowshare shaped. The

ambulacral grooves are narrow, and the marginal

plates are large and blocklike. The abactinal skeleton is reticulate or imbricate, and the actinal plates

are in regular transverse rows, with one or two

large spines or a group of small spinelets.

Only one species, of the monotypic genus Leilaster, is represented in the Caribbean; all other members of the family are confined to the polar regions.

Leilaster A. H. Clark

Leilaster A. H. Clark, 1938:1-7. [Type, by original designation,

Korethraster radians Perrier.]

Leilaster radians (Perrier)

PLATE 36: FIGURES A, B

Korethraster radians Perrier, 1881a: 12.

Korethraster hispidus Perrier [non Wyville Thomson], 1884:

212, pi 6: figs. 9-11.

Solaster radians.—Perrier, 1884:275 [in explanation of plates],

pi. 6: figs. 9-11.

Lophaster radians—Perrier, 1884:167-170 [in list of species].

Korethraster ? radians.—Sladen, 1889:459, 463.

Leilaster radians.-A. H. Clark, 1938:1-7, pi. 1.

This small form has a broad disc and five moderately long, broad arms. The disc is strongly

arched upward and the downward-directed arms,

high in the midregion, slope off steeply at the sides.

The tumid dorsal plates are oblong except at the

disc center, where the primary plates are round.

Papulae are single and confined to the radial areas.

The plates on the arms are in regular longitudinal

rows, a carinal and two adradials extending the

full length of the arm; between the second adradial

row and the superomarginals are one or two additional rows which do not reach the end of the arm.

The second adradials are larger than the carinals

and the first adradials. The plates imbricate upon

one another in the direction of the disc. The large

blocklike superomarginals are rectangular and vertical. The inferomarginal plates are paxilliform,

with the plate produced in a stout column extending downward. All abactinal plates bear short

clavate spinules with thick, rounded, thorny heads,

so the general appearance of the dorsum is openly
NUMBER 126 81

granular. The spinules of the dorsum are so spaced

that conspicuous channels occur between the plates,

and a broad sulcus is present in the interradius.

The inferomarginals bear, on the outer edge, spinules of quite a different character: large, stout, and

conical.

There is a single row of small actinal plates between the inferomarginals and the adambulacral

plates, unornamented or bearing one or two small

spinules. The adambulacral plates bear a furrow

series of three long acute spines, webbed together

and nearly parallel to the groove, the proximal

spine shorter than the other two. On the actinal

face of the plate is a comb of three longer spines

in transverse series, incompletely webbed. The

mouth plates are long and broad, and bear six long,

subacute, divergent spines on the margin. In the

specimens on hand, the actinal face of the mouth

plate is bare. The madreporite is small, inconspicuous, and slightly sunken between two of the

primary dorsal plates. The oculars are large and

cordiform.

This odd little species is a classificatory puzzle;

Clark placed it in the Ganeriidae, and Fisher

(1911a) does not disagree, although he points out

its affinities with the genera Mirastrella and Kampylaster, of the family Asterinidae. The character of

the inferomarginal plates is quite like the genus

Lophaster, of the family Solasteridae. The ill-defined Ganeriidae are a rather unsatisfactory family

whose systematic position is open to question.

This species has previously been reported from

Cuba, Barbados, and Puerto Rico, in 56-140 fathoms.

MATERIAL EXAMINED.—Oregon Station 4226 (6^

[R=8 mm. r=4 mm, Rr=l:2].

Family PORANIIDAE Perrier, 1894

It is interesting that this family, commonly represented in colder waters by medium to large species,

should apparently be limited in the Caribbean and

Gulf of Mexico to very small, immature-looking

species. Verrill (1915) reported Poraniella regularis (R=12 mm), Poraniella echinulata (R=5

mm), Marginaster pectinatus (R=5 mm), and Porania ? austera (R=17 mm), and speculates that

they may be juveniles of some species of Porania.

I find this hard to believe, as at least some of the

specimens were sexually mature, and no larger

Poraniidae have been taken in this area. All of

the above species were taken in 23-163 fathoms.

Systematically, this whole family is in need of

revision, consisting as it does of very diverse forms,

related variously to the Asterinidae, the Echinasteridae, the Ganeriidae, and the Oreasteridae, far

removed in the currently accepted classification.

Poraniella, as Verrill pointed out, is very close to

Asterina, in the family Asterinidae; Spoladaster,

according to Fisher, is close to Perknaster, in the

family Ganeriidae; and Pseudoporania and Sphaeriaster seem close to some of the Oreasteridae (notably Culcita). Chondraster and Tylaster may lack

any skeleton except for certain radial elements, and

it is hard to see why they should be placed in this

family rather than practically any other. As presently constituted, the family Poraniidae defies reasonable definition.

Poraniella Verrill

Poraniella Verrill, 1914:19. [Type, by original designation,

P. regularis Verrill.]

Poraniella regularis Verrill

PLATE 36: FIGURES C, D

Poraniella regularis Verrill, 1914:19; 1915:70, pi. 7: figs. 1-la,

pi. 15: figs. 5-56.

This small stellate starfish has a broad disc and

five short, broad-based, tapering arms. It is depressed, and the dorsum is sunken interradially,

throwing the radial areas into prominence. The

large abactinal plates are in regular longitudinal

rows. The primary radials are large and roughly

rectangular; their distal ends are overlapped, as are

the rest of the carinals, by the pentagonal or stellate first adradials. All abactinal plates are slightly

tipped up at the proximal end, and a large papular

pore appears under the proximal edge of each radial plate; there are no interradial papular pores.

The upraised edge of the plate also bears 1-4 small

acute spinules. The superomarginal plates are

large, oval, and (unlike Verrill's type specimen)

broader than long; they bear a few small acute

spinules near the outer edge. The inferomarginal

plates project beyond the superomarginals, but do

not lie side by side with them as they do in the
82 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

type, where the two series of marginals form a

double border around the abactinal surface. The

inferomarginals bear, on their outer edge, a row

of about half a dozen small acute spines and above

them 1—3 smaller spines. The actinal interradials

are large, roundish, slightly tumid, in regular chevrons with an unpaired interradial in all but the

first row.

They usually bear a small acute spinule near the

distal edge. The adambulacral plates bear, on the

furrow margin, three (four proximally, two distally) short, blunt, palmate spines and on the actinal face of the plate two longer, erect, thorny

spines. The mouth plates bear two large, blunt,

slightly flattened apical spines, the first larger than

the second, and on the side margin, three small

spines similar to the adambulacral furrow spines.

There is a single large suboral spine on each mouth

plate. The madreporite is small, round, and deeply

channeled; the oculars are oval; and there are no

pedicellariae. Actinal, abactinal, and marginal

plates are covered with minute glassy tubercles.

This species has been found off west Florida, the

Bahamas, and Cuba, in 36-169 fathoms.

MATERIAL EXAMINED.—Silver Bay Station 961 (1)

[R=17 mm, r=10 mm, Rr= 1:1.7].

Marginaster Perrier

Marginaster Perrier, 1881a: 16. [Type, by subsequent designation, M. pectinatus Perrier (Sladen, 1889:364).]

Cheilaster Bell, 1892:81. [Type, by original designation,

Marginaster fimbriatus Sladen.]

Marginaster pectinatus Perrier

PLATE 37: FIGURES A, B

Marginaster pectinatus Perrier, 1881a:16; 1884, p. 229, pi. 1:

figs. 4, 5; 1894, p. 167.-Sladen, 1889, p. 366.-Verrill, 1914,

p. 18; 1915, p. 76.

This small subpentagonal form has an arched dorsum and the actinal surface is plane. There is a

ridge down each midarm, and the abactinal interradial areas are sunken. The carinal plates are

large and the primary interradials enormous, but

other dorsal plates are generally small and irregular. Most plates bear 1-8 small, stumpy, thorny

spinules at the periphery of the plate. Papular

pores occur in the area between midarm and midinterradius. The superomarginal plates increase in

size from interradius to the end of the arm, becoming broader than long, and the last three or four

plates are separated on the midarm only by a single

carinal plate. In the interradial area, they bear

spinules similar to those of the dorsal plates (they

are scarcely distinguishable from the dorsals), but

distally the superomarginals are bare. They are

broader and imbricating near the top, but narrower

and separated by a papular pore at the bottom,

between the superomarginals and the inferomarginals.

The inferomarginal plates project beyond the

superomarginals. They are about twice as broad

as long and bear on the outer edge 4 or 5 moderately long, stout, ridged, and rather thorny subhyaline spines, slightly flattened dorsoventrally.

Above these is a row of 3 or 4 much shorter similar

spines. The actinal face of the inferomarginals is

plane and bare. The actinal interradial plates are

in 3 or 4 rows parallel to and alternating with the

inferomarginals; the central plate in each row is

larger than the others. The plates are irregularly

discoid and bare. The adambulacral plates bear

three moderately long, sturdy, blunt spines in transverse series, the inner furrow spine smallest. The

mouth plates, longer than broad, bear a stout,

blunt, flattened preoral spine at the apex, a similar

but smaller spine beside it, and two small spines

at the side; there are 2 or 3 larger suboral spines.

The madreporite is small, flat, round, and deeply

channeled. The oculars are square, with a rounded

distal edge which tilts up slightly.

This uncommon species occurs in the West Indies, in about 100 fathoms.

MATERIAL EXAMINED.—Oregon Station 4226 (5)

[R=ll mm, r=6.5 mm, Rr = 1:1.8].

Family ECHINASTERIDAE Verrill, 1867

The family Echinasteridae is characterized by having a well-developed abactinal skeleton, with plates

in more or less irregular longitudinal and trans

verse series forming an open meshwork, the plates

bearing isolated spines or groups of spinules. The

marginals are inconspicuous; the ambulacral

grooves are narrow; and the suckered tube feet are

in two rows. The mouth plates are small, and there

are no pedicellariae.
NUMBER 126 83

In the Gulf of Mexico and the Caribbean Echinasteridae, there is some confusion not only about

the species, but also the genera. Of the eight genera

in the family, only Henricia, Echinaster, and Thyraster have been reported from this area. Henricia

is distinguished from the other two (so-called) genera in having groups or bundles of spinules on

the plates, rather than isolated spines or spinelets.

Thyraster (type species by monotypy— T. serpentarius) is supposedly distinguished from Echinaster

by having rectangular plates which imbricate at the

corners; however, the plates of specimens in this

collection range from rectangular to triangular, to

polygonal to round, and other characters of this

monotypic genus are specific characters, not generic

ones. I believe that the genus Thyraster should be

dropped and T. serpentarius returned to its original position in the genus Echinaster, where it was

first described by Muller and Troschel.

The western Atlantic species of Echinaster could

then be readily distinguished from other genera in

the family by the presence of a patch of glassy tubercles on the plates. This would necessitate the

removal of Echinaster spinulosus, which lacks these

patches of glassy tubercles, from the genus, and a

new genus is here erected for this common Gulf

and Caribbean echinasterid, which I propose to

call Verrillaster, for A. E. Verrill, whose report on

the starfishes of Florida, the West Indies, and Brazil has been an invaluable aid in this present study.

Key to the Genera of Echinasteridae

1. With a patch of glass tubercles, concealed by skin, on most principal plates Echinaster

Without glassy tubercles on plates 2

2. With a bundle or group of spinules on principal plates Henricia

With isolated spines or spinelets Verrillaster, new genus

Henricia Gray

Henricia Gray, 1840:184. [Type, by original designation, H.

oculata Gray (—Asterias sanguinolenta O. F. Muller).]

Linckia Forbes [non Nardo], 1839:120. [Type, by original

designation, Asterias oculata Pennant.]

Cribella Forbes [non Cribrella L. Agassiz, 1835b], 1841:100.—

Lapsus calami.

Cribrella Lutken [non L. Agassiz, 1835b], 1859:93. [Type, by

original designation, C. sanguinolenta (—Astcrias sanguinolenta O. F. Muller).]

Madgalenaster Koehler, 1907:19. [Type, by original designation, M. arcticus Koehler.]

Cyllaster A. H. Clark, 1916:61. [Type, by original designation,

C. setninuda Clark.]

Spinohenricia [subgenus] Heding, 1936:8. [Type, by original

designation, Henricia (Spinohenricia) scabrior (zzCribrella

sanguinolenta forma scabrior Michailovski, 1903).]

Closely meshed skeleton bearing numerous spinelets in groups or scattered along ridges of skeleton;

marginal plates more or less distinguishable; adambulacral plates with one or more spinelets deep

within furrow.

Key to the Species of Henricia

1. With five arms 2

With six arms, or more H. sexradiata

2. Two to five inner furrow spines, one above the other in single series H. antillarum

Six inner furrow spines, not one above the other in single series (may be roughly in two

series of three each) Henricia species

Henricia sexradiata (Perrier)

PLATE 37: FIGURES C, D

Cribrella sexradiata Perrier, 1881a:8; 1884:209, pi. 4: fig. 6.

Henricia sexradiata.-Verrill, 1914:371; 1915:47, pi. 11: fig. 7.

H. L. Clark, 1941:58.

This small species is the most distinctive Henricia

of the Caribbean and Gulf of Mexico, quite apart

from its six or more arms. The disc is small and

the arms long, rounded, evenly tapering to a subacute point. The irregularly shaped abactinal
84 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

plates—elongate, lobed, or rounded—imbricate upon

one another to form a close reticulum. The papulae are small, and usually there are several to each

pore area. The abactinal plates are covered with a

dozen or more small, erect, thorny spinules. Both

series of marginals are large, distinct, equal, and

imbricate; they bear spinules similar to those of the

abactinals and are separated from the adambulacra]

plates by a single row of actinals extending less

than one third the length of the arm. Papular

pores occur between the marginals and between the

inferomarginals and the adambulacral plates. Actinai plates are few and rounded, with papular

pores extending to the mouth plates; they bear

spinules similar to those of the marginals and abactinals.

The adambulacral plates are broader than long,

rectangular, and bear, on the strongly carinate furrow face, three short appressed furrow spines, one

above the other, deep within the narrow ambulacral

groove. On the outer furrow margin are three

stout, thorny, subclavate spines, about twice the

length of the actinal spinules, and behind them is

a second series of 2 or 3 similar, slightly shorter

spines. The actinal face of the plate is covered

with spinules similar to those of the marginals.

The mouth plates are narrow and triangular, with

one or two stout, blunt, flattened preoral spines;

the rest of the spinulation is like that of the adambulacral plates. There is usually more than one

madreporite, raised, flattened, and bearing numerous spinules. The oculars are small, round, and

covered with small thorny spinules. This is a fissiparous species.

This species ranges from the northern Gulf of

Mexico to Surinam, in 75-190 fathoms.

MATERIAL EXAMINED.—Oregon Stations: 2289 (1)

[R=46 mm, r=6 mm, Rr=l:8 (three long arms,

four short arms)]; 36 (1) [R=49 mm, r=10 mm.

Rr=l:5 (six equal arms)].

Henricia antillarum (Perrier)

PLATE 38: FIGURES A, B

Cribrella antillarum Perrier, 1881a:8; 1884:207, pi. 3: fig. 3.

Henricia antillarum.-Verrill, 1914:210; 1915:46.-H. L. Clark,

1941:56.

Henricia microspina Verrill, 1915:48.—H. L. Clark, 1941:57.

This species has a small disc and five long, slender,

emaciated-looking arms. The arms are broad at the

base, but beyond the basal one fourth or one fifth

portion they are extremely attenuate. The abactinal plates are mostly C-, Y-, or X-shaped, with

many small round plates scattered among them.

They are covered with minute, glassy-tipped spinules, mostly in irregular rows on the plates. The

papular pores are large and the papulae are single.

Superomarginal plates are smaller than the nearly

square inferomarginals, but distally the two series

tend to become more equal. They bear spinules

like those of the abactinal plates. A single row of

actinals may extend a short distance beyond the

disc; the interradial actinals are few, crowded, and

small. Papular pores occur to the adambulacral

plates. Actinal spinules are smaller than those of

the abactinal surface.

The adambulacral plates have a sharply carinate

furrow face bearing 2-5 small furrow spines, one

above the other. On the furrow margin are 4-6

long, stout, blunt spines, and the actinal face of

the plate bears up to twenty short, stout, aciculate

spinules, becoming increasingly similar to those of

the inferomarginals. The long narrow mouth plates

bear one or two stout, blunt preoral spines at the

apex and 2 or 3 minute spines on the side margin,

well within the mouth. Behind the preoral spines

are 2-5 similar spines, not quite so stout. The rest

of the plate is covered with short, stout, aciculate

spinules. The single madreporite is rounded,

raised, and covered with spinules.

I believe Verrill's Henricia microspina is the

same as H. antillarum, despite H. L. Clark's assertion that the two species were distinguishable on

the basis of length of spinules and size of skeletal

meshes. I have examined the types of both species

and can find no specific differences. Allowing for

the difference in size of the two specimens (H. antillarum, R=54 mm, vs H. microspina, Rr=20 mm),

there is no essential difference in the length of

spinules or size of mesh of the skeleton. The number, size, type, and arrangement of the adambulacral spines, and the general form of the starfish

seem to be more reliable characters for distinguishing the species of Henricia.

However, there is a third species of Henricia in

the Caribbean. Because the types of so many species are not available, or are in very poor condition

as regards essential features, it is probably not de-
NUMBER 126 85

sirable to add still another name to this already

overcrowded genus without more material.

Henricia antillarum probably does not occur

south of about 15° N. Its depth range is 175-390

fathoms.

MATERIAL EXAMINED.— A laminos Station 25/70-

A-10 (1) [R=55 mm, r=10 mm, Rr=l:5.5].

Oregon Stations: 489 (1) [R=26 mm, r=4 mm,

Rr—1:6]; 491 (1) [R=28 mm, r=5 mm, Rr

= 1:5.5].

Henricia species

PLATE 38. FIGURES C, D

This species has a small disc and five long, tapering,

cylindrical arms, thick at the base, and slightly

constricted. The abactinal plates are thick, elongate, and imbricate. There are also some small

irregularly rounded plates. All bear numerous

small, divergent, thorny-headed spinules. The

meshes of the skeleton are large, as are the papulae, and the papulae are usually single, but there

may be up to four per mesh. The superomarginal

plates are small and triangular, and proximally

they may be separated from the inferomarginals by

a few small rounded plates. The inferomarginals

are broader than long, and about twice the size of

the superomarginals. The spinulation is like that

of the abactinals. The adambulacral plates are

about twice as broad as long. On the broadly carinate furrow surface, they bear, deep within the

furrow, about six short, stumpy spines, roughly in

two vertical rows of three each. Above them, on

the outer furrow margin of the adambulacrals, are

about 6-10 long, thick, blunt spines; owing to the

sloping face of the furrow margin, they do not

reach much beyond the height of the short, thick,

thorny spinules on the actinal face of the plate,

which are numerous but spaced. The spines on the

apical margin of the mouth plates are thick, blunt,

and crowded.

The plates themselves are long, narrow, and triangular, and the actinal face is evenly covered with

short, thick, thorny, spaced spinules. The madreporite is small, round, raised, and covered with radiating gyri; the smaller of the two specimens below

had three madreporites.

MATERIAL EXAMINED.—Oregon Stations: 35 (1)

[R=30 mm, r=6 mm, Rr=l:5]; 2644 (1) [R=62

mm. r—12 mm, Rr=l:5].

Echinaster Muller and Troschel

Echinaster Muller and Troschel, 1840:102. [Type, by subsequent designation, Asterias seposita Lamarck (non Retzius)

(=Asterias sagena Retzius, 1805) (Fisher, 1913:195).]

Othilia Gray, 1840:281. [Type, by original designation, Asterias spinosa Retzius (=Pentadactylosaster spinosus

Linck).]

Rhopia Gray, 1840:282. [Type, by original designation, Asterias seposita Retzius (=Pentadactylosaster reticulatus

Linck).]

Henricides Verrill, 1914:210. [Type, by original designation,

Henricia heteractis H. L. Clark, 1909.]

Thyraster Ives, 1890:329. [Type, by original designation,

Echinaster serpentarius Muller and Troschel.]

Disc small, five arms long, more or less cylindrical;

spines on plates isolated, never in groups or bundles; patch of glassy tubercles on most principal

plates; skeleton an open meshwork of plates connected by secondary plates.

This genus is perhaps the most confusing one

occurring in the tropical and subtropical Atlantic.

Not only are the species ill-defined, but the genus

itself has undergone various taxonomic changes.

Fisher (1919), in attempting to define the status of

the genus, compounded the confusion by making

several unwarranted assumptions and overlooking

important references. His arguments for separating the American species from the European ones

and placing them in another genus, Othilia, are so

involved and seem to me so feeble that it would be

preferable to retain Echinaster for these very common Gulf and Caribbean species, a name well

known and firmly established in the literature.

In examining the Echinasters in this collection,

I found it relatively easy to distinguish what seem

to be eight species. Echinaster braziliensis, E. sentus, E. echinophorus, E. modestus, and E. serpentarius could be identified definitely. Rather than

add what are probably quite unnecessary names to

the genus, the other three species I have distinguished have here been given letters in the hope

that this will simplify matters for some future reviser. Of the three species I have been unable to

identify, none seem to fit Tommasi's (1970) new

species. E. densispinus or E. nudus.
86

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to the Species of Echinaster

1. Five rows of primary plates 2

More than five rows of primary plates 6

2. Arms tapering '

Arms thick and blunt 5

3. Numerous small calcareous deposits under skin Echinaster species A

No calcareous deposits under skin 4

*. Glassy tubercles few, sunken in pits in plates Echinaster species B

Glassy tubercles not sunken in pits

 E

- modestus

5. Adambulacral plates with one spine within furrow and one on margin E. echinophorus

Adambulacral plates with 3 or 4 spines, 1 or 2 in furrow and 1 or 2 on margin E. sentus

6. Two to five spines on each primary plate E. serpentarius

One spine on each primary plate 7

7. Arms thick and blunt Echinaster species C.

Arms tapering

 £

« brasiUends

Echinaster serpentarius Muller and Troschel

PLATE 39: FIGURES A, B

Echinaster serpentarius Muller and Troschel, 1842:24.—

Perrier, 1875c:370.-Lutken, 1871:285.

Thyraster serpentarius.-l\es, 1890:329, pi. 8: figs. 19-21-

Verrill, 1915:44.-Tommasi, 1970:15, fig. 40.

This stellate species has a small disc and five moderately long, smoothly tapering arms. The regular

longitudinal rows of plates distinguish it from other

Echinasteridae. The plates are rectangular to

round, and imbricate. A carinal row of plates alternates left and right of the median dorsal line

on the arm; the adradial rows on either side of

the carinals (usually only one on each side, but in

the largest specimen there are two) appear to cross

each other at the base of the arm and continue

down the opposite side of the adjacent arm. The

superomarginal plates define the ambitus, and the

inferomarginals, larger than the other plates, adjoin the adambulacral plates. Frequently, there is

an incomplete row of plates between the adradials

and the superomarginals and between the superomarginals and the inferomarginals. All plates except the adambulacrals have a large irregularly

rounded patch of glassy tubercles on the surface

and bear, near the distal edge, 2-5 short, thick,

stubby spinules.

The papular areas between the plates may bear

up to ten large papulae, or more on the largest

specimens. The adambulacral plates bear, within

the furrow, a single slender spine, and two thicker,

obliquely situated spines on the furrow margin.

On the actinal face, they usually bear one or two

short spinules. The adambulacral plates are

broader than long, and spaces between them are

about the same length as the plate. The mouth

plates are small and narrow and bear one or two

large, stout subclavate preoral spines, one or two

similar large marginal spines, and usually a small

acute suboral spinule. The round, slightly raised

madreporite is covered with radiating gyri with

small spinules. The small oculars are round or

oval. The tegument is fairly thick.

This species has hitherto been considered a

strictly Mexican species, distributed from Vera Cruz

around the Yucatan Peninsula; the specimens below are all from northern Florida north. It is a

shallow-water species, not found in depths much

below about 50 fathoms.

MATERIAL EXAMINED.—Oregon Stations: 2205 (1)

fR=29 mm, r=5 mm, Rr=l:6]; 1672 (1) [R=43

mm, r=9 mm, Rr=l:5]. Silver Bay Stations: 1970

(1) [R=20 mm, r=5 mm, Rr=l:4]; 2775 (4) [R

= 4 0 mm, r=9 mm, Rr=1.4]; 1710 (2) [R=92 mm,

r=20 mm, Rr= 1:4.5].

Echinaster echinophorus (Lamarck)

PLATE 39: FIGURES C, D

Asterias spinosa [part ?] Retzius, 1805:18 [non Asterias

spinosa Pennant, 1777].

Asterias enchinophora Lamarck, 1816:560.

Othilia spinosa.—Gray, 1840:281; 1866:12.

Echinaster spinus Muller and Troschel [part], 1842:22.—

Lutken, 1859:90.-Perrier, 1869:57.

Echinaster (Othilia) crassispina Verrill, 1868:368, pi. 4: fig. 7.

Echinaster crassispinus.-Lutken, 1871:283 [61].—Ives, 1890:

326.

Echinaster echinophorus.-Perrier, 1875c:364.—Rathbun, 1879;

147.-Ives, 1889:171.-Verrill, 1915:42.-Tommasi, 1970:16,

figs. 41-43.
NUMBER 126 87

This species has a small disc and five short straight

arms, terminating abruptly. The rows of plates are

carinal, adradial, superomarginal, inferomarginal,

and adambulacral, with no intermediate plates except a few small actinal interradial plates on the

ventral surface of the disc. Only a few plates bear

spines, but those which do are tumid, almost mammiliform, and the spines are huge, movable, and

acute. Usually, the carinal and superomarginal

plates lack spines.

All plates without spines bear a large round

patch of coarse glassy tubercles. Papular areas between the plates are large and papulae numerous.

A large tissue-covered area at the center of the disc

dorsum contains one or two small "floating" plates

and a small, off-center anus raised in a minute epiproctal cone. The small madreporite is raised,

round, and the gyri are concealed by small spinules.

The ventral surface of the disc is without spines,

and the papulae on the ventral surface are small,

single, and scattered. The adambulacral plates

bear a single small acute spine within the furrow

and a single larger subacute spine on the margin.

The small mouth plates bear a single epioral spine

common to each pair, and on the margin of each

plate are two similar, widely separated spines; these

spines are small and blunt.

This is mainly a Brazilian species, and has not

previously been reported north of Yucatan. It is

confined to shallow water.

MATERIAL EXAMINED.—Oregon Station 5456 (1)

[R=37 mm. r=9 mm, Rr=l:4].

Echinaster sentus (Say)

PLATE 40: FIGURES A, B

Asterias sentus Say, 1825:143.

Othilia aculeata Gray, 1840:281; 1866:12.

Echinaster spinosus [part] Muller and Troschel, 1842:22.—

Verrill, 1867:343.-Ives, 189O:S25.-H. L. Clark, 1898a:6.

Othilia spinosa.—A. Agassiz, 1869:308.

Echinaster sentus.—Lutken, 1871:60 [284].—Perrier, 1875a:

366.- A. Agassiz, 1877:97, pi. 10: figs. l-6.-Rathbun, 1879:

147.-Verrill, 1915:36, pi. 29: fig. 2.-Tommasi, 1970:17. figs.

46-48.

The small disc and five thick, stumpy arms of this

species are distinctive. The rows of plates are:

carinal, adradial, superomarginal, inferomarginal,

and adambulacral. Each plate bears a large cqnical

acute spine, and many plates, particularly the secondaries (which lack a spine), bear a small round

flat patch of glassy tubercles. The papular areas

are large and the papulae numerous, less numerous

on the actinal surface. The adambulacral plates

bear one or two slender spines within the furrow,

and one or two heavier, subacute spines, one behind the other, on the marginal and actinal surfaces.

The mouth plate bears a heavy, blunt apical

spine, 2 or 3 smaller marginal spines, and a small

acute suboral spine. The adambulacral marginal

spines are webbed together longitudinally. The

madreporite is small, round, with radiating gyri.

and small spinules. Scattered over the entire dorsal surface, not confined to the papular areas, are

large discrete globules of a dark reddish brown, of

unknown nature and function (photoreceptive pigment spots ?).

This species is widely distributed from North

Carolina to Brazil, in shallow water.

MATERIAL EXAMINED.—Oregon Stations: 1936 (2)

[R=62 mm, r=15 mm, Rr=l:4]; 1937 (3) [R=70

mm, r=20 mm, Rr=l:3.5]; 1938 (3) [R=67 mm,

r=16 mm, Rr=l:4]; 1934 (5) [R=75 mm, r=18

mm, Rr=l:4].

Echinaster modestus Perrier

PLATE 40: FIGURES C, D

Echinaster modestus Perrier, 1881a:7; 1884:206, pi. 3: fig. 7.—

Verrill, 1915:43.

The disc of this species is small, and the five arms

are slender, moderately long, and cylindrical, tapering to a subacute terminus. The longitudinal rows

of plates are: carinal, adradial, superomarginal, inferomarginal, and adambulacral. There are few

actinal interradial plates between the superomarginals and the inferomarginals. Each plate bears,

on the distal edge, one or two small, stout, blunt

spines (regularly two on the inferomarginals, usually two on the superomarginals). The plates are

irregularly round and imbricate on the distal edge,

while the proximal surface of the plates bears a

round patch of glassy tubercles. The plates become

small, crowded, and decidedly imbricate at the end

of the arm. The adambulacral plates bear a small

slender spine deep within the furow, and two subequal blunt, thick spines set obliquely on the furrow margin; on the actinal face, well separated
88 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

from the furrow margin spines, is a single short,

blunt spine. The papulae are large, 6-8 per papular area. The mouth plates are small and narrow,

with one median and two marginal spines on each

plate; the spines are round, thick, and clavate. The

madreporite is small, round, raised, and covered

with small blunt spinules.

The oculars are small, rounded, and raised. The

covering tegument is quite thick.

The species distribution is from Florida to the

Lesser Antilles, in 13—123 fathoms.

MATERIAL EXAMINED.—Silver Bay Station 1909 (1)

[R=49 mm, r=ll mm, Rr=l:4.5]. Combat Stations: 90 (1) [R=52 mm, r=ll mm, Rr=l:5];

339-340 (1) [R=48 mm, r=ll mm, Rr=l:4.5).

Echinaster species A

PLATE 41: FIGURES A, B

The small disc and five moderately long, smoothly

tapering cylindrical arms which terminate in an

acute point are covered with a smooth, thick dermis. The most conspicuous feature of this species

is the numerous small, irregular calcareous deposits

under the skin. The plates are in regular longitudinal rows; carinal, adradial (connected by small

secondary ossicles), superomarginals (connected to

the adradials by small elongate secondary ossicles),

inferomarginals (close to, and corresponding with,

superomarginals), and adambulacrals. Actinal interradial plates are few or none. The round patch

of glassy tubercles on each plate (except the secondaries) is particularly well developed. A single,

small, somewhat spatulate spine occurs on the distal edge of many, but not all, plates. The adambulacral plates bear 1-3 small slender spines deep

within the furrow, and 2 or 3 stouter blunt spines,

more or less at right angles to the furrow, on the

furrow margin. The papular areas are large and

distinct, and the large papulae 6-20 per area dorsally, but fewer ventrally. The small, raised, flat

madreporite bears coarse radiating gyri without

spinules. The oculars are broader than long and

covered with glassy tubercles.

MATERIAL EXAMINED.—Oregon Station 4086 (8)

[R=84 mm. r=15 mm, Rr=l:5.5].

Echinaster brasiliensis Muller and Troschel

PLATE 42: FICURES C, D

Echinaster brasiliensis Muller and Troschel [part], 1842: 22.

-Lutken, 1859:9 [67].-Verrill, 1868:343, 368; 1915:41, pi.

26:1.-Perrier, 1875c:367. - Rathbun, 1879:148.-Ludwig,

1882:7.-Ives, 1890:324, 326, pi. 8: figs. 16-18.-Tommasi,

1970:17, figs. 44-45.

Othilia brasiliensis.-A. Agassiz, 1869:308.

The disc of this species is small, and the five arms

are stout, cylindrical, and taper to a subacute tip.

The regular rows of roughly triangular plates are

connected by oval or rectangular secondary plates

and bear a flat, triangular patch of glassy tubercles

and a single small, conical, blunt, erect spine on

the distal edge of the plate. The spines of the marginals are larger and more thornlike than those of

the other plates. The rows are: carinal, adradial,

dorsolateral (with a spine, but without a patch of

glassy tubercles), superomarginal, intermarginal

(short, not extending much beyond the disc, without spines or glassy tubercles), inferomarginal, and

adambulacral.

Large papulae cover the entire surface except

where interrupted by plates. There are 1—3 slender,

blunt adambulacral spines deep within the furrow,

and two larger, stouter spines, one behind the

other, at the furrow margin (1-3, usually two).

The one closest to the groove is the largest. The

mouth plates are small, with a short, stubby epioral

spine at the apex of each, two similar, slightly

longer spines on the side margin, and, frequently, a

suboral spine. The madreporite is small, oval,

raised, covered with radiating gyri, and without

spinules or with a few small ones. The tips of

the arms turn up, and the oculars are small and

broader than long.

This species is supposedly distributed from

Florida to Brazil; the Florida record is doubtful.

Its depth range is 11—57 fathoms.

MATERIAL EXAMINED.—Oregon Stations: 5739 (2)

[R=41 mm, r=10 mm, Rr=1.4]; 5723 (1 with four

arms) [R=57 mm, r=10 mm, Rr=l:5.7]; 2267 (1)

[R=48 mm, r=10 mm, Rr=l:4.8]; 2000 (1) [R

=27 mm, r=4 mm, Rr=l:7]; 2245 (1) [R=25 mm,

r =5 mm, Rr=l:5]; 2249 (1) [R=40 mm, r=6 mm,

Rr=l:7]; 2329 (2) [R=49 mm, r=10 mm, Rr=

1:4.9]; 2240 (2) [R=62 mm, r=12 mm, Rr=l:6];

2345 (1) [R=42 mm, r=10 mm, Rr=l:4.2]; 2343

(2) [R=38 mm, r=9 mm, Rr=l:4]; 1865 (1) [R

=60 mm, r=ll mm, Rr=l:6]. Oregon II Stations:

10212 (1) [R=65 mm, r=14 mm, Rr=l:4.5]; 10561

(1) [R=45 mm, r=6 mm, Rr=l:7].
NUMBER 126 89

Echinaster species C

PLATE 42: FIGURES A, B

This robust species has a small disc and five thick,

bluntly terminated arms. The rows of plates are:

carinal, two adradials, superomarginals, inferomarginals, and adambulacrals. Each plate bears a single large, robust, acute, movable spine mounted on

a mammiliform plate which is connected to the

adjoining plates by small secondary plates bearing

a small round patch of glassy tubercles. The large

papular areas bear numerous papulae, usually more

than twenty per area. All plates are covered with a

thick, heavily pigmented tegument. The adambulacral plates bear two slender spines within the

furrow, the innermost one the smallest, and a

larger, more robust, subacute spine on the furrow

margin. The mouth plates bear a preoral spine and

3 or 4 larger marginal spines that are webbed together at the base. The madreporite is small,

round, raised, and bears a double corona of small

spinules.

This may possibly be an aberrant E. brasiliensis,

from which it differs mainly in the number of

adambulacral furrow margin spines (two vs one),

and in the size of the abactinal spines, which are

very large and thornlike, in contrast to the small

conical spines of E. brasiliensis.

MATERIAL EXAMINED.—George M. Bowers Station

85 (1) [R=72 mm, r=15 mm. Rr=l:5].

Echinaster species B

PLATE 41: FIGURES C, D

This robust species has a moderate disc and five

stout, tapering arms. The plates are in regular

longitudinal rows: carinals (regularly alternating

left and right of the midline), adradials, superomarginals, inferomarginals, and adambulacrals. Between the adradials and the superomarginals, and

between the superomarginals and the inferomarginals are one or two incomplete rows of secondary

plates. All plates are small and connected to one

another by small secondary plates. The primary

plates bear a single small, straight, blunt spine distally, and proximally there is a minute patch of a

few glassy tubercles sunken in a small pit in the

plate. There are no spines on the actinal surface

of the disc. There are two small slender spines on

the adambulacral plates, deep within the furrow,

and on the furrow margin are two stout, blunt

spines of moderate length, slightly curved and

webbed together. There are three short, stout,

blunt spines of moderate length, slightly curved

and webbed together. There are three short, stout,

blunt spines on each narrow mouth plate, and also

a small suboral spinelet. The madreporite is round

and plane, with radiating gyri which probably bore

small spinules; however, the surface of the madreporite in this specimen is much eroded. The upturned oculars are small and broadly oval. The

covering tegument is thick.

In the regular arrangement of plates, this species

resembles E. serpentarius, but the shape of the

specimen is quite different, with a much larger disc

and thicker arm. Also, the size and ornamentation

of the plates differs, as well as the spinulation of

the adambulacral plates.

MATERIAL EXAMINED.—Silver Bay Station 48 (1)

[R=65 mm, r=23 mm, Rr=l:3].

VerrilUtster, new genus

TYPE-SPECIES.—Echinaster spinulosus Verrill,

1869.

DIAGNOSIS.—Single isolated spine on each abactinal plate; no patch of glassy tubercles; tegument

relatively thin, lifted in tentlike peaks by spines.

Verrillaster spinulosus (Verrill)

PLATE 43: FIGURES A, B

Echinaster spinulosus Verrill, 1869:386; 1915:40, pi. 4: figs.

1, 2.-Lutken, 1871:285 [61].-Ives. 1890:326.

This species has a moderate-sized disc and five

broadly based arms tapering to an acute tip. There

are more than six rows of plates between the carinal series and the adambulacrals at the base of the

arm. The rows of plates are: an irregular row of

carinals, an adradial, one or two dorsolaterals, superomarginals, and short intermarginal, inferomarginal, 1—3 actinolaterals, and adambulacrals. The

plates are small, lobate, imbricate, and without

glassy tubercles. Each carries a small, erect, acute

or blunt spine. The papular areas are numerous

and contain 1—6 papulae; they are scarce or lacking below the inferomarginals. The thin tegument

is lifted in tentlike peaks by the spines.
90 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The central anus, although small, is conspicuous

and surrounded by numerous small spinules. The

small, raised, round madreporite is covered with

coarse radiating gyri and surrounded by a corona

of small spinules. The very small oculars are upturned and more or less rounded. The spines of

the actinal surface are smaller and fewer than those

of the abactinal surface, many plates being entirely

without spines. The small adambulacral plates

bear one or two long, slender spines deep within

the furrow, and two larger, stouter, subacute spines

at the margin; on the actinal face are one or two

smaller spines. The small, narrow mouth plates

bear a stout preoral spine at the apex, 2-4 similar

but smaller spines on the side margin, and one or

two short suboral spines.

This is a description of one specimen from Oregon Station 5023. A second specimen from the

same station differs in having more numerous papulae in the papular areas, oculars completely obscured by spines, three rows of dorsolaterals, two

rows of intermarginals, and adambulacral marginal

spines with a broad, flattened tip. A specimen from

Oregon Station 3210-3216 has narrow arms, not

broad-based; very small spines, usually 3 or 4 per

plate; intermarginal plates confined to the disc; and

the actinal spines are a little larger than those of

the abactinal surface. Another specimen, from

Oregon Station 5934, would almost justify erecting

a separate subspecies, having roughly two carinal

rows of plates; the spines, particularly those of the

marginals, are much larger than those of the previous specimens, and the papulae per area are more

numerous. The spines of the ventral surface are

particularly long, numerous, and acute, and there

are no suboral spines on the mouth plates. A second, smaller specimen from the same station agrees

in every particular. A specimen without data is

similar to the second specimen from Oregon Station 5023.

Verrill gives the range of this species as west

coast of Florida and the northern Gulf of Mexico,

in 23-32 fathoms. The specimens here from the

Lesser Antilles thus represent a range extension,

both vertical and horizontal.

MATERIAL EXAMINED.—Oregon Stations: 5023 (2)

[R=65 mm, r=15 mm, Rr=l:4]; no station (1)

[R=60 mm, r=14 mm, Rr=l:4]; 3210-3216 (1)

[R=46 mm, r=9 mm, Rr=l:5]; 5934 (2) [R=47

mm, r=12 mm, Rr=l:4].

Order FORCIPULATIDA Perrier, 1884

In this order, pedicellariae, straight and/or crossed,

are almost always present. The dorsal skeleton is

reticulate, forming a more or less open meshwork.

The tube feet are suckered, and the ampullae single. Papulae occur on all surfaces. The plates are

never paxillate, and the spines occur singly, never

in groups or bundles. The marginals are inconspicuous and mostly irregular. The ambulacral and

adambulacral plates are crowded and compressed.

Only one family is represented here.

Family ASTERIIDAE Gray, 1840

This family is poorly represented in the Caribbean

and the Gulf of Mexico. This collection contains

only four genera and five species, all confined to

the northern Gulf of Mexico. Asteriids are characterized by having a more or less swollen body (i.e.,

neither actinal nor abactinal surface plane); five or

more arms; an open reticulate skeleton; numerous

crowded and compressed ambulacral and adambulacral plates; two or four rows of tube feet; and,

usually, numerous crossed and/or straight pedicellariae. The plates bear spines or spinules and are

never paxilliform.

Ampheraster Fisher

Ampheraster Fisher, 1923:253. [Type, by original designation,

Sporasterias mariana Ludwig, 1905.]

Pedicellariae unguiculate; no accessory inferomarginal spinelet; adambulacral spines predominantly

monacanthid; two enlarged interbrachial superomarginal plates overlapping two corresponding inferomarginal plates; superomarginals three- or

four-lobed; prominent inferomarginal spines, conspicuously larger than superomarginal spines; first

pair of postoral adambulacral plates narrowly separated or touch only at their adoral corners; ray$

five or six; species not fissiparous.

The five species previously assigned to this genus

by Fisher (1928b) are all from the eastern Pacific;

this is the first species in the genus reported from

the Atlantic. It seems to be most closely related

to A. hyperoncus (Clark), from Lower California,
NUMBER 126 91

Key to the Genera of Asteriidae

1. Adambulacral plates diplacanthid throughout, or alternately diplacanthid and monacanthid 2

Adambulacral plates monacanthid throughout 4

2. Arms usually five S

Arms 9-11 Coronaster

3. Disc small; arms long, narrow, angular Sclerasterias

Disc moderately large; arms stout, inflated Asterias

4. Inferomarginal plates with one spine, conspicuously larger than that of the superomarginals

Ampheraster

Inferomarginal plates with two spines, not conspicuously larger than that of the superomarginals Coscmasterias

but differs strikingly from this and other species in

the genus in several important respects.

Ampheraster alaminos Downey

PLATE 43: FIGURES C, D

Ampheraster alaminos Downey, 1971c:51-54.

This species has six arms, whereas other species in

the genus are regularly five armed. The skeleton

is weaker and more irregular than that of A. hyperoncus, and the arms are weakly attached to the

disc. On the dorsal surface, there is a large clear

patch of thin, uncalcified tissue between the base of

the arms and the disc, with, at most, one or two

small carinal plates between the arm and the disc.

In contrast, the enlarged interbrachial superomarginals, warped and bent and rather thin, form,

together with the heavy calcareous interbrachial

septa, six strong and rigid areas around the periphery of the disc. The superomarginal plates are generally four lobed, and imbricate in straight (not

zigzag) series. The upper lobe is usually reduced,

while the lower lobe is greatly elongated, overlapping the corresponding inferomarginal plate. The

small disc has a dorsum of thin tissue and a circlet

of elongate primary plates, with a few unattached

elongate plates in the center. Each plate bears a

few short, thorny, aciculate spines, and the madreporite, borne on one of the interradial primaries, is

of moderate size, covered with deep coarse gyri,

and bearing around its periphery a circlet of

spines.

Elsewhere the disc is covered with large straight

unguiculate pedicellariae. The surface of the arms

is completely covered with small crossed pedicellariae. The small carinal plates are more or less

irregular, and there are two series of meshes on

either side of the carinal row. The adambulacral

plates bear a single (occasionally two) long ridged,

aciculate spine, and the inferomarginal plates bear

a similar, somewhat shorter spine; above the inferomarginals, the superomarginals, dorsolaterals,

and carinals may bear a single, short spine, or no

spine. Straight unguiculate pedicellariae are abundant on the actinal disc interradii, and also occur

between the proximal adambulacral plates and the

inferomarginals. A few smaller pedicellariae of this

type occur within the groove. The long, narrow

mouth plates bear two long, slender, divergent

spines on the side of the straight bare apex, and

2 or 3 similar suboral spines, one behind the other.

The mouth plates also bear a number of small

unguiculate pedicellariae.

All of the specimens of this species are from

depths of 1,450 to 1,750 fathpms, in the north central Gulf of Mexico, where the bottom is mainly

foraminiferal.

MATERIAL EXAMINED.—Alaminos Stations: 3C/68-

A-7 (1 holotype) [R=62 mm, r=8 mm, Rr=l:7];

4E/68-A-7 (1, pluss 44 arms, 1 disc) [R=47 mm,

r=4 mm, Rr=l:ll]; 4A/68-A-7 (9) [R=42 mm,

r=4 mm, Rr=l:10]. Oregon Station 2547 (1) [R

=50 mm. r=5 mm. Rr=l:10].

Sclerasterias Perrier

Sclerasterias Perrier, 1891b:1227. [Type, by original designation, 5. guernei Perrier.]

Eustolasterias Fisher, 1923:255. [Type, by original designation,

Coscmasterias (Distolasterias) euplecta Fisher.]
92 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Dorsal skeleton regular, with two series of skeletal

meshes on either side of prominent carinal ridge;

inferomarginal plates with two spines, only outer

one carrying cluster of crossed pedicellariae; adambulacral plates diplacanthid throughout; straight

pedicellariae lanceolate; outer inferomarginal

spines webbed together at base; alternate superomarginals usually spineless; carinal, marginal, and

actinal plates in regular longitudinal series; small

obtuse spines above inferomarginals wreathed by

collar of small crossed pedicellariae; adults regularly five-armed, but young may be six-armed and

fissiparous.

Only one species occurs in the area covered by

this report. S. tanneri (Verrill), a more nothern

species, is generally much smaller and bears a spine

on every carinal plate, rather than every other one.

S. subangulosa (Verrill) (=Orthasterias (Stylasterias) subangulosa Vemll=Asterias angulosa Perrier) was rightly considered the young of S. contorta

by Fisher (1928a).

Sclerasterias contorta (Perrier)

PLATE 44: FIGURES A, B

Asterias contorta Perrier, 1881a: 1.

Asterias angulosa Perrier [non Muller], 1881a:3; 1884:202.

Orthasterias contorta .-Verrill, 1914:48; 1915:18.

Orthasterias (Stylasterias) subangulosa Verrill, 1914:168, 370.

Orthasterias subangulosa.—Verrill, 1915:16, pi. 2: figs. 1, 2, pi.

9: figs. 1, la.

Eustolasterias subangulosa.—Fisher, 1923:255.

Eustolasterias contorta.—Fisher, 1923:255.

Sclerasterias subangulosa.—Fisher, 1928a: 107, 108.

Sclerasterias contorta.—Fisher, 1928a: 107, 108.

This species has a small disc and five long, narrow

arms, angular and straight sided, tapering to an

acute point. The disc dorsum has an irregular

pentagon of plates, each bearing one or two spines.

The carinal plates on the arm form a prominent

ridge; they are lobed and bear a single, large, aciculate spine surrounded by a dense wreath of small,

blunt, crossed pedicellariae. They connect by a

series of elongate ossicles to the prominent superomarginal plates. There is an incomplete series of

dorsolaterals, many without spines. Every other

superomarginal bears a spine similar to those of

the carinals on the abactinal portion of the plate,

and each plate may bear, on the abactinal portion,

a large round patch of low glassy tubercles. These

usually tend to become fewer and smaller with

growth. A long lobe on the superomarginal connects it to the corresponding inferomarginal, leaving a wide, bare space, devoid of spines or other

ornamentation, between the two series of plates.

The inleromarginal plates bear two spines; those

of the outer (or abactinal) series are webbed together and bear, on the abactinal side, a dense

patch of pedicellariae similar to those wreathing

the upper spines. The inner (or actinal) series of

inferomarginal spines are free and without pedicellariae. There is a single row of broad, low actinal

plates between the inferomarginals and the adambulacral plates. Between the actinals is a single

papula; elsewhere, above the inferomarginals, the

papulae occur usually in pairs between the plates,

but there may be as many as 5 or 6 in each area.

The adambulacral plates bear two long, slender,

divergent spines which form a double row down

each side of the ambulacral groove. The groove

is wide, and the tube feet are in four rows. The

small, narrow mouth plates project into the peristome and bear an apical, forward-directed spine and

a smaller oblique marginal spine on each half;

there are two sets of suboral spines, the first of

one spine and the next of two. Large, isolated, lanceolate pedicellariae occur here and there on both

surfaces, mostly on or near the disc. The madreporite is of moderate size, irregularly round, and

plane to concave.

This species differs from S. tanneri, which occurs

from Cape Hatteras north, in having a more prominent carinal ridge, an incomplete row of dorsolaterals, fewer large lanceolate pedicellariae, and

one or two spines on the dorsal disc plates, instead

of a number of small spines, as well as in having

angular, straight-sided arms, pentagonal in cross

section (the arms of S. tanneri are almost round

in cross section).

This species has previously been recorded from

Florida to Barbados in 11—188 fathoms.

MATERIAL EXAMINED.—Oregon Station 4731 (1)

[R=90 mm, r=9 mm, Rr=l:10].

Asterias Linnaeus

Asterias Linnaeus, 1758:661. [Type, by subsequent designation, A. rubens Linnaeus (Norman, 1865:126).]

Stellonia Nardo, 1834:716. [Type, by subsequent designation,

Asterias rubens Linnaeus (Fisher, 1913:639).]
NUMBER 126 93

Uraster L. Agassiz, 1835a: 191. [Nomen nov. for Stellonia Nardo.]

Asteracanthion Muller and Troschel, 1840:102. [Type, by original designation, Asterias rubens Linnaeus.]

Allasterias Verrill, 1909:65. [Type, by original designation,

A. rathbuni Verrill.]

Parasterias Verrill, 1914:53. [Type, by original designation,

P. albertensis Verrill.]

Large disc; five or six broad, rather short arms;

aboral skeleton forms an open network, with spines

in nearly regular series on carinal and marginal

plates, but irregular elsewhere.

Only one species is known from the Gulf and

Caribbean.

Asterias forbesi (Desor)

PLATE 44: FIGURES C, D

Asteracanthion forbesi Desor, 1848:67.

Asterias arenicola Stimpson, 1851:268.

Asteracanthion berylinus A. Agassiz, 1863:3; 1877:94, pi. 9.

Asterias forbesi.—Verrill, 1866:345.-H. L. Clark, 1902:552, pi.

1: ags. 1, 2, pi. 4: figs. 14, 15.-Fisher, 1930:205.-Gray,

Downey, and Cerame-Vivas, 1968:156, fig. 34.

Asteracanthion novae boracensis Perrier, 1869:41, pi. 1: fig. 9a.

This is probably the commonest shallow-water

species on the east coast of North America; certainly, it is the best known, for it is the one

commonly used in biology classes. The disc is

small, and the five arms are moderately long, thick,

and constricted at the base. There is an irregular

carinal row of plates down the middle of each arm.

The superomarginal plates define the ambitus and

are connected to the inferomarginals by an elongate

ossicle; the inferomarginal plates are connected to

3 or 4 adambulacral plates by a triangular, unarmed

ossicle. All plates from the carinals to the inferomarginals are connected by small, elongate secondary ossicles, forming an open meshwork, and bear

a single, thick, upright, thorny-tipped spine, the

spines of the superomarginals being slightly larger

than those of the other plates. The spines are all

surrounded, about halfway up, by a wreath of

small, crossed pedicellariae. The inferomarginals

bear two thicker, blunter spines. The small,

crowded adambulacral plates bear either one or

two spines, alternately; they are usually chisel

shaped and often grooved. Between the spines, over

the entire surface, and on many of the adambulacral

spines are straight, flat, oval or cordiform pedicellariae, larger than the crossed pedicellariae which

wreath the spines. The papulae between the plates

are small and numerous.

The long, narrow mouth plates consist of 3—7

adambulacral plates which meet across the interradius; each bears a long, straight, slender, somewhat flattened spine. The madreporite is large, irregularly round, raised, and wartlike, covered with

many fine gyri.

This species ranges from Maine to the Gulf of

Mexico, in shallow water. It is the southern form

of Asterias vulgaris, which ranges from Labrador to

the Carolinas (and, rarely, to Florida). It is sometimes difficult for even a specialist to distinguish

these two species, and hybridization is not uncommon, particularly in the Cape Cod area. This

is one of the many cases among marine invertebrates which compels us to realize how very

unsatisfactory are any of the proposed definitions

of "species."

MATERIAL EXAMINED.—Oregon Stations: 5894 (1)

[R=:83 mm, r=21 mm, Rr=l:4]; 6300 (12) [R=45

mm, r=ll mm, Rr=l:4]; 5882 (1) [R=22 mm,

r=6 mm, Rr=l:4]. Silver Bay Stations: 1564 (1)

[R=20 mm, r=5 mm, Rr=l:4]; 1710 (1, with four

arms) [R=42 mm, r=10 mm, Rr=l:4]; 1952 (4,

two with four arms) [R=80 mm, r=18, Rr=l:4].

Coscinasterias Verrill

Cosinasterias Verrill, 1867:248. [Type, by original designation,

C. muricata Verrill.]

Stolasterias (subgenus) Sladen, 1889:583. [Type, by subsequent

designation, Asterias tenuispina Lamarck, 1816 (Fisher,

1923:128).]

Lytaster Perrier, 1894:09. [Type, by original designation, L.

inaequalis Perrier ( = Asterias tenuispina Lamarck, 1816).]

Polyasterias Perrier, 1894:108. [Type, by original designation,

Asterias tenuispina Lamarck, 1816.]

Disc small; variable number (5-12) of moderately

long, angular rays; single series of spiniferous

actinal plates; adambulacral plates monacanthid

throughout; only outer of two inferomarginal

spines carries cluster of crossed pedicellariae; fissiparous.

Only one species is known from the western

Atlantic.

Coscinasterias tenuispina (Lamarck)

PLATE 45: FIGURES A, B

Asterias tenuispina Lamarck, 1816:561.—Perrier, 1875c:306.
94 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Asteracanthion tenuispinus.—Muller and Troschel, 1842:16,

pi. 1: fig. 1.—Perrier. 1869:32, pi. 1: figs. 3-3c—Lutken,

1859:95.

Asterias (Stolasterias) /enuispina.—Sladen, 1889:565, 583.-

Verrill, 1907:324 [280], pi. 34: fig. 2.

Coscinasterias tenuispina.—Verrill, 1914:45; 1915:19, pi. 26:

fig. 2, pi. 27: fig. 4.—Gray, Downey, and Cerame-Vivas, 1968:

159, fig. 38.

Asterias atlantica [part] Verrill, 1868:368.-Rathbun, 1879:144.

This species has a small disc and a variable number (5—10, usually seven) of long, angular arms of

unequal length. It is fissiparous. The dorsal plates

are thick and large and generally four lobed, forming an open meshwork. The carinal row is well

marked, and most of the carinal plates bear a long,

strong, movable, aciculate spine. The specimen

before me is a juvenile, and has only a few dorsolateral plates, but larger specimens usually have one

or two regular rows of dorsolaterals, armed like

the carinal plates. The superomarginal plates bear

an irregular patch of fine glassy tubercles, and alternate plates bear a sturdy aciculate spine, longer

than that of the dorsal plates. The inferomarginals

are trilobate and bear two oblique spines similar

to those of the superomarginals. This juvenile

specimen has no interactinal plates, but larger

specimens usually have a single row of unarmed

interactinals. The small, crowded adambulacral

plates bear a single, long, slender spine. All spines

above the inferomarginals bear a dense wreath of

small, crossed pedicellariae; this juvenile specimen

has only a few small, straight pedicellariae within

the groove, but in larger specimens these are frequently large and abundant on all surfaces. The

adambulacral spines and the inner, or actinal, inferomarginal spines are without pedicellariae. The

disc dorsum is plated with large, heavy, irregularly

shaped plates, most bearing a single large spine.

On the specimen before me, there are six round,

raised madreporites, covered with coarse gyri.

This tropical-subtropical amphi-Atlantic species

is common in the Mediterranean, and has been

found at most of the islands in the Atlantic from

Bermuda to St. Helena. Its depth range is unknown,

but is quite possibly very great.

MATERIAL EXAMINED.—Silver Bay Station 2010 (])

[R=30 mm, r=4 mm, Rr= 1:7.5].

Coronaster Perrier

CoronaUer Perrier, 1885c: 13. [Type, by original designation,

C. parfaiti Perrier.]

Heterasterias Verrill, 1914:46. [Type, by original designation,

Asterias (Stolasterias) volsellata Sladen.]

Rays 9-11; prominent, well-spaced acicular abactinal and marginal spines; large, straight, spatulate,

unguiculate pedicellariae; abactinal skeleton with

large square meshes; conspicuous wreath of small,

crossed pedicellariae around spines; abactinal

skeleton much reduced.

Only one species is known from the western

Atlantic.

Coronaster briareus (Verrill)

PLATE 45: FIGURES C, D

Asterias briareus Verrill, 1882:220; 1884:659; 1895:209.

cf. "Asterias volsellata".—Nutting, 1895:168, fig. 3.

Coronaster briareus Verrill, 1915:31, pi. 1: figs. 1, 2, pi. 9:

figs. 4-4c.-Gray, Downey, and Cerame-Vivas, 1968:158, fig.

37.

This species has a small disc and eleven (9—12)

long, slender arms. The disc is well covered with

large, irregularly shaped and arranged plates, most

of them bearing one or two long aciculate spines

encircled by a dense wreath of small, crossed pedicellariae.

Elsewhere on the disc are large, isolated, straight,

spatulate unguiculate pedicellariae. The madreporite is small, round, smoothly convex, and covered with deep gyri. The arms are high and inflated

near the base, and the skeleton is reduced to a

carinal row of lobate plates connected by long,

slender, transverse ossicles to similar superomarginals, which in turn are connected by elongate

lobes to trilobate inferomarginals. The inferomarginal plates are connected to one another by imbricate secondary plates. Most of the carinals,

superomarginals, and inferomarginals, and a few

of the secondary connective plates bear a long,

slender, aciculate spine with a dense wreath of

small, crossed pedicellariae encircling it. Between

the narrow plates, the meshes are large, square, and

contain a few papulae. The adambulacral plates

bear two long, slender, divergent spines without

pedicellariae. Within the groove, and also elsewhere

on the surface, are large, solitary, straight unguiculate pedicellariae, expanded and spatulate near the

tip, like little clasped hands. Smaller straight pedicellariae are also present. The large tube feet are
NUMBER 126 95

in two rows. The mouth plates bear a large acicular

oral spine and a smaller marginal spine, as well as

a number of small, straight pedicellariae. Behind

the mouth plates, two or three adambulacral plates

are united across the interradius.

The peristome is wide open, and there is a rather

broad peristomial membrane. Within the peristome

of the specimen before me were two tiny crabs of

the family Calapidae.

The known distribution of this species is from

New Jersey to the Florida Keys, in 31-373 fathoms.

MATERIAL EXAMINED.—Alaminos Station 20/65-

A-9 (1) [R=125 mm. r=7 mm, Rr=l:1.3].

Order ZOROCALLIDA Downey, 1970b

In this order, the disc is more or less domed, with

a regular arrangement of enlarged primary plates.

There is a prominent median carina on the arms,

and all arm plates are in compact and imbricating

longitudinal and transverse series. The mouth frame

is deeply sunken in the actinostome. The adambulacral plates are alternating carinate and noncarinate, and at least some of the tube feet are

large and conical, with a tiny suckered disc. There

is but one living family in the order.

Family ZOROASTERIDAE Sladen, 1889

The disc is small, and the five arms are long,

slender, and tapering. The dorsal disc plates are

enlarged, in a regular arrangement of primary

radials, interradials, centrodorsal, and sometimes

additional plates. There is a prominent row of

carinal plates along the midarm, and the other

arm plates are in regular longitudinal and transverse series; in most specimens, the number of rows

decreases from base to arm tip. The adambulacral

plates are alternately carinate and noncarinate.

Superambulacral plates are present in all zoroasterids, although they may be present only proximally in most genera, and in Zoroaster they are

greatly reduced. Large, duck-billed pedicellariae

are present in most genera, but may be absent in

Mammaster. Small, straight pedicellariae are

numerous.

The tube feet are in four rows proximally, two

distally; within the peristomial cavity, they are

straight, with a large suckered disc, but on the

arms they become stout and conical, with a very

small suckered disc.

Key to the Genera of Zoroasteridae

1. Disc plates conspicuously enlarged, more or less convex, bare, or at least not covered with

spinules; disc and upper part of arms skin-covered 2

Disc plates plane or, if convex, closely covered with spinules; no skin-covering on dorsal

surface Xoroasttr

2. Disc plates distinctly stellate, not much swollen Doraster

Disc plates decidedly swollen, more round than stellate Mammaster

Doraster Downey

Doraster Downey, 1970b:5. [Type, by monotypy, D. constellatus Downey.]

Doraster constellatus Downey

PLATE 46: FIGURES A, B

Doraster constellatus Downey, 1970b:5-12, figs. 2c, 3-11.

The plates of the disc dorsum are large, flat, stellate,

smooth, and skin covered. The central plate usually

bears a stubby tubercle and a similar tubercle sometimes occurs on other primary disc plates. The

anus, surrounded by small spinelets, is located in

one of the angles of the central plate. Surrounding

the centrodorsum are five primary interradial

plates, and between and underneath them are five

radial plates on which the five pointed lobes of the

centrodorsum rest; just outside and partly covering

the primary interradials are five large primary

radials; a pair of interradial plates, entirely concealed by skin and the overlapping primary radials,

lie between and beneath the primary radials and

four enlarged imbricating (in large specimens,

fused) adradials that occupy the interradial angle.

Small pedicellariae and tiny granuliform platelets

occur between the plates of the disc. The madreporite is slightly smaller than the disc plates,
96 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

raised, covered with irregular channels, and located

between a primary interradial and the enlarged interradial adradials; it actually rests on a pair of

concealed interradials.

The first carinal is enlarged, about the same

size as the interradial adradials. The jaw is one

solid piece (made up of two pairs of adambulacral

plates), projecting into the peristome. Behind it

are four pairs of adambulacrals joined across the

interradius by a tooth-and-socket arrangement

reminiscent of the way the ambulacral plates are

joined at the top of the ambulacral groove. Internally, the front of the jaw is flaired like the

carinate adambulacral plates and bears, on each

side, three subequal spines completely covered with

tiny, straight pedicellariae, which thus form a

wreath around the peristome. External to these

three pairs of oral spines is a single pair of long,

sharp spines. The next two pairs of adambulacral

plates behind the jaw each bear a single pair of

long sharp spines, and the two pairs behind them

bear 4—6 similar spines. The seventh and eighth

pairs of interradial adambulacrals are separated

by a large duck-billed pedicellaria. The first two

pairs of ambulacral plates are also strongly fused

and greatly swollen. The ambulacrals curve sharply

upward under the disc and thus support the disc

in a domed position. The two fused pairs are

buttressed against the body wall by a solid ridge

formed of the first two (enlarged) superambulacral plates. Superambulacral plates of two kinds

occur, alternately, but do not continue to end

of the arm; rather, they diminish in size and

eventually disappear. A straight, triangular superambulacral plate alternates with a thin, flaired

superambulacral, which terminates in a broad,

flat disc resting on the ambulacral plates.

They gradually become similar, small, and granuliform, and do not continue much beyond the

middle of the arm. The tube feet are in four rows

proximally, two distally. Around the mouth, they

are straight and conspicuously annulated and

terminate in a large suckered disc, but beyond the

confines of the disc, they are stout and conical and

taper to a tiny disc. On the arms, the carinal plates

and two rows of plates on either side (adradials

and superomarginals) are covered with skin, devoid

of spines or spinules, but may have small pedicellariae, especially proximally. The carinals may have

a central, nonspine-bearing tubercle. Adradial

plates overlap carinals and superomarginals. The

carinals and adradials are about equal in size,

somewhat cruciform, wider than long; the superomarginals are slightly smaller, lobed, or, in large

specimens, somewhat triangular, tapering toward

the actinal surface. On large specimens, an irregular

row of tiny granuliform platelets occurs between

the carinals and the adradials; a few also occur

sporadically on the disc. Proximally, there are five

rows of plates between the superomarginals and

the adambulacrals, covered with small sacculate

spinules; the row adjoining the superomarginals

is the inferomarginal row. About halfway out on

the arm, one row of plates drops out, then farther

out another, until at the end of the arm only the

carinals, very small adradials, superomarginals, and

inferomarginals remain.

The row of plates adjoining the adambulacrals

is very narrow, longer than broad, and only the

edge of the plate may be visible. Most of the

actinolaterals carry a long, flattened, appressed

spine, directed upward and attached to a prominent

tubercle. All of these arm plates are in regular

longitudinal and transverse rows; however, the

latter do not correspond to the adambulacrals.

Papulae occur singly or in groups of up to five

between the plates; none occur below the inferomarginals. The terminal ocular plates are cordiform, covering the arm tip, with three or more

coarse terminal spines. Adambulacral plates are

of two alternating sorts, carinate and noncarinate;

the carinate adambulacrals project strongly into

the ambulacral groove and bear four movable

spines on a transverse row of tubercles. The spines

are stout and rounded at the point of attachment,

tapering rapidly to a point; they may be curved.

The first or second spine usually bears a large

duck-billed pedicellaria, and there are numerous

small, straight pedicellariae. The noncarinate

adambulacrals bear 3-5 small spinelets on their

outer surface and do not project into the furrow;

they may also bear small, straight pedicellariae.

This species seems to be limited to the northern

Gulf of Mexico, in 190-350 fathoms.

MATERIAL EXAMINED.—Oregon Stations: 3583 (1)

[R=80 mm, r=12 mm, Rr=l:6.6]; 382 (6) [R=

122 mm, r=12 mm, Rr=l:10]; 2780 (2) [R=239

mm, r=38 mm, Rr=l:7]. Alaminos Station 21/68-
NUMBER 126 97

A-13 (1) [R=121 mm, r=15mm, Rr=l:9]. Oregon

II Station 10602 (1) [R=183 mm, r=23 mm.

Rr=l:9].

Mammaster Perrier

Mammaster Perrier, 1894:125. [Type, by monotypy, Zoroaster

sigsbeei Perrier.]

Mammaster sigsbeei (Perrier)

PLATE 46: FIGURES C, D

Zoraster sigsbeei Perrier, 1881a:5.-Sladen, 1889:417, 418, 423,

790.

Mammaster sigsbeei.-Pertier, 1894:125. H. L. Clark, 1941:

67.-Downey, 1970b: 12.

The arrangement of plates on the disc is definite

and unmistakable: a centrodorsal plate, five interradial plates, then five larger radial plates. In each

interradial arc are two large somewhat crowdedlooking plates, and over the base of each arm are

three large, tumid plates. All these plates are

raised, tumid, round, and somewhat bare (although

they may have small spinelets around the edge).

The madreporite is small and inserted between

the two interradial arc plates and the interradial

primary plates. The jaws bear three transverse

rows of 2, 2, and 3 or 4 long, acute spines, and

inside the mouth, not visible without dissection,

are two more pairs, the central pair short and

blunt, and the outer pair curved away from the

jaws like cow's horns, both pairs covered with

small pedicellariae. Both small and large straight

pedicellariae are present in the ambulacral groove

and on the dorsal surface, especially on the disc.

The number of rows of tube feet diminishes

from four to two about half to three quarters of

the way down the arm. Only the carinal plates

and one row of plates on each side extend all the

way to the end of the arm. The terminal plate is

as broad as long, with an indentation on the proximal side. The carinals are overlapped on each

side by the plates of the next adjoining row. These

three rows of plates are comparatively bare, with

only a few spinules on the distal edges of the

plates, and are probably covered by a thin skin.

The next row of plates is covered with tiny spinules,

is overlapped by the adradials, and in turn overlaps

the similar row of plates below it. Both of these

rows diminish and disappear before reaching the

end of the arm, the lower row first, then the upper.

Next come two rows of narrow elongate plates

which disappear distally before the rows above.

Each of the plates in these four spinose rows bears

one larger spine on a tubercle. The adambulacral

plates are alternately carinate and noncarinate,

with every other one projecting strongly into the

furrow and bearing four stout, movable spines in a

transverse row. These spines are frequently curved

and the first or second often bears one or two large

pedicellariae. The adambulacral plates between do

not project into the furrow and have no furrow

spines, but bear two to four somewhat flattened

spines distally, side by side in pairs. The difference

between the alternating adambulacrals becomes

less distinct distally.

JUVENILE.—A specimen from Combat Station 450

has a major radius of 9 mm. The centrodorsal

plate is by far the largest plate and bears a blunt

tubercle. There are only two rows of plates on the

arms beyond the carinals proximally, and only

one row distally. There is no appreciable alternation of the adambulacral plates. There are only

two rows of tube feet the whole length of the arm.

The primary plates of the disc, except for the

centrodorsal plate, are not appreciably raised and

tumid. There are no pedicellariae.

This species apparently occurs throughout the

West Indies, from Key West to Trinidad, in 195—

350 fathoms.

MATERIAL EXAMINED.—Combat Station 450 (10)

[R=47 mm, r=8 mm, Rr=l:6]; juvenile, [R=9

mm, r=2 mm, Rr= 1:4.5]. Oregon Stations: 2775

(1) [R=29 mm, r=5 mm, Rr=l:6]; station unknown, Florida Keys, 200 fms (5) [R=60 mm. r=9

mm, Rr=l:6].

Zoroaster Thomson

Zoroaster Thomson, 187S: 154. [Typ*e, by original designation,

Z. fulgens Thomson.]

All of the Zoroasters in the western Atlantic appear

to belong to one species (see Downey, 1970b).

Zoroaster fulgens Thomson

PLATE 47: FIGURES A, B

Zoroaster fulgens Thomson, 1873:154— Studer, 1884:2^

den, 1889:694.-Bell, 1893:88.-Perrier, 1894:116.-Verrill.
98 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1895:205.-Koehler, 1895:442; 1896:42.-Chun, 1900:489, fig.

- H. L. Clark, 1901:237.-Nichols, 1903:251.-Schmidt, 1905:

22.-Grieg, 19O7a:46; 1932:24.-Downey, 1970b: 15.

Zoroaster ackleyi Perrier, 1880:436; 1881b:59; 1881a:272;

1881b: 1; 1894:117.-Verrill, 1895:205.-H. L. Clark, 1901:237.

-Fisher, 1928a:491.

Non Zoroaster ackleyi.—Wood-Mason and Alcock, 1891:14.

Zoroaster trispinosus Koehler, 1895:442; 1896:42, figs. 14, 15.

Zoroaster bispinosus Koehler, 1909b: 136.

A five-lobed centrodorsum, eight primary radials

(in overlapping pairs), and five large interradials,

plus five large first carinals, make up the disc

dorsum. The anus is between the centrodorsum

and two primary radials, which are single rather

than being paired like the other six. The anal pore

is surrounded by small, flattened spines. Small

pedicellariae may be scattered over the disc. The

madreporite is small, flat, channeled, and located

just distal to a primary interradial. Each carinal

plate on the arms has two lobes on each side and

overlaps the carinal plate proximal to it. Transversely, the carinals overlap the adradials, and the

superomarginals overlap both the adradials and

the inferomarginals. Below the inferomarginals

are 6—2 rows of actinolateral plates (six near the

disc, becoming two by the end of the arm). Each

actinolateral plate bears a long, slender, appressed

spine, directed upward, and mounted on a horseshoe-shaped tubercle, plus numerous small sacculate spinelets. The inferomarginals are similarly

armed. The superomarginals and carinals frequently (but not always) bear a stout, erect central

spine or tubercle, and these plates, as well as the

adradials, are covered with small spinelets. The

carinals are broader than long proximally and

longer than broad distally.

The superambulacral plates are very small and

do not extend beyond about the middle of the arm.

The tube feet are in four rows proximally, becoming two rows less than halfway out on the arm.

They are stout, conical, terminating in a small

suckered disc (except for those actually within the

peristomial cavity, which have large suckered discs),

and the ampulae are double. The adambulacral

plates are alternately carinate and noncarinate;

the carinate adambulacrals bear five spines, two

long, slender furrow spines with many small,

straight pedicellariae (a large duck-billed pedicellaria may replace the many small ones) and three

outer spines which are similar to the three spines

of the noncarinate adambulacrals; these small

spines do not bear pedicellariae. The jaw of four

fused adambulacral plates bears two short, stout

oral spines on each side, in a row along the oral

edge of the jaw, covered with small, straight pedicellariae, and behind them are two pairs of long,

acute spines.

This species seems to occur all over the North

Atlantic, in 200-2,000 fathoms.

MATERIAL EXAMINED.—Alaminos Stations: 3C/68-

A-7 (1) [R=54 mm, r=7 mm, Rr=1.8]; 1/68-A-13

(2) [R=86 mm, r=8 mm, Rr=l.ll]; 10B/68-A-3

(2) [R=85 mm, r=6 mm, Rr=l:14]. Oregon Stations: 1537 (2) [R=75 mm, r=9 mm, Rr=l:8];

2574 (4) [R=85 mm, r=7 mm, Rr=l:12]; 3561 (1)

[R=200 mm (est.), r=21 mm, Rr=l:10]; 3560 (2)

[R=210 mm, r=18 mm, Rr=l:ll]; 5929 (5) [R=

149 mm, r=14 mm, Rr=l:10]; 4413 (3) [R=177

mm, r=22 mm, Rr=l:8]; 2571 (1); 2202 (2);

2779 (1); 2824 (2); 2652 (10); 1538 (6); 382 (6);

2650 (5); 3654 (6) [R=71 mm, r=7 mm, Rr=l:10].

Oregon II Station 10619 (1) [R=75 mm, r=7 mm,

Rr=l:10].

Order EUCLASTERIDA Tortonese, 1958

With the characters of the family Brisingidae; the

only family in the order is Brisingidae Sars, 1875.

Family BRISINGIDAE Sars, 1875

The Brisingidae have many deciduous arms, sharply

distinct from the small circular disc. The vertebralike ambulacral and adambulacral plates are not

compressed, and the tube feet are biserial. The

dorsal skeleton, confined to the proximal third of

the arm, is weak and never reticulate. The abundant pedicellariae are crossed, never straight. The

mouth plates are expanded, and the actinostome

is large, with a broad peristomial membrane. Papulae are lacking or confined to the disc and/or

genital region of the arms. The marginal and

adambulacral spines are long, delicate, acicular, and

ensacculate.

Key to the Genera of Brisingidae

Papulae numerous on disc and genital region of arms Odinia

Papulae limited to one pair on each radial area of disc, near disc margin Midgardia
NUMBER 126 99

Odinia Perrier

Odinia Perrier, 1885c:9. [Type, by original designation, O.

semicoronata Perrier.]

Numerous papulae on disc and genital region of

arms; mouth plates broad and fan shaped toward

actinostome, nearly closing entrance to ambulacral

furrow; genital region of arms with transverse

skeletal arches; tegument between costae contains

numerous immersed plates and papular pores; arms

with regularly spaced transverse combs of slender

lateral spines; adambulacral plates, higher than

long, bear single prominent, frequently truncate,

subambulacral spine; first 3-5 adambulacral plates

united interradially, as are marginals above them.

Only one species has been reported from the

Caribbean and Gulf of Mexico.

Odinia antillensis Clark

PLATE 47: FIGURES C. D

Odinia antillensis A. H. Clark, 1934:1, pi. 1.

This species has a small disc with a plane, abactinal

surface and high, abrupt sides sloping sharply to

the base of 15-17 long, thin arms. The surface of

the disc is covered with small polygonal plates

bearing 2—16 short spinelets. In the interradial

areas of the disc, the plates form a solid bare

channel which extends outward between the fused

bases of the arms for a short distance. The arms

are constricted at the base, and about the first 5—10

adambulacral and marginal plates are closely

joined to those of the adjoining arms to form the

bare channels mentioned above. Just beyond the

constricted and partly fused basal area, the arms

swell abruptly in the high and inflated genital area.

This area, which tapers gradually distally, is about

a quarter of the arm length or less. The genital

region is plated with small polygonal plates, most

bearing one or two small spinelets. The costae on

the proximal half of the genital region are irregular and scarcely distinguishable from the general

plating; they correspond to every third adambulacral plate and are mainly distinguishable by the

band of large plates bearing single large spines

across the arm.

On the distal half of the genital region, the

costae are distinct and form conspicuous bands or

ridges across the arm. Beyond the genital region,

the costae are reduced to 3 or 4 plates just above

every fourth adambulacral plate, each bearing a

single long spine, and the dorsal portion of the arm

is covered by a thin plateless tegument through

which the ambulacral plates are clearly visible.

Papulae are numerous between the plates of the

disc and the genital region of the arms. The adambulacral plates are cylindrical and each bears a

prominent furrow spine; proximally, these are very

long and heavy, and the tip is expanded and may

be forked or chisel shaped. Beyond the genital

region, the adambulacral spines are long and

acicular. The mouth plates are large and closely

connected both radially and interradially, forming

a nearly closed ring around the broad peristome.

They bear about five very short spines. Small,

crossed pedicellariae are abundant on all surfaces

and cover the spines. The small, round madreporite is on the sloping side of the disc at the edge

of the abactinal surface.

This species was previously known only from

off Puerto Rico.

MATERIAL EXAMINED.—Oregon Stations: 4480 (1)

1408 (1) [R=304 mm (est.), r=10 mm, Rr=l:30];

4574 (2) [R=100 mm, r=5 mm, Rr=l:20].

Midgardia Downey

Midgardia Downey, 1971d:421-426. [Type, by monotypy, M.

xandaros Downey.]

Midgardia xandaros Downey

PLATE 48: FIGURES A, B, C

Midgardia xandaros Downey, 1971d:

This species has a small, circular, plane disc and

twelve very long, attenuate arms. The disc dorsum

is covered with a thin membrane in which are

immersed many small, irregular plates bearing one

or two tiny spinules. In each radial area, near the

edge of the disc dorsum, is a pair of small papulae;

there are no other papulae. The madreporite is

large, situated on the edge of the disc, forming a

stout, heavily calcified cone with few gyri in a

pit on top. At the periphery of the disc, the first

pair of ambulacral plates are united by a syzygy,

and the oval interbrachial adambulacral plates

between are separated by the intrusion from above

of a pair of partially fused marginals. On the
100 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ventral surface, the actinostome is wide (1 cm

radius), and the expanded mouth plates form a complete wall around the inside of the peristome;

each mouth plate bears 2 or 3 small, fine, acicular

spines on the apical margin. The peristomial membrane is rather small and fragile. No pedicellariae

were noted on the disc. The arms, constricted at

the base, swell rapidly in the genital region, which

constitutes less than one fifth of the total arm

length. The largest diameter of the arm, about 1

cm, is in this area.

The costae of the slightly inflated genital region

are more or less irregular and crowded near the

base of the arm, becoming less crowded and more

distinct distally. They are made up of elongate

overlapping plates, most of which bear a tiny

aciculate spine with an expanded bulbous base.

The costal plates become smaller and fewer beyond

the middle of the genital region, and beyond this

region are represented only by bands of pedicellariae; even these disappear at about midarm, and

the distal half of the arm is covered only by a thin

tegument. The tegument between the costae is

filled with immersed platelets, thin, spineless,

fenestrated, and somewhat imbricate. The arms

become extremely attenuate at the tip, terminating

in a small, broader than long, down-curved plate

(less than 0.4 mm wide), well-armed on the actinal

edge with small spines, so that it looks like a tiny

cat's paw with claws extended. Laterally, on the

arm, the first few marginals are fused together to

form a smooth channel between the bases of the

arms. Beyond the arm base, the small inferomarginal plates are irregularly spaced, averaging one

about every third or fourth adambulacral plate. In

the genital region, the costae arise from these inferomarginals. Each inferomarginal plate bears a single

very long, slender, acicular spine with a flap of

tissue at the tip. Beyond the genital region these

spines increase in length, becoming as long at 2.5

cm beyond the middle of the arm.

On the actinal side of the arms, the somewhat

cylindrical adambulacral plates bear a single long,

slender spine on the actinal face, slightly shorter

than the inferomarginal spine, and within the furrow they bear two pairs of tiny, fine spines that

project toward the middle of the furrow. The very

long suckered tube feet are biserial and stiffened

at the base by a calcareous collar; they rest in large

basins between the adambulacral plates. In dorsal

aspect, the ambulacral pores are covered with a

thin membrane which appears to be only slightly

expandable; indeed, it can hardly be called an

ampulla. The tube feet were very long and agile

in the live specimen and therefore probably had

limited ability to contract. Sars (1875), in his study

of the structure of the related genus Brisinga,

noted that the tube feet were not so highly contractile as in other (families of) starfishes. The

gonads appear to be united in a single strand on

each side of the arm, but it was not possible to

determine whether the gonoducts were single or

double. Tiny hyaline pedicellariae cover the costae

and clothe the spines of the arms to their tips.

MATERIAL EXAMINED.—A laminos Station 58/69-

A-ll (1) [R=680 mm, r=13 mm, Rr=l:60].

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Sluiter, C. P.

1895. Die Asteriden Sammlung des Museums zu Amster-
NUMBER 126 109

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1951. Early Palaeozoic Starfish. Philosophical Transactions of the Royal Society of London, series B, 235

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10 figures, 1 plate.

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Ulrey, A. B.

1918. The Starfishes of Southern California. Bulletin of

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1963. Asteroids from the Netherlands Antilles and Other

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1959. Coloured Illustrations of Seashore Animals of Japan. xvii-f-227 pages, volume 167, 64 colored plates,

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pages 105-122, placts 53-61.]

Verrill, A. E.

1866. On the Polyps and Echinoderms of New England,
110 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

with Descriptions of New Species. Procedings of the

Boston Society of Natural History, 10:333-375.

1867. Notes on Radiata. Transactions of the Connecticut

Academy of Arts and Sciences, 1 (2) :247-613, plates

4-10.

1868. Notice of the Corals and Echinoderms Collected

by Prof. C. F. Hartt, at the Abrolhos Reef, Province of Bahia, Brazil, in 1867, No. 4. Transactions

of the Connecticut Academy of Arts and Sciences,

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Corals. Proceedings of the Boston Society of Natural

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1871. Brief Contributions to Zoology from the Museum

of Yale College, No. 15: Descriptions of Starfishes

and Ophiurans from the Altantic Coasts of America

and Africa. American Journal of Science, 2:130-133.

1878. Notice of Recent Additions to the Marine Fauna

of the Eastern Coast of North America, nos. 1 and

2. American Journal of Science and Arts (3), 16:

207-215, 371-378.

1880. Notice of Recent Additions to the Marine Invertebrata of the Northeastern Coast of America, with

Description of New Genera and Species and Critical Remarks on Others. Proceedings of the United

States National Museum, 2:165-205.

1882. Brief Contributions to Zoology from the Museum of

Yale College, No. 49: Notice on the Remarkable

Marine Fauna Occupying the Outer Banks off the

Southern Coast of New England, No. 3. American

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1884. Notice of the Remarkable Marine Fauna Occupying

the Outer Banks off the Southern Coast of New

England, and of Some Additions to the Fauna of

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States Commisisoner of Fish and Fisheries for 1882,

26 pages.

1885. Results of the Explorations Made by the Steamer

Albatross off the Northern Coast of the United

States in 1883. Annual Report of the Commissioner

for Fish and Fisheries for 1883: 107 pages, (503-

699), 44 plates.

1894. Descriptions of New Species of Starfishes and

Ophiurans, with a Revision of Certain Species

Formerly Described. Proceedings of the United States

National Museum, 17:243-297.

1895. Distribution of the Echinoderms of Northeastern

America. American Journal of Science, 49:127-141,

199-212.

1899. Revision of Certain Genera and Species of Starfishes. Transactions of the Connecticut Academy of

Arts and Sciences, 10:145-234, plates 24-30.

1901. Additions to the Fauna of the Bermudas from the

Yale Expedition of 1901. Transactions of the Connecticut Academy of Arts and Science, 11:15-62.

1902. Additions to the Fauna of the Bermudas from the

Yale Expedition of 1901, with Notes on other

Species. Transactions of the Connecticut Academy

of Arts and Sciences, 11:15-62, 9 plates, 6 figures.

1907. The Bermuda Islands. Part 4, Geology and Paleontology; Part 5, An Account of the Coral Reefs.

Transactions of the Connecticut Academy of Arts

and Sciences, 12:45-348, plates 16-40.

1909. Remarkable Development of Starfishes on the

Northwest American Coast; Hybridism; Multiplicity

of Rays; Teratology; Problems in Evolution; Geographical Distribution. American Naturalist, 43:

542-555.

1914. Monograph of the Shallow-Water Starfishes of the

North Pacific Coast, from the Arctic Ocean to

California, with Revisions of Various Extra-Limital

Genera and Species. Harriman Alaska Series, 14:

420 pages, 110 plates.

1915. Report on the Starfishes of the West Indies, Florida,

and Brazil, Including Those Obtained by the Bahama Expedition from the University of Iowa in

1893. Bulletin of the State of University of Iowa,

7: 232 pages, 29 plates.

Vevers, H. G.

1956. Observations on Feeding Mechanisms in Some

Echinoderms. Proceedings of the Zoological Society

of London, 126:484-485.

Viguier, C.

1879. Anatomie Comparee du Squelette des Stellerides.

Archives de Zoologie Experimentale et Generate, 7:

33-250, plates 5-16.

Whitelegge, T.

1897. The Echinodermata of Funafuti. (In The Atoll of

Funafuti . . . Based on Collections Made by Mr.

Charles Hedley, etc. Part 2). Memoirs of the Australian Museum, 3:155—162.

Wilson, D. P.

1955. In Illustrated London News, 227:2-4.

Wood-Mason, J., and A. Alcock.

1891. Natural History Notes from H. M. Marine Survey

Steamer Investigator. Annals and Magazine of Natural History, series 6, 7:12-15, 427-433.
NUMBER 126 111

PLATE 1.—A, Luidia senegalensis, abactinal; B, same, actinal (actual size=50 mm R); c, Luidia

clathrata, abactinal; D, same, actinal (actual size= 77 mm R).
112 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATF 2.—A, Luidia alternata, abactinal; B, same, actinal (actual size=115 mm R); c, Luidia

sagamtna, abactinal; D, same, actinal (actual size=8 mm R).
NUMBER 126 113

PLATE 3.—A, Luida barbadensis, abactinal; B, same, actinal (actual size=75 mm R); c, Luidia

elegans, ahactinal; D, same, actinal (actual size =77 mm R).
114 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 4.—A, Astropecten americanus, abactinal; B, same, actinal (actual size —72 mm R); c,

Astropecten articulatus, abactinal; D, same, actinal (actual size=84 mm R).
NUMBER 126 115

PLATE 5.—A, Astropecten cingulatus, abactinal; B, same, actinal (actual size —31 mm R); c,

Astropecten comptus, abactinal; D, same, actinal (actual size=:61 mm R).
116 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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PLATE 6.—A, Aslropecten duplicates, abactinal; B, same, actinal (actual size=45 mm R); c,

Astropecten marginatus, abactinal; D, same, actinal (actual size=55 mm R).
NUMBER 126 117

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PLATE 7.—A, Astropecten nitidus, abactinal; B, same, actinal (actual size=40 mm R); c, Psilasttr

cassiope, abactinal; D, same, actinal (actual size=82 mm R).
118 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 8.—A, Psilaster patagiatus, abactinal; B, same, actinal (actual size=64 mm R); c,

Persephonaster echinulatus, abactinal; D, same, actinal (actual size=45 mm R).
NUMBER 126 119

I

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PLATE 9.-A, Plutonaster intermedius, abactinal; B, same, actinal (actual size=lll mm R);

c, Dytaster insignis, abactinal; D, same, actinal (actual size=86 mm R).
120 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 10.—A, Blakiaster conicus, abactinal; B, same, actinal (actual size=26J> mm R); c,

Tethyaster grandts, abactinal; D, same, actinal (actual size=105 mm R).
NUMBER 126 121

A B

PLATE 11.—A, Dtpsacaster antillensis, abactinal; B, same, actinal (actual size—12 mm R); c,

Goniopecten demonstrans, abactinal; D, same, actinal (actual size=138 mm R).
122 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 12.—A, Prionaster elegans, abactinal; B, same, actinal (actual size=37 mm R); c,

Benthopecten simplex, abactinal; D, same, actinal (actual size—48 mm R).
NUMBER 126 123

PLATE 13.—A, Cheiraster mirabilis, abactinal; B, same, actinal (actual size=120 mm R); c,

Cheiraster echinulatus, abactinal; D, same, actinal (actual size=35 mm R).
124 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 14.—A, Cheiraster enoplus, abactinal; B, same, actinal (actual sizez=104 mm R); c.

Pectinaster mixtus, abactinal; D, same, actinal (actual size—41 mm R).
NUMBER 126 125

PLATE 15.—A, Pecttnaster gracilis, abactinal; B, same, actinal (actual size=13 mm R); c,

Odontaster hispidus, abactinal; D, same, actinal (actual size—37 mm R).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 16.—A, Odontaster setosus, abactinal; B, same, actinal (actual size=28 mm R) ; c, Goniaster

tessellatus, abactinal; D, same, actinal (actual size=71 mm R).
NUMBER 126 127

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D

PLATE 17.—A, Anthenoides piercei, abactinal; B, same, actinal (actual size=82 mm R); c,

Ceramaster grenadensts, abactinal; D, same, actinal (actual size—21 mm R).
128 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 18.—A, Peltaster nidarosiensis, abactinal; B, same, actinal (actual size=75 mm R) ; c,

Peltaster placenta, abactinal; D. same, actinal (actual size=75 mm R).
NUMBER 126 129

PLATE 19.—A, Plinthaster dentatus, abactinal; B, same, actinal (actual size=51 mm R); c,

Tessellaster notabilis, abactinal; D, same, actinal (actual size—76 mm R).
130 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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PLATE 20.—A, Tana parva, abactinal; B, same, actinal (actual size=18 mm R); c, Rosaster

alexandri, abactinal; D, same, actinal (actual size —24 mm R).
NUMBER 126 131

PLATE 21.—A, Circeaster americanus, abactinal; B, same, actinal (actual size=76 mm R); c,

Litonotaster intermedius, abactinal; D, same, actinal (actual size—17 mm R).
132 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 22.—A, Paragonaster subtilis, abactinal; B, same, actinal (actual size=65 mm R); c,

Nymphaster arenatus, abactinal; D, same, actinal (actual size= 84 mm R).
NUMBER 126 133

PLATE 23.—A, Nymphaster subsptnosus, abactinal; B, same, actinal (actual size=70 mm R);

c, Pseudarchaster gracilis, abactinal; D, same, actinal (actual size—190 mm R).
134 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 24.—A, Oreaster reticulatus, abactinal; B, same, actinal (actual size=91 mm R); c,

Chaetaster nodosus, abactinal; v, same, actinal (actual size=60 mm R).
NUMBER 126 135

PLATE 25.—A, Drachmaster bullisi, abactinal; B, same, actinal (actual size —33 mm R); c, Tamaria

floridae, abactinal; D, same, actinal (actual size z= 25 mm R).
136

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 26.—A, Tamaria halperni, abactinal; B, same, actinal (actual size=53 mm R); c, Tamaria

passiflora, abactinal; D, same, actinal (actual size—24 mm R).
NUMBER 126 137

A * D Ml

Q B

D

PLATE 27.—A, Linckia guildingii, abactinal; B, same, actinal (actual size=38 mm R); c, Linckia

nodosa, abactinal; D, same, actinal (actual size —36 mm R).
138 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 28.—A, Linckia bouvieri, abactinal; B, same, actinal (actual size =134 mm R); c,

Ophidiaster guildingii, abactinal; D, same, actinal (actual size—33 mm R).
NUMBER 126 139

I

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PLATE 29.—A, Narcissia trigonaria, abactinal; B, same, actinal (actual size—74 mm R); c, Solaster

notophrynus, abactinal; D, same, actinal (actual size—114 mm R).
140 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE SO.—A, Solaster caribbaeus, abactinal; B, same, actinal (actual size—58 mm R); c,

Lophaster verrilli, abactinal; D, same, actinal (actual size=38 mm R).
NUMBER 126 141

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PLATE 31.—A, Hymenaster rex, abactinal; B, same, actinal (actual size=81 mm R); c, Hymenaster

modestus, abactinal; D, same, actinal (actual size=30 mm R).
142

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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PLATE 32.—A, Hymenaster anomalus, abactinal; B, same, actinal (actual size=14 mm R); c,

Pteraster personatus, abactinal; D, same, a^tin^i (actual size=70 mm R).
NUMBER 126 14S

PLATE 33.—A, Pteraster caribbaeus, abactinal; B, same, actinal (actual size=36 mm R); c,

Pteraster rugosus, abactinal; D, same, actinal (actual size=24 mm R).
144 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 34.—A, Pteraster militaroides, abactinal; B, same, actinal (actual size=22 mm R); c,

Pteraster acicula, abactinal; D, same, actinal (actual size—10 mm R).
NUMBER 126 145

PLATE 35.—A, Pteraster species, abactinal; B, same, actinal (actual size=26 mm R); c,

Calyptraster personatus, abactinal; D, same, actinal (actual size=20 mm R).
146 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 36.—A, Leilaster radians, abactinal; B, same, actinal (actual size=8 mm R).

c, Poraniella regularis, abactinal; D, same, actinal (actual size =17 mm R).
NUMBER 126 147

PLATE 37.—A, Marginaster pectinatus, abactinal; B, same, actinal (actual size=ll mm R); c,

Henricia sexradiata, abactinal; o, same, actinal (actual size=46 mm R).
148 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Pi. VIE 38.—A, Henricia antillarum, abactinal; B, same, actinal (actual size=55 mm R); c,

Henricia species, abactinal; n, same, actinal (actual size—62 mm R).
NUMBER 186 149

PLATE 39.—A, Echinaster serpentarius, abactinal; B, same, actinal (actual size=43 mm R); c,

Echinaster echinophorus, abactinal; D, same, actinal (actual size = 37 mm R).
150 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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PLATE 40.—A, Echinaster sentus, abactinal; B, same, actinal (actual size=70 mm R); c,

Echinaster modestus, abactinal; D, same, actinal (actual size=49 mm R).
NUMBER 126 151

PLATE 41.—A, Echinaster species A , abactinal; B, same, actinal (actual size —84 mm R); c,

Echinaster species B, abactinal; D, same, actinal (actual size —65 mm R).
152 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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PLATE 42.—A, Echinaster species c, abactinal; B, same, actinal (actual size = 72 mm R); c,

Echinaster brasiliensis, abactinal; D, same, actinal (actual size=65 mm R).
NUMBER 126 153

PLATE 43.—A, Verrillaster spinulosus, abactinal; B, same, actinal (actual sizei=65 mm R); c,

Ampheraster alaminos, abactinal; D, same, actinal (actual size=62 mm R).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 44.—A, Sclerasterias contorta, abactinal; B, same, actinal (actual size=90 mm R); c,

Astertas forbesi, abactinal; D, same, actinal (actual size=45 mm R).
NUMBER 126 155

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PLATE 45.—A, Coscinasterias tenuispina, abactinal; B, same, actinal (actual size—30 mm R); c,

Coronaster briareus, abactinal; D, same, actinal (actual size=125 mm R).
156 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 46.—A, Doraster constellatus, abactinal; B, same, actinal (actual size=80 mm R); c,

Mammaster sigsbeei, abactinal; D, same actinal (actual size=47 mm R).
NUMBER 126 157

PLATE 47.—A, Zoroaster fulgens, abactinal; n, same, actinal (actual size=85 mm R); c, Odinia

antillensis, abactinal; D, same, actinal (actual size —47 mm R).
158 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 48.—A, Midgardia xandaros, abactinal; B, same, actinal; c, abactinal view of disc (actual

size=680 mm R).

•Cl U. S. GOVERNMENT PRINTING OFFICEl l»7 » 48«-3«/35
Publication in Smithsonian Contributions to Zoology

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titles will carry a final line indicating the higher categories to which the taxon is referable:

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For references within the text, use the author-date system: "(Jones 1910)" and "Jones

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Illustrations (line drawings, maps, photographs, shaded drawings) can be intermixed

throughout the printed text. They will be termed Figures and should be numbered consecutively; however, if a group of figures is treated as a single figure, the components should be

indicated by lowercase italic letters on the illustration, in the legend, and in text references:

"Figure 96." If illustrations (usually tone photographs) are printed separately from the text as

full pages on a different stock of paper, they will be termed Plates, and individual components

should be lettered (Plate 9b) but may be numbered (Plate 9: figure 2). Never combine the

numbering system of text illustrations with that of plate illustrations. Submit all legends on

pages separate from the text and not attached to the artwork. An instruction booklet for the

preparation of illustrations is available from the Press on request.

In the bibliography (usually called "Literature Cited"), spell out book, journal, and

article titles, using initial caps with all words except minor terms such as "and, of, the." For

capitalization of titles in foreign languages, follow the national practice of each language.

Underscore (for italics) book and journal titles. Use the colon-parentheses system for volume,

number, and page citations: "10(2) :5-9." Spell out such words as "figures," "plates," "pages."

For free copies of his own paper, a Smithsonian author should indicate his requirements

on "Form 36" (submitted to the Press with the manuscript). A non-Smithsonian author will

receive 50 free copies; order forms for quantities above this amount with instructions for payment will be supplied when page proof is forwarded.