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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 488

Living and Fossil Genera of the Clypeasteroida (Echinoidea: Echinodermata): An Illustrated Key and Annotated Checklist

Rich Mooi

SMITHSONIAN INSTITUTION PRESS

Washington, D.C.

1989

ABSTRACT

Mooi, Rich. Living and Fossil Genera of the Clypeasteroida (Echinoidea: Echinodermata): An Illustrated Key and Annotated Checklist. Smithsonian Contributions to Zoology, number 488, 51 pages, 34 figures, 1989.—The illustrated key allows identification of 26 living and 49 fossil clypeasteroid genera, plus the tiny fossil, Togocyamus. It incorporates internal and external characteristics of the test, Aristotle's lantern, spines, pedicellariae, and podia. Figures depict features used in the key, and provide an illustrated glossary of clypeasteroid terminology. A checklist groups the genera into three suborders (the Clypeasterina, Laganina, and Scutellina) and summarizes, for each genus, information on authorities, type species, number of included species, distribution, and stratigraphic occurrence.

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

Library of Congress Cataloging in Publication Data

Mooi, Rich

Living and Fossil genera of the Clypeasteroida (Echinoidea, Echinodermata).

(Smithsonian contributions to zoology ; no. 488)

Bibliography: p.

Supt. of Docs. no.: SI 1.27:488

1. Sand Dollars—Identification. I. Title. II. Series.

QL1.s54 no. 488 591 s [593.9′ 89–600113 [QL384.E2]

Living and Fossil Genera of the Clypeasteroida (Echinoidea: Echinodermata): An Illustrated Key and Annotated Checklist

Rich Mooi

____________

Rich Mooi, Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

Introduction

The order Clypeasteroida A. Agassiz, 1872 is composed of some 150 living and 750 fossil species of irregular echinoids commonly known as the sand dollars, keyhole urchins, and sea biscuits. There have been no comprehensive keys to the genera of this order since Mortensen's (1948) great monograph. Since the publication of that work, several workers (for example, Nisiyama, 1963; Philip and Foster, 1971; and especially Durham, 1954,1955; and Durham et al., 1966) have described new genera and redescribed some of the lesser known fossil taxa covered by Mortensen. Therefore, there is need for a revised key that draws upon this new information, and allows identification of specimens without resorting to the out-dated generic keys scattered throughout Mortensen's large monograph. The new checklist provides brief notes on authorities, type species, number of species, geographic range, stratigraphic occurrence, and selected representative figures gleaned from the literature and personal observations. The key and checklist represent an uncritical tabulation of the known clypeasteroid genera, and are merely intended to allow identification, and to summarize current knowledge of these genera. Although the key is, of course, artificial, it relies on many characters used in studies like that of Mooi (1987; in press), which provide phylogenetic classifications above the subordinal level as well as generic and familial revisions of each suborder.

The specimens used in the formulation of this key (and the pending phylogenetic analyses) were provided in part by the following institutions: National Museum of Natural History, Smithsonian Institution (NMNH), Washington, D.C., which houses the collections of the former United States National Museum (USNM); the British Museum (Natural History) (BMNH), London; the Museum of Comparative Zoology (MCZ), Cambridge, Massachusetts; the American Museum of Natural History (AMNH), New York; the Muséum National d'Histoire Naturelle (MNHN), Paris; the Western Australia Museum (WAM), Perth; the Royal Ontario Museum (ROM), Toronto; and the Florida Department of Natural Resources (FDNR), St. Petersburg. Additional specimens were examined from the collections in the laboratory of Malcolm Telford, University of Toronto (UT).

CONVENTIONS USED IN THE KEY.—The new key relies on characteristics of both the test (apical system, peristome and periproct position, gonopores, hydropores, internal buttresses, Aristotle's lantern, lantern supports, and plate patterns) and external structures (pedicellariae, spines, podia) to provide as complete characterizations of adult morphology of both fossil and living genera as possible. Ambulacra and interambulacra are numbered according to Lovén's system (See Illustrated Glossary, Figure 31). Fossil genera in the key are marked with a dagger (†). A brief comment on the range (“terra typica”) of both living and fossil genera is provided to aid in identifying specimens for which collection data are available.

Unfortunately, it was necessary to resort to characteristics of internal structure in some cases in order to make the couplets unambiguous. With the increasing use of radiography in taxonomic work, it is hoped that dissection of specimens will not always be necessary to reveal useful characters in these couplets. As many opposing characters as possible are used in each couplet to maximize the possibility for accurate identification of incomplete specimens and poorly preserved fossil material. Because fossil material seldom displays the podial, pedicellarial and spine features sometimes given in the couplets, an attempt has been made to provide alternatives to these types of characters whenever fossils are involved.

The terminology associated with echinoid morphology can be complex. To aid the reader in the interpretation of these terms, an Illustrated Glossary is provided at the end of the paper. This type of glossary is provided instead of a written glossary because (1) an image is more useful to someone trying to interpret the key, (2) images tend to remain in the mind of the reader, and (3) verbal descriptions can often become jargon-laden themselves, leading the reader on a potentially circular chase for definitions. The first time a term occurs in the key, it is accompanied by a reference to the figure(s) in the Illustrated Glossary (denoted in the text by “I.G.” and a figure number) that explicates that term. If the reader prefers, a written glossary can be found in Durham et al. (1966:U253–U256) and Davies (1971:161–165), and a general guide to echinoid morphology can be found in Smith (1984:1–6).

NOTE ON TAXA AND CLASSIFICATION USED IN THE KEY AND CHECKLIST.—The present key incorporates a total of 75 clypeasteroid genera (26 living, 49 fossil), plus Togocyamus, which has here been tentatively recognized as lying outside the order Clypeasteroida. Togocyamus occupies this special position partly because of a lack of certain features generally felt to be diagnostic of the Clypeasteroida, most notably the greater width of ambulacra relative to interambulacra, the distribution of accessory podia in fields, and the enclosure of the sphaeridia within the test (Kier, 1982; Mooi, in press). Eight fossil genera (Fibulina, Peronellites, Tetradiella, Scutulum, Samlandaster, Proescutella, Kieria, and Mennerella), and one living genus (Marginoproctus) are omitted from the key (but included in the checklist) because their characteristics are too poorly known to allow placement. Stephenson (1968:136) synonymized Fibulina Tornquist, 1904, with Fibularia Lamarck, 1816 and Peronellites Hayasaka and Morishita, 1947 has recently been synonymized with Peronella Gray, 1855 by Wang (1982b:143). Runa L. Agassiz, 1841 and Tournoueraster Lambert, 1914 are not considered in either key or checklist, as they are probably internal casts of previously described forms (Durham, 1955:187). The living genus Taiwanaster Wang, 1984 is, as far as can be determined, identical with Sinaechinocyamus Liao, 1979, and is listed with Sinaechinocyamus in the key. Because type material from these genera has not yet been examined in order to make a decision regarding synonymy, these two genera are listed separately in the checklist.

Three of the four suborders recognized by Durham et al. (1966) are employed as major subsections of the key: the Clypeasterina, Laganina, and Scutellina. For the purposes of the key, the suborder Rotulina has been incorporated into the Laganina, as many rotuline characters are shared with the Laganina. One other modification has been introduced to the basic scheme of Durham et al. (1966). Four of the fossil genera (Scutellina L. Agassiz, 1841; Porpitella Pomel, 1883; Lenita Desor, 1847; Eoscutum Lambert, 1914) that have long been considered members of the family of tiny laganines, the Fibulariidae, have little in common with this family, apart from small size. This is reflected by their position in the key, as they do not even fall within the suborder Laganina. They have here been referred to the suborder Scutellina, in both the key and the annotated checklist. Apart from this relatively minor provisional change, no attempt has been made to formally revise clypeasteroid classification, or group genera into families.

ACKNOWLEDGMENTS.—I would like to thank Malcolm Telford, who provided encouragement and advice during the formative years of a project that laid the groundwork for this key—truly a man in an iconoclass of his own. David Pawson and Cindy Ahearn, of the NMNH, facilitated access to specimens and information, and helped convince me that this undertaking was indeed necessary. I thank Steve Beadle for valuable new information on stratigraphic occurrences of western American clypeasteroids. I am also grateful to the following for hospitality extended during pillaging raids: Frederick Collier and Jann Thompson (Paleobiology, NMNH); Ailsa Clark, Gordon Paterson, Andrew Smith, David Lewis, and Richard Jefferies (BMNH); Robert Woollacott and R.C. Eng (MCZ); Catherine Vadon, Jean Roman, and Y. Gayrard (MNHN); Loisette Marsh and Kenneth McNamara (WAM); Harold Feinberg (AMNH); Janet Waddington (ROM); Sandra Farrington and David Camp (FDNR). Personal support was provided by a Natural Sciences and Engineering Research Council of Canada Post-doctoral Fellowship held while I was a Post-doctoral Fellow at the NMNH.

An Illustrated Key to the Living and Fossil Genera of the Clypeasteroida

(† = fossil taxa)

FIGURE 1.—Sphaeridia and podial distribution in ambulacrum III: a, Togocyamus seefredi Oppenheim, USNM, no number (plate pattern not known); b, Monostychia australis Laube, USNM 96251; c, Fibularia ovulum Lamarck, UT, Enewetak. (Buccal and accessory podial pores indicated by solid dots; “open” sphaeridia indicated by solid circle, sphaeridia enclosed within test by dotted circle; plate pattern shown for ambulacrum III only, pattern unknown in a; peristome cross-hatched; scale bars: a and c = 1 mm; b = 5 mm.)

Order CLYPEASTEROIDA A. Agassiz, 1872

FIGURE 2.—Top (aboral) view of Aristotle's lantern in clypeasteroids: a, Arachnoides placenta (Linné), WAM 271.77; b, Laganum laganum (Leske), WAM 2082.25; c, Encope emarginata (Leske), UT, Belize; d, Echinodiscus bisperforatus (Leske), UT, New Guinea. (Anterior towards top of page; scale bars = 5 mm; abbreviations: ew = external wing; iw = internal wing; sa = supra-alveolar process; sd = subdental process.)

FIGURE 3.—Internal view of lantern supports and basicoronal plate patterns in clypeasteroids: a, Fellaster zelandiae (Gray), adapted from Kier (1970, pl. 23); b, Echinarachnius parma (Lamarck), adapted from Kier (1970, pl. 23). (Peristome in solid black, interambulacra stippled, lantern supports in heavy outline; anterior towards top of page; scale bars = 10 mm.)

Suborder CLYPEASTERINA A. Agassiz, 1872

FIGURE 4.—Food grooves of clypeasterines: a, Oral surface of Clypeaster subdepressus (Gray), UT, Florida Keys; b, oral surface of Arachnoides placenta (Linné), UT, New Guinea; c, aboral surface of Arachnoides placenta (Linné), UT, New Guinea. (Respiratory podial rows, peristomes, and periproct in solid black; food grooves and periproctal grooves stippled; anterior towards top of page; scale bars = 30 mm.)

FIGURE 5.—Plate patterns on the oral surface of clypeasterines: a, Ammotrophus cyclius H.L. Clark, Paratype, MCZ 3352; b, Monostychia australis Laube, after Durham (1955:126); c, Arachnoides placenta (Linné), UT, New Guinea; d, Fellaster zelandiae (Gray), AMNH 2302; e, Scutellinoides patella Tate, USNM 96254; f, Willungaster scutellaris Philip and Foster, after Philip and Foster (1971:689). (Interambulacra shaded; anterior towards top of page; scale bars: a–d = 50 mm; e,f = 5 mm.)

FIGURE 6.—Demiplates and primary plates in the petaloids of clypeasteroids: a, Arachnoides placenta (Linné), UT, New Guinea; b, Neorumphia elegans (Sánchez Roig), after Durham (1954:683); c, Rotula deciesdigitata (Leske), USNM 6991; d, Monophoraster darwini (Desor), ROM 5578. (All figures show a portion of a petaloid in a half ambulacrum; ambitus towards bottom of page; respiratory podial pores in solid black; scale bars = 20 mm; abbreviations: d = demiplate; p = Primary plate.)

FIGURE 7.—Interambulacral plate pattern adjacent to the apical system of clypeasteroids: a, Peronella lesueuri (Valenciennes). BMNH 1981.2.6.86; b, Rotula deciesdigitata (Leske), USNM 2307; c, Tarphypygus clarki (Lambert), USNM 2522; d, Fibulariella oblonga (Gray), MCZ 7565; e, Thagastea wetterlei Pomel, ROM 4100 (ambulacrum V obscured by matrix); f, Scutellina lenticularis (Lamarck), BMNH 31187. (Details of ambulacral plating and petaloids omitted; anterior towards top of page; scale bars in mm.)

FIGURE 8.—Arrangement of respiratory podia in petaloids of clypeasteroids: a, Condition typical of laganines, Laganum laganum (Leske), UT, New Guinea; b, condition typical of clypeasterines and scutellines, Echinodiscus bisperforatus (Leske), UT, New Guinea. (Both figures show portion of a petaloid in a half ambulacrum; pore pairs solid black, conjugation groove indicated by dashed lines, plate sutures by solid lines; ambitus towards bottom of page; scale bars = 20 mm.)

FIGURE 9.—Basicoronal plate patterns of clypeasteroids: a, Peronella lesueuri (Valenciennes), BMNH 1981.2.6.86; b, Neolaganum archerensis (Twitchell), after Durham (1954:679); c, Rotula deciesdigitata (Leske), MCZ 2551 (note that paired plates adjacent to peristome are not basicoronals, but first post-basicoronals); d, Eoscutella coosensis (Kew), after Durham (1955:98); e, Iheringiella patagonensis (Desor), after Durham (1955:126); f, Scutella subrotunda (Leske), after Durham (1955:98). (Interambulacral plates shaded, peristomes in solid black; anterior towards top of page; scale bars = 10 mm.)

FIGURE 10.—Keels and peristomial points in the food groove of oral ambulacrum III in clypeasteroids: a, Wythella eldridgei (Twitchell), USNM Acc. 268937; b, Periarchus lyelli (Conrad), USNM 312506; c, Scaphechinus mirabilis A. Agassiz, UT, Vostok Bay. (Ends of ambulacral basicoronal plates indicated by arrows; Food grooves represented by dotted lines, keels and peristomial points in solid black; anterior towards top of page: scale bars = 10 mm.)

FIGURE 11.—Miliary spines of clypeasteroids: a, Heliophora orbiculus (Linné), BMNH 1953.1.29.197; b, Echinarachnius parma (Lamarck), UT, New Brunswick; c, Leodia sexiesperforata (Leske), UT, Florida Keys. (Semitransparent views showing epidermis, basal musculature, and internal calcite skeleton. Heavy line at base of spine represents surface of test; scale bar = 500 μm.)

FIGURE 12.—Rotules from the Aristotle's lanterns of clypeasteroids: a, Echinocyamus pusillus (O.F. Müller), UT, Scotland; b, Heliophora orbiculus (Linné), BMNH 1904.10.28.5-8); c, Mellita quinquiesperforata, UT, North Carolina; d, Echinodiscus bisperforatus (Leske), UT, New Guinea. (In each of a to d, center of lantern is to left; upper figure shows side, and lower figures top of a single rotule; scale bar = 1 mm.)

Suborder LAGANINA Mortensen, 1948

FIGURE 13.—Food grooves of laganines: a, Rotula deciesdigitata (Leske), USNM 2307; b, Laganum laganum (Leske), UT, New Guinea; c, Fibulariella oblonga (Gray), MCZ 4094. (Food grooves, peristome, and periproct in solid black; scale bars: a and b = 30 mm; c = 5 mm.)

FIGURE 14.—Plate patterns on the oral surface of rotulid laganines: a, Rotuloidea fimbriata Etheridge, after Durham (1955:99); b, Heliophora orbiculus (Linné), USNM 32900; c, Rotula deciesdigitata (Leske), USNM 2307. (Interambulacra shaded; anterior towards top of page; scale bars = 30 mm.)

FIGURE 15.—Respiratory podial pores and spines of rotulids: a, Petaloid in ambulacrum IV of Heliophora orbiculus (Linné), AMNH 2298; b, petaloid in ambulacrum IV of Rotula deciesdigitata (Leske), USNM 2307; c, aboral primary spine of Heliophora orbiculus (Linné), BMNH 1953.1.29.197; d, aboral primary spine of Rotula deciesdigitata (Leske), MNHN EcEn997. In a and b, ambitus is shown at bottom of drawing, and scale bars = 10 mm; in c and d, scale bar = 500 μm.)

FIGURE 16.—Plate patterns on the oral surface of laganid and neolaganid laganines: a, Weisbordella caribbeana (Weisbord), after Durham (1954:678); b, Wythella eldridgei (Twitchell), after Durham (1954:683); c, Neolaganum archerensis (Twitchell), after Durham (1954:678); d, Cubanaster torrei (Lambert), after Durham (1954:678); e, Tarphypygus clarki (Lambert), USNM 2522; f, Peronella peronii (L. Agassiz), BMNH 48.5.8.2-7; g, Hupea decagonale (Lesson), after Durham (1955:140); h, Jacksonaster conchatus (M'Clelland), USNM E6999; i, Laganum laganum (Leske), UT, New Guinea. (Interambulacra shaded; anterior towards top of page; scale bars = 20 mm.)

FIGURE 17.—Aboral surfaces of neolaganids: a, Female Pentedium curator Kier, after Kier (1967:Plate 129); b, Durhamella floridana (Twitchell), after Kier (1968:34). (Anterior towards top of page; scale bars = 3 mm.)

FIGURE 18.—Madreporic plate, ocular plates, hydropores, and gonopores of laganines and scutellines: a, Wythella eldridgei (Twitchell), USNM Acc. 268937; b, Neolaganum durhami Cooke, Holotype, USNM 562290a; c, Peronella pellucida Döderlein, USNM 34258; d, Laganum laganum (Leske), WAM 2082.75; e, Fibulariella oblonga (Gray), MCZ 4590; f, Fibularia ovulum Lamarck, UT, Eniwetok; g, Echinocyamus pusillus (O.F. Müller), UT, Scotland; h, Porpitella hayesiana (Desmoulins), MNHN, no number; i, Eoscutum doncieuxi (Lambert), MNHN. (Anterior towards top of page; scale bars = 500 μm.)

FIGURE 19.—Aboral surface and transverse profiles of neolaganids: a, Cubanaster acunai (Sánchez Roig), after Durham et al. (1966:U474); b, Sanchezella sanchezi (Lambert), USNM, no number. (Anterior towards top of page; scale bars = 30 mm.)

FIGURE 20.—Petaloids of neolaganids: a, Neolaganum dalli (Twitchell), BMNH E75279; b, Neorumphia elegans (Sánchez Roig), after Durham et al. (1966:U474). (Anterior towards top of page; scale bars; a = 10 mm; b = 30 mm.)

FIGURE 21.—Radiographs showing internal buttress systems of clypeasteroids: a, Peronella lesueuri (Valenciennes). BMNH 1981.2.6.86; b, Echinocyamus pusillus (O.F. Müller), UT, Scotland; c, Periarchus lyelli (Conrad), USNM; d, Scutellina lenticularis (Lamarck), BMNH 31187; e, Lenita patellaris (Leske), MNHN, no number; f, Taiwanaster mai Wang, paratype, USNM E32412; g, Monophoraster darwini (Desor), USNM Acc. 339378; h, Encope aberrans Martens, FDNR 118975. (Anterior towards top of page; scale bars: a,c,d,g, and h = 10 mm; b,e, and f = 5 mm.)

FIGURE 22.—Tuberculation on oral surface of laganines and a scutelline: a, Lenicyamidia compta Brunnschweiler, after Durham et al. (1966:U470); b, Leniechinus herricki Kier, after Kier (1968:6, pl. 1, fig. 4); c, Lenita patellaris (Leske), MNHN, no number. (Peristome and periproct in solid black; anterior towards top of page; plate pattern shown only in naked zone (ambulacrum III and interambulacrum 5); scale bars = 3 mm.)

FIGURE 23.—Spines, pedicellariae, and podial spicules of laganines: a, Aboral primary spine of Fibularia ovulum Lamarck, MNHN EcEk131; b, aboral primary spine of Fibulariella oblonga (Gray), MCZ 7565; c, intact globiferous pedicellaria with poison glands (left), and single, cleaned valve (right) of Fibulariella oblonga (Gray), MCZ 7565; podial spicules of Fibulariella oblonga (Gray), after Mortensen (1948:219). (Scale bars: a to c = 100 μm; d = 50 μm.)

FIGURE 24.—Plate patterns and tuberculation of periproctal membrane of laganines: a, Fibularia ovulum Lamarck, MNHN EcEs5756; b, Echinocyamus pusillus (O.F. Müller), UT, Scotland; c, Mortonia australis (Desmoulins), after Mortensen (1949:160). (Anterior towards top of page; scale bars = 1 mm.)

Suborder SCUTELLINA Haeckel, 1896

FIGURE 25.—Pressure drainage channels and food grooves of scutellines: a, Protoscutella missippiensis (Twitchell), USNM, no number; b, Periarchus lyelli (Conrad), USNM 264066; c, Iheringiella patagonensis (Desor), BMNH E43225; d, Scutellaster oregonensis major (Kew), after Durham (1955:79); e, Kewia blancoensis (Kew), USNM 264125 (partially reconstructed); f, Echinarachnius parma (Lamarck), UT, New Brunswick; g, Dendraster excentricus (Eschscholtz), UT, Alaska; h, Merriamaster perrini (Weaver), BMNH E39170; i, Scaphechinus mirabilis A. Agassiz, UT, Vostok Bay; j, Remondella waldroni Wagner, USNM 181153; k, Monophoraster darwini (Desor), MCZ 3369; l, Leodia sexiesperforata (Leske), UT, Florida. (Pressure drainage channels stippled; peristomes and periprocts in solid black; anterior towards top of page; scale bars = 10 mm.)

FIGURE 26.—plate patterns on the oral surface of scutellines: a, Protoscutella mississippiensis (Twitchell), after Durtham (1955:154); b, Periarchus lyelli (Conrad), USNM Acc. 326367; c, Scutellina obovata L. Agassiz, MNHN, no number, d, Eoscutella coosensis (Kew), after Durham (1955:98); e, Iheringiella patagonensis (Desor), after Durham (1955:126); f, Scutella subrotunda (Leske), after Durham (1955:98); g, Parascutella bonali (Tournouer), MNHN, no number, h, Parmulechinus subtetragona (Grateloup), after Durham (1955:98); i, Nipponaster nipponicus (Nisiyama), after Durham (1955:154); j, Vaquerosella norrisi (Pack), after Durham (1955:103); k, Scaphechinus mirabilis A. Agassiz, UT, Vostok Bay; I, Remondella waldroni Wagner, USNM 181153; m, Monophoraster darwini (Desor), MCZ 3369; n, Karlaster pirabensis Marchesini Santos, adapted from Durham et al. (1966:U484); o, Mellita quinquiesperforata (Leske), UT, North Carolina; P, Leodia sexiesperforata (Leske), UT, Florida. (Interambulacrs shaded; anterior towards top of page; scale bars: a,b and d–p = 20 mm; c = 5 mm.)

FIGURE 27.—Aboral surface of scutellines showing test shape and petaloids: a, Mortonella quinquefaria (Say), adapted from Durham et al. (1966:U479); b, Periarchus lyelli (Conrad), USNM 312506; c, Scutella subrotunda (Leske), adapted from Durham (1953a:352); d, Parascutella bonali (Tournouer), MNHN, no number, e, Parmulechinus subtetragona (Grateloup), ROM 4122; f, Astrodapsis spatiosus Kew, USNM 15575; g, Abertella aberti (Conrad), USNM, no number; h, Dendraster excentricus (Eschscholtz), UT, Alaska; i, Scutaster andersoni Pack, USNM 371777; j, Astriclypeus manni Verrill, BMNH 81.8.2.3; k, Echinodiscus bisperforatus Leske, BMNH 97.6.10.1–7; i, Amplaster coloniensis Martínez, adapted from Martíez (1985:506); m, Mellita quinquiesperforata (Leske), UT, North Carolina; n, Leodia sexiesperforata (Leske), UT, Florids; o, Mellitella stokesii (L. Agassiz), USNM E15164; p, Encope micropora L. Agassiz, UT, Panama. (Petaloids in solid black; anterior towards top of page; scale bars = 40 mm.)

FIGURE 28.—Trailing podia and plate patterns of petaloids in scutellines: a, Aboral ambulacrum IV of Iheringiella patagonesis (Desor), ROM 5433M (plate pattern partially obscured); b, aboral ambulacrum II of Parascutella sp., UT, France. (Ambitus towards bottom of page; scale bars = 5 mm.)

FIGURE 29.—Spines and podial spicules of scutellines: a, Locomotory spine (left) and geniculate spine (right) from the oral surface of Dendraster excentricus (Eschscholtz), UT, Alaska; b, locomotory spine (left) and ambulacral spine (right) from the oral surface of Scaphechinus mirabilis A. Agassiz, BMNH 1902.12.4.1–5; c, single spicule from tip of an accessory podium of Astriclypeus manni Verrill, MCZ 2624; d, single spicule from the tip of an accessory podium of Mellita quinquiesperforata (Leske), UT, North Carolina; e, aboral primary spine of Mellita quinquiesperforata (Leske), UT, North Carolina; f, aboral primary spine of Leodia sexiesperforata (Leske), UT, Florida. (Scale bars: a and b = 1 mm; c and d = 30 μm; and e and f = 300 μm.)

FIGURE 30.—Lunule structure in scutellines: a, Oblique aboral view of cross-linked lunule wall from ambulacrum I of Echinodiscus bisperforatus (Leske), UT, New Guinea (entire specimen at left); b, oblique aboral view of festooned lunule wall from ambulacrum I of Mellita quinquiesperforata (Leske), UT, North Carolina (entire specimen at left). (Scale bars = 10 mm.)

Annotated Checklist of the Clypeasteroid Genera

The following is a checklist of all the known genera currently placed in the order Clypeasteroida, plus Togocyamus, which represents an entirely extinct sister group to that order (Mooi, in press). Although the generic synonymies of Mortensen (1948), Durham (1955), Durham et al. (1966), Philip and Foster (1971), and Kier and Lawson (1978) are adhered to in this list (interested readers are referred to these works for that information, it is not reiterated here), no attempt has been made to follow any particular family level classification, pending phylogenetic revision of the group. For that reason, the clypeasteroid genera are grouped into three major suborders, the Clypeasterina, Laganina, and Scutellina, and listed alphabetically in those suborders. Authorities for the genera, the type species, approximate number of species (because of the uncertain status of some taxa, this is not intended to be an exhaustively accurate estimate, but is merely intended to suggest the size, and to a certain extent, the relative taxonomic importance of the genus), distribution, stratigraphic range, and selected sources of figures (in the present, and other works) that illustrate general morphology are given for each entry.

Sister Group to the Clypeasteroida

Togocyamus Oppenheim, 1915

TYPE SPECIES.—Echinocyamus (Togocyamus) seefriedi Oppenheim, 1915.

NUMBER OF SPECIES.—Two (Kier, 1982:6).

DISTRIBUTION.—Known only from French West Africa, Togo, Senegal, and the “Gold Coast.”

STRATIGRAPHIC OCCURRENCE.—Paleocene.

FIGURES.—Figure 1a, ambulacrum; Kier (1982, pl. 1), podial pores, aboral and oral surfaces.

Order CLYPEASTEROIDA A. Agassiz, 1872

Suborder CLYPEASTERINA A. Agassiz, 1872

Ammotrophus H.L. Clark, 1928

TYPE SPECIES.—Ammotrophus cyclius H.L. Clark, 1928.

NUMBER OF SPECIES.—Two or three known living species, with at least one fossil species (Durham, 1955:127–128).

DISTRIBUTION.—Southern and Western Australia; Tasmania.

STRATIGRAPHIC OCCURRENCE.—Pleistocene to Recent.

FIGURES.—Figure 5a, oral surface plate pattern; Durham et al. (1966:U468), aboral and oral surfaces.

Arachnoides Leske, 1778

TYPE SPECIES.—Echinus placenta Linné, 1758.

NUMBER OF SPECIES.—Mortensen (1948:144) lists two living species, but no species are known only as fossils.

DISTRIBUTION.—Andaman Islands to Queensland, Western Australia; Java; Sumatra; New Guinea; Philippines.

STRATIGRAPHIC OCCURRENCE.—Pliocene to Recent.

FIGURES.—Figure 4b, oral surface; Figure 4c, aboral surface; Figure 5c, oral surface plate pattern; Figure 6a, petaloid plate pattern; Durham et al. (1966:U467), aboral and oral surfaces.

Clypeaster Lamarck, 1801

TYPE SPECIES.—Echinus rosaceus Linné, 1758.

NUMBER OF SPECIES.—Including fossil taxa, about 400 species names have been used for members of this genus (Mortensen, 1948:19). Approximately 40 living species are known.

DISTRIBUTION.—Living forms are common from the Caribbean, tropical Atlantic, Indian, eastern and western Pacific Oceans, especially the East Indies, Australia, and Hawaii, but not from the Mediterranean, where it is known only as a fossil.

STRATIGRAPHIC OCCURRENCE.—Upper Eocene (Auversian) to Recent.

FIGURES.—Figure 4a, oral surface with food grooves; Durham et al. (1966:U466), oral surface plate patterns, profiles, internal view of test, aboral surface.

Fellaster Durham, 1955

TYPE SPECIES.—Arachnoides zelandiae Gray, 1855.

NUMBER OF SPECIES.—The type, and a fossil species have been described.

DISTRIBUTION.—Once thought endemic to New Zealand, a fossil species has been described from southeastern Australia (Foster and Philip, 1980:155–156; Sadler and Pledge, 1985: 175–176).

STRATIGRAPHIC OCCURRENCE.—Durham (1955:127) listed it from the Pliocene, but later (Durham et al., 1966:464) suggested that Arachnoides occurs from the Oligocene to Recent.

FIGURES.—Figure 3a, lantern supports, basicoronal plates; Figure 5d, oral surface plate pattern; Durham et al. (1966:U467), aboral and oral surfaces.

Fossulaster Lambert and Thiéry, 1925

TYPE SPECIES.—Fossulaster halli Lambert and Thiéry, 1925.

NUMBER OF SPECIES.—Philip and Foster (1971:682–687) list two.

DISTRIBUTION.—South Australia.

STRATIGRAPHIC OCCURRENCE.—Upper Oligocene or Lower Miocene (Janjukian or Longfordian).

FIGURES.—Philip and Foster (1971, figs. 5, 7, pls. 125, 127, 128, 130–132), aboral and oral surface plate patterns, internal view of test, aboral and oral surfaces, side views.

Monostychia Laube, 1869

TYPE SPECIES.—Monostychia australis Laube, 1869.

NUMBER OF SPECIES.—Two or three (Durham, 1955:128).

DISTRIBUTION.—Southern Australia; Tasmania.

STRATIGRAPHIC OCCURRENCE.—Miocene.

FIGURES.—Figure 1b, ambulacrum; Figure 5b, oral surface plate pattern; Durham et al. (1966:U468), aboral and oral surfaces.

Scutellinoides Durham, 1955

TYPE SPECIES.—Scutellina patella Tate, 1891.

NUMBER OF SPECIES.—Probably only one (Philip and Foster, 1971:682).

DISTRIBUTION.—South Australia.

STRATIGRAPHIC OCCURRENCE.—Miocene.

FIGURES.—Figure 5e, oral surface plate pattern; Durham et al. (1966:U468), aboral and oral surfaces.

Willungaster Philip and Foster, 1971

TYPE SPECIES.—Willungaster scutellaris Philip and Foster, 1971.

NUMBER OF SPECIES.—Only the type species is known.

DISTRIBUTION.—South Australia.

STRATIGRAPHIC OCCURRENCE.—Upper Oligocene or Lower Miocene (Janjukian or Longfordian).

FIGURES.—Figure 5f, oral surface plate pattern; Philip and Foster (1971, fig. 8, pls. 127, 131, 133), aboral and oral surface plate patterns, aboral and oral surfaces, side views, internal view of test

Suborder LAGANINA Mortensen, 1948

Cubanaster Sánchez Roig, 1952

TYPE SPECIES.—Jacksonaster torrei Lambert in Sánchez Roig, 1962.

NUMBER OF SPECIES.—Approximately ten (Durham, 1954:681; Kier and Lawson, 1978:66).

DISTRIBUTION.—Cuba.

STRATIGRAPHIC OCCURRENCE.—Upper Eocene.

FIGURES.—Figure 16d, oral surface plate pattern; Figure 19a, aboral surface and transverse profile; Durham et al. (1966:U474), madreporite, aboral and oral surfaces.

Cyamidia Lambert and Thiéry, 1914

TYPE SPECIES.—Echinocyamus nummulitica Duncan and Sladen, 1884.

NUMBER OF SPECIES.—Mortensen (1948:227) suggests that there are two or three.

DISTRIBUTION.—India; Pakistan; Australia.

STRATIGRAPHIC OCCURRENCE.—Eocene.

FIGURES.—Durham et al. (1966:U469), madreporite, aboral and oral surfaces.

Durhamella Kier, 1968

TYPE SPECIES.—Laganum ocalanum Cooke, 1942.

NUMBER OF SPECIES.—Two (Kier, 1968:23–38).

DISTRIBUTION.—Georgia, Florida, USA.

STRATIGRAPHIC OCCURRENCE.—Middle Eocene.

FIGURES.—Figure 17b, aboral surface; Cooke (1959, pl. 20), aboral and oral surfaces, side view; Kier (1968, figs. 31–33, 37–39, 40–42, pls. 6–10), aboral and oral surfaces, apical system, internal view of test, lantern supports, aboral and oral surface plate patterns.

Echinocyamus van Phelsum, 1774

TYPE SPECIES.—Echinocyamus pusillus O.F. Müller, 1776.

NUMBER OF SPECIES.—Approximately 50, including fossil forms. Mortensen (1948:176–177) lists 16 living species, not counting members of his subgenus Mortonia, herein considered a separate genus.

DISTRIBUTION.—Caribbean; eastern Atlantic; northwestern Europe; Mediterranean; Indian Ocean; Australia; East Indies; Philippines; Malaysia; Hawaii; worldwide as a fossil.

STRATIGRAPHIC OCCURRENCE.—Durham et al. (1966:U469) say Upper Cretaceous (Senonian) to Recent, but Kier (1982:4) was apparently unable to find any clypeasteroids in strata before the Paleocene.

FIGURES.—Figure 18g, apical system; Figure 21b, internal buttress system; Figure 24b, periproctal membrane; Durham et al. (1966:U469), aboral and oral surfaces, internal view of test.

Fibularia Lamarck, 1816

TYPE SPECIES.—Fibularia ovulum Lamarck, 1816 (subsequent designation by the International Commission of Zoological Nomenclature [ICZN], 1950).

NUMBER OF SPECIES.—Probably at least 20, including fossils. Mortensen (1948:207) lists four living species, not counting members of his subgenus Fibulariella, herein considered a separate genus.

DISTRIBUTION.—Australia; East Indies; Indian Ocean; Red Sea; worldwide as a fossil.

STRATIGRAPHIC OCCURRENCE.—Upper Cretaceous (Senonian)(?) to Recent (See Echinocyamus).

FIGURES.—Figure 1c, ambulacrum; Figure 23a, aboral primary spine; Figure 24a, periproctal membrane; Durham et al. (1966:U469), aboral and oral surfaces.

Fibulariella Mortensen, 1948

TYPE SPECIES.—Fibularia acuta Yoshiwara, 1898.

NUMBER OF SPECIES.—Four or five, all living (Mortensen, 1948:207).

DISTRIBUTION.—Malaysia; south, west, and north Australia.

STRATIGRAPHIC OCCURRENCE.—Recent.

FIGURES.—Figure 7d, plate pattern adjacent to apical system; Figure 13c, food grooves; Figure 18e, apical system; Figure 23b, aboral primary spine; Figure 23c, globiferous pedicellaria; Figure 23d, podial spicules.

Fibulaster Lambert and Thiéry, 1914

TYPE SPECIES.—Sismondia michelini Cotteau, 1861.

NUMBER OF SPECIES.—Apparently only the type species has been ascribed to this poorly known genus.

DISTRIBUTION.—Europe, France.

STRATIGRAPHIC OCCURRENCE.—Eocene.

FIGURES.—Durham et al. (1966:UU469), aboral and oral surfaces, internal view of test.

Fibulina Tornquist, 1904

TYPE SPECIES.—Fibulina gracilis Tornquist, 1904.

NUMBER OF SPECIES.—Only the type species has been described. Stephenson (1968) synonymized Fibulina gracilis with Fibularia voeltzkowi Tornquist, 1904.

DISTRIBUTION.—Madagascar.

STRATIGRAPHIC OCCURRENCE.—Eocene.

FIGURES.—Durham et al. (1966:U472), aboral surface. Not in key.

Heliophora L. Agassiz, 1840

TYPE SPECIES.—Echinus orbiculus Linné, 1758.

NUMBER OF SPECIES.—Two (Durham, 1955:186), with one known living species.

DISTRIBUTION.—Tropical west coast of Africa; Cape Verde Islands; Ascension Island.

STRATIGRAPHIC OCCURRENCE.—Miocene to Recent.

FIGURES.—Figure 11a, miliary spine; Figure 12b, rotule; Figure 14b, oral surface plate pattern; Figure 15a, petaloid; Figure 15c, aboral primary spine; Durham et al. (1966:U490), aboral and oral surfaces, oral surface plate pattern.

Hupea Pomel, 1883

TYPE SPECIES.—Laganum decagonalis Lesson, 1883.

NUMBER OF SPECIES.—Only the type species is known.

DISTRIBUTION.—Malaysia; Polynesia.

STRATIGRAPHIC OCCURRENCE.—Pliocene to Recent.

FIGURES.—Figure 16g, oral surface plate pattern; Durham et al. (1966:U472), aboral and oral surfaces.

Jacksonaster Lambert and Thiéry, 1914

TYPE SPECIES.—Echinarachnius conchatus M'Clelland, 1840.

NUMBER OF SPECIES.—Kier and Lawson (1978:64–65) list two or three species that are probably attributable to this genus, with one known living species. New world species are not attributable to this genus.

DISTRIBUTION.—Java; Indonesia; Red Sea.

STRATIGRAPHIC OCCURRENCE.—Miocene to Recent.

FIGURES.—Figure 16h, oral surface plate pattern; Durham et al. (1966:U473), aboral and oral surfaces.

Laganum Link, 1807

TYPE SPECIES.—Echinodiscus laganum Leske, 1778.

NUMBER OF SPECIES.—At least 30 species have been named, including fossils. Mortensen (1948:307–308) lists about 11 living species.

DISTRIBUTION.—Philippines; New Guinea; Malaysia; Polynesia; north Australia; Japan; Indian Ocean; Red Sea; eastern Africa; Madagascar.

STRATIGRAPHIC OCCURRENCE.—Oligocene to Recent.

FIGURES.—Figure 2b, Aristotle's lantern; Figure 8a, respiratory podial pores; Figure 13b, food grooves; Figure 16i, oral surface plate pattern; Figure 18d, apical system; Durham et al. (1966:U472), madreporite, aboral and oral surfaces.

Lenicyamidia Brunnschweiler, 1962

TYPE SPECIES.—Lenicyamidia compta Brunnschweiler, 1962.

NUMBER OF SPECIES.—Only the type species is known.

DISTRIBUTION.—Western Australia.

STRATIGRAPHIC OCCURRENCE.—Lower Eocene (Cuisian or Lutetian).

FIGURES.—Figure 22a, oral surface tuberculation; Durham et al. (1966:U470), aboral and oral surfaces.

Leniechinus Kier, 1968

TYPE SPECIES.—Leniechinus herricki Kier, 1968.

NUMBER OF SPECIES.—Only the type species is known (Kier, 1968:5–12).

DISTRIBUTION.—Georgia, USA.

STRATIGRAPHIC OCCURRENCE.—Lower Middle Miocene.

FIGURES.—Figure 22b, oral surface tuberculation; Kier (1968, figs. 1–4, pls. 1, 2), aboral and oral surface plate patterns, peristome, aboral and oral surfaces, side view, internal view of test.

Marginoproctus Budin, 1980

TYPE SPECIES.—Marginoproctus djakonovi Budin, 1980.

NUMBER OF SPECIES.—Only the type species has been described. Although Budin (1980:305) placed this genus in the laganine family Fibulariidae, he was correct in noting affinities to Scutellina, which has here been listed in the suborder Scutellina.

DISTRIBUTION.—East Kamchatka, Commander and Kurile Islands, Sea of Okhtosk “to depths of 60 to 800 m” (Budin, 1980:308).

STRATIGRAPHIC OCCURRENCE.—Recent.

FIGURES.—Budin (1980, figs. 1–8), aboral and oral surfaces, side view, spines, and pedicellariae, Not in key.

Mortonia Gray, 1852

TYPE SPECIES.—Fibularia australis Desmoulins, 1837.

NUMBER OF SPECIES.—Mortensen (1948:197) lists two species, both living.

DISTRIBUTION.—Hawaii; Eniwetok; central Pacific.

STRATIGRAPHIC OCCURRENCE.—Recent.

FIGURES.—Figure 24c, periproctal membrane; Durham et al. (1966:U470), aboral and oral surfaces, internal view of test.

Neolaganum Durham, 1954

TYPE SPECIES.—Laganum archerensis Twitchell, 1915.

NUMBER OF SPECIES.—At least two (Durham, 1954:680–681).

DISTRIBUTION.—Gulf of Mexico, Florida.

STRATIGRAPHIC OCCURRENCE.—Upper Eocene.

FIGURES.—Figure 9b, basicoronal plates; Figure 16c, oral surface plate pattern; Figure 18b, apical sytem; Figure 20a, aboral surface; Cooke (1959, pl. 21), aboral and oral surfaces, side view; Durham et al. (1966:U474), aboral and oral surfaces, side view.

Neorumphia Durham, 1954

TYPE SPECIES.—Rumphia elegans Sánchez Roig, 1949.

NUMBER OF SPECIES.—Three (Durham, 1954:681–682).

DISTRIBUTION.—Cuba.

STRATIGRAPHIC OCCURRENCE.—Upper Oligocene.

FIGURES.—Figure 6b, petaloid plate pattern; Figure 20b, aboral surface; Durham et al. (1966:U474), petaloid plate pattern, aboral and oral surfaces.

Pentedium Kier, 1967

TYPE SPECIES.—Pentedium curator Kier, 1967.

NUMBER OF SPECIES.—Kier (1967) described only the type species.

DISTRIBUTION.—Georgia, USA.

STRATIGRAPHIC OCCURRENCE.—Lower Middle Miocene.

FIGURES.—Figure 17a, aboral surface; Kier (1967, figs. 2, 3, pls. 129, 130), juveniles, aboral and oral surfaces, oral surface plate pattern, internal view of test; Kier (1968, pl. 4) aboral surface.

Peronella Gray, 1855

TYPE SPECIES.—Laganum peronii L. Agassiz, 1841.

NUMBER OF SPECIES.—At least 30 species are known, including fossils. Mortensen (1948:258–259) lists about 15 living species.

DISTRIBUTION.—Indian Ocean; Australia; Japan; Taiwan; Malaysia; Philippines; Polynesia; East Indies.

STRATIGRAPHIC OCCURRENCE.—Upper Miocene to Recent.

FIGURES.—Figure 7a, plate pattern adjacent to apical system; Figure 9a, basicoronal plates; Figure 16f, oral surface plate pattern; Figure 18c, apical system; Figure 21a, internal buttress system; Durham et al. (1966:U473), aboral and oral surfaces.

Peronellites Hayasaka and Morishita, 1947

TYPE SPECIES.—Peronellites ovalis Hayasaka and Morishita, 1947.

NUMBER OF SPECIES.—Only the type species is known. Wang (1982b: 143, 144) synonymized Peronellites with Peronella.

DISTRIBUTION.—Taiwan.

STRATIGRAPHIC OCCURRENCE.—Miocene.

FIGURES.—Not in key, but see Peronella (sensu Wang, 1982b:144).

Proescutella Pomel, 1883

TYPE SPECIES.—Scutella cailliaudi Cotteau, 1861.

NUMBER OF SPECIES.—Two or three.

DISTRIBUTION.—France.

STRATIGRAPHIC OCCURRENCE.—Middle Miocene.

FIGURES.—Durham et al. (1966:U490), aboral and oral surfaces, side view. Not in key.

Rotula Schumacher, 1817

TYPE SPECIES.—Echinodiscus octiesdigitatus Leske, 1778 (= Echinodiscus deciesdigitatus Leske, 1778).

NUMBER OF SPECIES.—Only the type is known.

DISTRIBUTION.—West Africa, Angola.

STRATIGRAPHIC OCCURRENCE.—Miocene to Recent.

FIGURES.—Figure 6c, petaloid plate pattern; Figure 7b, plate pattern adjacent to the apical system; Figure 9c, basicoronal plates; Figure 13a, food grooves; Figure 14c, oral surface plate pattern; Figure 15b, petaloid; Figure 15d, aboral primary spine; Durham et al. (1966:U490), aboral and oral surfaces.

Rotuloidea Etheridge, 1872

TYPE SPECIES.—Rotuloidea fimbriata Etheridge, 1872.

NUMBER OF SPECIES.—Two other known fossil species, besides the type.

DISTRIBUTION.—West Africa, Morocco.

STRATIGRAPHIC OCCURRENCE.—Miocene to Pliocene.

FIGURES.—Figure 14a, oral surface plate pattern; Durham et al. (1966:U490), oral surface plate pattern.

Rumphia Desor, 1858

TYPE SPECIES.—Laganum rostratum L. Agassiz, 1841.

NUMBER OF SPECIES.—Only the type is known.

DISTRIBUTION.—East Indies; Australia.

STRATIGRAPHIC OCCURRENCE.—Miocene to Recent.

FIGURES.—Durham et al. (1966:U473), aboral and oral surfaces.

Sanchezella Durham, 1954

TYPE SPECIES.—Jacksonaster sanchezi Lambert, 1926.

NUMBER OF SPECIES.—Only the type species is known.

DISTRIBUTION.—Cuba.

STRATIGRAPHIC OCCURRENCE.—Upper Eocene.

FIGURES.—Figure 19b, aboral surface and transverse profile; Durham et al. (1966:U474), oral surface plate pattern, aboral and oral surfaces.

Sismondia Desor, 1858

TYPE SPECIES.—Scutella occitana Defrance, 1827.

NUMBER OF SPECIES.—Mortensen (1948:234) claims that some 30 species have been ascribed to this genus.

DISTRIBUTION.—Europe; Africa; East Indies; Australia; Japan.

STRATIGRAPHIC OCCURRENCE.—Eocene to Miocene.

FIGURES.—Durham et al. (1966:U473), aboral and oral surfaces, side view; Kier (1982, pl. 2), aboral and oral surfaces, side view.

Tarphypygus Arnold and Clark, 1927

TYPE SPECIES.—Tarphypygus ellipticus Arnold and Clark, 1927.

NUMBER OF SPECIES.—At least five (Kier and Lawson, 1978:63).

DISTRIBUTION.—Cuba; Jamaica.

STRATIGRAPHIC OCCURRENCE.—Eocene.

FIGURES.—Figure 7c, plate pattern adjacent to apical system; Figure 16e, oral surface plate pattern; Durham et al. (1966:U471), aboral and oral surfaces, side view.

Tetradiella Liao and Lin, 1981

TYPE SPECIES.—Tetradiella sinica Liao and Lin, 1981.

NUMBER OF SPECIES.—One.

DISTRIBUTION.—China, Guangxi.

STRATIGRAPHIC OCCURRENCE.—Late Tertiary, probably Plioene.

FIGURES.—Liao and Lin (1981, figs. 1–2, pl. 1), aboral and oral surfaces, side view, internal buttress system. Not in key.

Thagastea Pomel, 1888

TYPE SPECIES.—Thagastea wetterlei Pomel, 1888.

NUMBER OF SPECIES.—One or two.

DISTRIBUTION.—Europe; northern Africa, Tunisia.

STRATIGRAPHIC OCCURRENCE.—Eocene.

FIGURES.—Figure 7e, plate pattern adjacent to apical system; Durham et al. (1966:U471), aboral and oral surfaces, side view.

Tridium Tandon and Srivastava, 1980

TYPE SPECIES.—Tridium kieri Tandon and Srivastava, 1980.

NUMBER OF SPECIES.—Only the type species has been described.

DISTRIBUTION.—Known only from Kutch, India.

STRATIGRAPHIC OCCURRENCE.—Middle Eocene.

FIGURES.—Tandon and Srivastava (1980, figs. 1–2, pl. 1), apical system, basicoronal plates, aboral and oral surfaces, petaloids, internal structure, periproct and peristome.

Weisbordella Durham, 1954

TYPE SPECIES.—Peronella caribbeana Weisbord, 1934.

NUMBER OF SPECIES.—Probably at least three (Durham, 1954:682).

DISTRIBUTION.—Gulf of Mexico; West Indies.

STRATIGRAPHIC OCCURRENCE.—Upper Eocene.

FIGURES.—Figure 16a, oral surface plate pattern; Cooke (1959, pl. 20), aboral and oral surfaces, side views.

Wythella Durham, 1954

TYPE SPECIES.—Laganum eldridgei Twitchell, 1915.

NUMBER OF SPECIES.—Only the type species is known.

DISTRIBUTION.—Gulf of Mexico, Georgia.

STRATIGRAPHIC OCCURRENCE.—Upper Eocene.

FIGURES.—Figure 10a, keel in food groove; Figure 16b, oral surface plate pattern; Figure 18a, apical system; Cooke (1959, pl. 21), aboral and oral surfaces, side view; Durham et al. (1966:U474), aboral and oral surfaces.

Suborder SCUTELLINA Haeckel, 1896

Abertella Durham, 1955

TYPE SPECIES.—Scutella aberti Conrad, 1842.

NUMBER OF SPECIES.—Durham (1953b:351; 1955:178) suggested that four species belong to this genus, including the type. He later added two more species (Durham, 1957:627).

DISTRIBUTION.—Eastern USA, Maryland; Guatemala; Mexico.

STRATIGRAPHIC OCCURRENCE.—Middle Miocene (Steven C. Beadle, pers. comm.).

FIGURES.—Figure 27g, aboral surface; Cooke (1959, pl. 16), aboral and oral surfaces, side view; Durham et al. (1966:U488), aboral and oral surfaces, side view.

Allaster Nisiyama, 1968

TYPE SPECIES.—Allaster rotundatus Nisiyama, 1968.

NUMBER OF SPECIES.—Only the type species is known.

DISTRIBUTION.—Hokkaido, Japan.

STRATIGRAPHIC OCCURRENCE.—Oligocene or Miocene.

FIGURES.—Nisiyama (1968, pls. 17, 18), aboral and oral surfaces.

Amphiope L. Agassiz, 1840

TYPE SPECIES.—Scutella bioculata Desmoulins, 1835.

NUMBER OF SPECIES.—At least six (Mortensen, 1948:413–414; Kier and Lawson, 1978:73).

DISTRIBUTION.—Europe; western Africa; India.

STRATIGRAPHIC OCCURRENCE.—Late Eocene or Oligocene to Miocene.

FIGURES.—Durham et al. (1966:U487), aboral and oral surfaces.

Amplaster Martínez, 1985

TYPE SPECIES.—Amplaster coloniensis Martínez, 1985.

NUMBER OF SPECIES.—Only the type species is known (Martínez, 1985).

DISTRIBUTION.—Known only from Uruguay.

STRATIGRAPHIC OCCURRENCE.—Upper Miocene.

FIGURES.—Figure 27l, aboral surface; Martínez (1985, figs. 1, 2), aboral and oral surfaces.

Astriclypeus Verrill, 1867

TYPE SPECIES.—Astriclypeus manni Verrill, 1867.

NUMBER OF SPECIES.—The type is the only living species. Wang (1983:116–117) recently added a new fossil species to the 3 fossil species already known from Japan.

DISTRIBUTION.—Cambodia to southern Japan.

STRATIGRAPHIC OCCURRENCE.—Durham et al. (1966:U489) give the range as Miocene to Recent, but Wang (1983:118) reports an Astriclypeus from the Late Oligocene.

FIGURES.—Figure 27j, aboral surface; Figure 29c, podial spicule; Durham et al. (1966:U487), aboral and oral surfaces.

Astrodapsis Conrad, 1856

TYPE SPECIES.—Astrodapsis antiselli Conrad, 1856.

NUMBER OF SPECIES.—Grant and Hertein (1938:68–78) list over 25 species from the middle Tertiary of California alone. Durham (1952) suggested that none of the species from Kamchatka or Sakhalin were true Astrodapsis, and relegated them to Nipponaster. With these accounted for, there are probably still in excess of 40 species ascribed to the genus Astrodapsis. Hall (1962:48) reduced the 59 named species, subspecies, and varieties to twelve.

DISTRIBUTION.—Western USA, California.

STRATIGRAPHIC OCCURRENCE.—Middle Miocene to Upper Miocene (Steven C. Beadle, pers. comm.).

FIGURES.—Figure 27f, aboral surface; Durham et al. (1966:U483), aboral and oral surfaces, side view.

Dendraster L. Agassiz, 1847

TYPE SPECIES.—Scutella excentricus Eschscholtz, 1831.

NUMBER OF SPECIES.—Raup (1956:685) reports that over 25 species and subspecies have been placed in the genus Dendraster. Mortensen (1948:381) lists four living species.

DISTRIBUTION.—Western North America, Pacific coast of Baja, Gulf of California (only as fossil?) north to southeastern Alaska.

STRATIGRAPHIC OCCURRENCE.—Latest Upper Miocene, or Pliocene to Recent (Steven C. Beadle, pers. comm.).

FIGURES.—Figure 25g, food grooves; Figure 27h, aboral surface; Durham et al. (1966:U480), aboral and oral surfaces plate patterns, food grooves.

Echinarachnius Gray, 1825

TYPE SPECIES.—Scutella parma Lamarck, 1816.

NUMBER OF SPECIES.—Grant and Hertlein (1938:56–64) list about five species that could still be referable to Echinarachnius, once those taxa now recognized as belonging to other genera such as Kewia and Vaquerosella are accounted for. Nisiyama (1968:95–104) lists seven fossil species from Japan, and Wagner (1974:109–116) three from Alaska. There are probably more than 15 fossil species attributable to this genus, as well as two known living species.

DISTRIBUTION.—Eastern North America from Labrador at least as far south as Chesapeake Bay and perhaps northern North Carolina; western North America from the Bering and Chukchi Seas along the Aleutian Islands north to Point Barrow and south to southeastern Alaska; northeast Asia, Kamchatka, Sakhalin, northern Japan.

STRATIGRAPHIC OCCURRENCE.—Upper or Middle Miocene to Recent.

FIGURES.—Figure 3b, lantern supports; Figure 11b, miliary spine; Figure 25f, food grooves; Durham et al. (1966:U483), aboral and oral surfaces.

Echinodiscus Leske, 1778

TYPE SPECIES.—Echinoglycus irregularis Leske, 1778 (= Echinodiscus bisperforatus Leske, 1778 by action of the ICZN, 1950).

NUMBER OF SPECIES.—Nisiyama (1968:132–134) lists two fossil species from Japan, but Wang (1982a:150) lists three fossil species, and adds two more (Wang, 1982a: 150, 1984a: 107). Kier (1972a:91–92) described two species from Saudi Arabia, so there are probably about nine known fossil species. Mortensen (1948:400) lists three living species.

DISTRIBUTION.—Red Sea; Madagascar and southeastern Africa; northern Indian Ocean; Philippines; Australia; New Caledonia; southern Japan.

STRATIGRAPHIC OCCURRENCE.—Wang (1984a:109) records a single specimen of a new species, Echinodiscus tiliensis, that he found “in the collection of the Museum of the Department of Geology, National Taiwan University…Probably the holotype of this new species comes from the middle or upper part of the Tachien Sandstone…as judged from the feature [sic] of the sandstone matrix (C.H. Chen, CGS, 1982, personal communication) although no calcareous nannofossils are found in the matrix” (Wang, 1984a:109). The Tachien Sandstone is “considered Eocene in age by Chen” (Wang, 1984a:109), but the “Chiayang Formation which overlies the Tachien Sandstone is Early to Middle Eocene…in age…then the Tachien Sandstone should be either Early Eocene or Late Paleocene in age” (Wang, 1984a:109). At present it would appear to be more profitable to investigate further and ascertain the suggested locality and age of strata in which the single specimen of E. tiliensis Wang was found than to revise present views (for example Durham et al., 1966; Kier, 1982) of the origins, not only of the Scutellina, but of the entire Clypeasteroida. Until corroboration is available, the attribution of Echinodiscus to the Late Paleocene or Early Eocene cannot be accepted without reservation, and the stratigraphic range of Echinodiscus is here regarded as Late Oligocene or Early Miocene to Recent.

FIGURES.—Figure 2d, Aristotle's lantern; Figure 8b, respiratory podial pores; Figure 12d, rotule; Figure 27k, aboral surface; Figure 30a, lunule; Durham et al. (1966:U487), aboral and oral surfaces, side view.

Encope L. Agassiz, 1840

TYPE SPECIES.—Encope grandis L. Agassiz, 1840.

NUMBER OF SPECIES.—From papers by Mortensen (1948:436), A.H. Clark (1946), H.L. Clark (1948), and others, a list of fifteen living species can be compiled. Approximately 25 fossil species have been described by workers such as Durham (1950), Jeannet (1928a,b) and Cooke (1961).

DISTRIBUTION.—Eastern North America as far north as North Carolina; Caribbean; Gulf of Mexico; northern and eastern South America to Rio de la Plata; western North America, Gulf of California south to Peru; Galápagos and Cocos Islands.

STRATIGRAPHIC OCCURRENCE.—Pliocene (perhaps Upper Miocene) to Recent.

FIGURES.—Figure 2c, Aristotle's lantern; Figure 21h, internal buttress system; Figure 27p, aboral surface; Durham et al. (1966:U468), aboral and oral surfaces.

Eoscutella Grant and Hertlein, 1938

TYPE SPECIES.—Scutella coosensis Kew, 1920.

NUMBER OF SPECIES.—Grant and Hertlein (1938:54–55) listed only the type species, and Mortensen (1948:386) suggested the existence of a second fossil species. Parma (1985:37–39) described one, and possibly a second new species of Eoscutella from Argentina.

DISTRIBUTION.—Western USA, California and Oregon; Patagonia, Argentina.

STRATIGRAPHIC OCCURRENCE.—Middle to Upper Eocene.

FIGURES.—Figure 9d, basicoronal plates; Figure 26d, oral surface plate pattern; Durham et al. (1966:U479), aboral and oral surfaces; Parma (1985, figs. 3, 4, pl. 1), adapical plate pattern, basicoronal plates, aboral and oral surfaces, side views.

Eoscutum Lambert, 1914

TYPE SPECIES.—Porpitella doncieuxi Lambert, 1905.

NUMBER OF SPECIES.—Mortensen (1948:231) suggests that five or six have been described.

DISTRIBUTION.—Europe, France.

STRATIGRAPHIC OCCURRENCE.—Eocene.

FIGURES.—Figure 18i, apical system; Mortensen (1948:230), aboral and oral surfaces, side view; Durham et al. (1966:U469), aboral and oral surfaces.

Faassia Schmidt, 1971

TYPE SPECIES.—Faassia globosa Schmidt, 1971 (in Schmidt and Sinel'nikova, 1971).

NUMBER OF SPECIES.—Only the type species has been described.

DISTRIBUTION.—Known only from western Kamchatka.

STRATIGRAPHIC OCCURRENCE.—Upper Miocene.

FIGURES.—Schmidt and Sinel'nikova (1971, fig. 1), aboral and oral surfaces, side view; Budin (1977, figs. 1, 2), growth rings, food grooves, oral surface plate pattern.

Iheringiella Berg, 1898

TYPE SPECIES.—Scutella patagoniensis Desor, 1847.

NUMBER OF SPECIES.—Apparently only the type species has been referred to this genus.

DISTRIBUTION.—Eastern South America, Argentina, Tierra del Fuego; Chile. Hotchkiss and Fell (1972:371) report the discovery of a fossil petaloid from Antarctica that might be referable to this genus.

STRATIGRAPHIC OCCURRENCE.—Durham (1955:171) records that the type species is from the Lower Miocene. Larrain (1984:30) extends this stratigraphic range from the Lower (Middle[?]) Eocene to the Miocene(?).

FIGURES.—Figure 9e, basicoronal plates; Figure 25c, food grooves; Figure 26e, oral surface plate pattern; Figure 28a, petaloid; Durham et al. (1966:U484), aboral surface.

Karlaster Marchesini Santos, 1958

TYPE SPECIES.—Karlaster pirabensis Marchesini Santos, 1958.

NUMBER OF SPECIES.—Only the type species is known.

DISTRIBUTION.—Brazil.

STRATIGRAPHIC OCCURRENCE.—Miocene.

FIGURES.—Figure 26n, oral surface plate pattern; Durham et al. (1966:U484), oral surface.

Kewia Nisiyama, 1935

TYPE SPECIES.—Scutella blancoensis Kew, 1920.

NUMBER OF SPECIES.—Wagner (1974:116–119) added four new Alaskan species to the six known Japanese species, and there are six more described from western North America in deposits south of Alaska, for a total of 16 named species (Linder et al., 1988).

DISTRIBUTION.—Western North America, Oregon, Washington, Alaska, California; Japan.

STRATIGRAPHIC OCCURRENCE.—Upper Eocene to Pliocene.

FIGURES.—Figure 25e, food grooves; Durham et al. (1966:U483), aboral and oral surfaces, side view.

Kieria Mihály, 1985

TYPE SPECIES.—Kieria semseyana Mihály, 1985.

NUMBER OF SPECIES.—Only the type species has been described. It is not clear whether the specimens upon which the description is based represent adult echinoids. Their small size suggests that they are juvenile astriclypeids, possibly Amphiope, but the presence or absence of gonopores cannot be ascertained from the plates or description in Mihály (1985:255, 261).

DISTRIBUTION.—Known only from a locality near Budapest, Hungary.

STRATIGRAPHIC OCCURRENCE.—Upper Badenian, Middle Miocene.

FIGURES.—Mihály (1985, pl. 4), aboral and oral surfaces.

Lenita Desor, 1847

TYPE SPECIES.—Echinites patellaris Leske, 1778.

NUMBER OF SPECIES.—One, or possibly two species can be referred to this genus.

DISTRIBUTION.—Europe, France, Belgium, Germany.

STRATIGRAPHIC OCCURRENCE.—Eocene.

FIGURES.—Figure 21e, internal buttress system; Figure 22c, oral surface tuberculation; Durham et al. (1966:U470), aboral and oral surfaces; Kier (1968, pl. 1), aboral and oral surfaces.

Leodia Gray, 1852

TYPE SPECIES.—Echinodiscus sexiesperforatus Leske, 1778.

NUMBER OF SPECIES.—Only the type species can legitimately be referred to this genus.

DISTRIBUTION.—Restricted to biogenic (carbonate) substrates of Florida Keys; Central and South America; some Caribbean Islands.

STRATIGRAPHIC OCCURRENCE.—Pliocene to Recent. A recently discovered specimen in the NMNH suggests that this genus occurs in the Pleistocene of Florida. Previous references to fossil Leodia are actually attributable to Mellita caroliniana (Ravenel).

FIGURES.—Figure 11c, miliary spine; Figure 25l, pressure drainage channels and food grooves; Figure 26p, oral surface plate pattern; Figure 27n, aboral surface; Figure 29f, aboral primary spine; Durham et al. (1966:U486), oral surface plate pattern.

Mellita L. Agassiz, 1841

TYPE SPECIES.—Echinodiscus quinquiesperforatus Leske, 1778.

NUMBER OF SPECIES.—Four living species were recognized by Mortensen (1948:422). Workers such as Durham (1961) and Kier (1963; 1972b) recognized five or six fossil species. The genus is currently being revised by Harold and Telford (in prep.).

DISTRIBUTION.—Restricted to terrigenous (siliceous) substrates of eastern North America as far north as Virginia; Caribbean; Gulf of Mexico; northern South America to Sao Paulo, Brazil; Gulf of California; west side of Isthmus of Panama.

STRATIGRAPHIC OCCURRENCE.—Pliocene (perhaps Upper Miocene) to Recent.

FIGURES.—Figure 12c, rotule; Figure 26o, oral surface plate pattern; Figure 27m, aboral surface; Figure 29d, podial spicule; Figure 29e, aboral primary spine; Figure 30b, lunule; Durham et al. (1966:U486), aboral and oral surfaces, side view.

Mellitella Duncan, 1889

TYPE SPECIES.—Encope stokesii L. Agassiz, 1841.

NUMBER OF SPECIES.—The type is the only living species recognized to date. One or two fossils are known (Durham, 1950; Cooke, 1961).

DISTRIBUTION.—Gulf of California; west side of Isthmus of Panama; northern South America.

STRATIGRAPHIC OCCURRENCE.—Pliocene (perhaps Upper Miocene) to Recent.

FIGURES.—Figure 27o, aboral surface; Durham et al. (1966:U486), aboral and oral surfaces.

Mennerella Schmidt, 1971

TYPE SPECIES.—Mennerella ovata Schmidt, 1971 (in Schmidt and Sinel'nikova, 1971).

NUMBER OF SPECIES.—Only the type species has been described.

DISTRIBUTION.—Known only from Kavran-Utkholok Bay, western Kamchatka.

STRATIGRAPHIC OCCURRENCE.—Upper Miocene.

FIGURES.—Schmidt and Sinel'nikova (1971, fig. 1), aboral and oral surfaces, side view. Not in key.

Merriamaster Lambert, 1911

TYPE SPECIES.—Scutella perrini Weaver, 1908.

NUMBER OF SPECIES.—At least three species are known (Durham, 1978; Durham and Morgan, 1978:301–303).

DISTRIBUTION.—Central and southern California; Baja California.

STRATIGRAPHIC OCCURRENCE.—Upper Miocene to Upper Pliocene.

FIGURES.—Figure 25h, food grooves; Durham et al. (1966:U480), aboral and oral surfaces, side view, food grooves.

Monophoraster Lambert and Thiéry, 1921

TYPE SPECIES.—Monophora darwini Desor, 1847.

NUMBER OF SPECIES.—Three species have been described (Mortensen, 1948:419).

DISTRIBUTION.—South America, Argentina. Although Durham (1955:169) was probably in error in recording this genus from Chile, Larrain provides “the first documented record of M. darwini in the Chilean Tertiary” (1984:27).

STRATIGRAPHIC OCCURRENCE.—Miocene, possibly Upper Eocene (Larrain, 1984).

FIGURES.—Figure 6d, petaloid plate pattern; Figure 21g, internal buttress system; Figure 25k, pressure drainage channels and food grooves; Figure 26m, oral surface plate pattern; Durham et al. (1966:U484), aboral and oral surfaces.

Mortonella Pomel, 1883

TYPE SPECIES.—Scutella quinquefaria Say, 1825.

NUMBER OF SPECIES.—Probably at least two.

DISTRIBUTION.—Southeastern United States; Cuba; Gulf of Mexico.

STRATIGRAPHIC OCCURRENCE.—Upper Eocene.

FIGURES.—Figure 27a, aboral surface; Durham et al. (1966:U479), aboral and oral surfaces, side view.

Nipponaster Durham, 1952

TYPE SPECIES.—Astrodapsis nipponicus Nisiyama, 1934.

NUMBER OF SPECIES.—Probably three or four species can be ascribed to this genus, following Durham's (1955) revision, plus two or three species originally described as members of the genus Pseudoastrodapsis, herein considered synonymized with Nipponaster following Durham et al. (1966:U482).

DISTRIBUTION.—Japan; Sakhalin(?) and Kamchatka(?).

STRATIGRAPHIC OCCURRENCE.—Miocene to Pliocene(?).

FIGURES.—Durham et al. (1966:U483), aboral and oral surfaces, oral surface plate pattern.

Parascutella Durham, 1953

TYPE SPECIES.—Scutella leognanensis Lambert, 1903.

NUMBER OF SPECIES.—Probably about ten (Durham, 1953b: 349–350; Kier and Lawson, 1978:66–67).

DISTRIBUTION.—Europe, France.

STRATIGRAPHIC OCCURRENCE.—Miocene.

FIGURES.—Figure 26g, oral surface plate pattern; Figure 27d, aboral surface; Figure 28b, petaloid; Durham et al. (1966:U478), aboral and oral surfaces.

Parmulechinus Lambert, 1910

TYPE SPECIES.—Scutella agassizi Oppenheim, 1902.

NUMBER OF SPECIES.—Five or six known (Durham, 1953b:349).

DISTRIBUTION.—Europe, northern Africa, Malta.

STRATIGRAPHIC OCCURRENCE.—Lower Miocene.

FIGURES.—Figure 26h, oral surface plate pattern; Figure 27e, aboral surface; Durham et al. (1966:U478), aboral and oral surfaces.

Periarchus Conrad, 1866

TYPE SPECIES.—Sismondia alta Conrad, 1865.

NUMBER OF SPECIES.—At least three (Durham, 1955:155; Kier, 1980:40–43).

DISTRIBUTION.—Southeastern and eastern USA as far north as North Carolina; Cuba; Gulf of Mexico.

STRATIGRAPHIC OCCURRENCE.—Upper Eocene.

FIGURES.—Figure 10b, keel and peristomial point in food groove; Figure 21c, internal buttress system; Figure 5b, food grooves; Figure 26b, oral surface plate pattern; Figure 27b, aboral surface; Cooke (1959, pls. 12–14), aboral and oral surfaces, side view; Durham et al. (1966:U479), aboral and oral surfaces, side view.

Porpitella Pomel, 1883

TYPE SPECIES.—Cassidulus Hayesianus Desmoulins, 1837.

NUMBER OF SPECIES.—At least three species have been ascribed to this genus (H.L. Clark, 1937; Mortensen, 1948:232–233).

DISTRIBUTION.—Europe, France, Alabama(?).

STRATIGRAPHIC OCCURRENCE.—Eocene.

FIGURES.—Figure 18h, apical system; Mortensen (1948:231), aboral and oral surfaces, side and rear views; Durham et al. (1966:U470), aboral surface.

Protoscutella Stefanini, 1924

TYPE SPECIES.—Scutella mississippiensis Twitchell, 1915.

NUMBER OF SPECIES.—Mortensen (1948:390) lists five fossil species.

DISTRIBUTION.—Southeastern and eastern USA as far north as North Carolina; Gulf of Mexico.

STRATIGRAPHIC OCCURRENCE.—Middle to Upper Eocene.

FIGURES.—Figure 25a, food grooves; Figure 26a, oral surface plate pattern; Cooke (1959, pl. 15), aboral and oral surfaces, side view; Durham et al. (1966:U479), aboral and oral surfaces.

Remondella Durham, 1955

TYPE SPECIES.—Clypeaster gabbi Rémond, 1863.

NUMBER OF SPECIES.—Three (Wagner, 1974:120–121; Budin, 1977:446).

DISTRIBUTION.—Western USA, California and Alaska. There are reports that Remondella is also found in Kamchatka (Budin, 1977:446).

STRATIGRAPHIC OCCURRENCE.—Upper Miocene in California to Lower Pliocene in Alaska (the Alaskan deposits might also be Upper Miocene, according to Steven C. Beadle, pers. comm.). It is found in the Upper Miocene in Kamchatka (Budin, 1977:446).

FIGURES.—Figure 25j, food grooves; Figure 26l, oral surface plate pattern; Durham et al. (1966:U484), oral surface plate pattern; Wagner (1974, pl. 3), aboral and oral surfaces, side view.

Samlandaster Lambert and Thiéry, 1914

TYPE SPECIES.—Scutella germanica von Beyrich, 1847.

NUMBER OF SPECIES.—Only the type species is known.

DISTRIBUTION.—Europe, Poland.

STRATIGRAPHIC OCCURRENCE.—Upper Eocene.

FIGURES.—Durham et al. (1966:U489), oral surface, internal view of test. Not in key.

Scaphechinus A. Agassiz, 1863

TYPE SPECIES.—Scaphechinus mirabilis A. Agassiz, 1863.

NUMBER OF SPECIES.—Nisiyama (1968:110–111) lists two fossil species, and Mortensen (1948:374) lists three living species.

DISTRIBUTION.—Northeastern Asia, Vostok Bay; Japan; Taiwan (as a fossil only?).

STRATIGRAPHIC OCCURRENCE.—Pliocene to Recent.

FIGURES.—Figure 10c, keel and peristomial point in food groove; Figure 25i, food grooves; Figure 26k, oral surface plate pattern; Figure 29b, spines from oral surface; Durham et al. (1966:U480), aboral and oral surfaces.

Scutaster Pack, 1909

TYPE SPECIES.—Scutaster andersoni Pack, 1909.

NUMBER OF SPECIES.—Two or three (Durham, 1955:180).

DISTRIBUTION.—Western USA, California.

STRATIGRAPHIC OCCURRENCE.—Lower Oligocene to Lower or Middle Miocene (Steven C. Beadle, pers. comm.).

FIGURES.—Durham et al. (1966:U488), aboral surface, oral surface plate pattern, food grooves.

Scutella Lamarck, 1816

TYPE SPECIES.—Echinodiscus subrotundus Leske, 1778.

NUMBER OF SPECIES.—Durham (1953b:349) feels that only the type species can be referred to this genus with any certainty. Kier and Lawson (1971:66–67) list more than 20 species of Scutella, some of which are probably not true Scutella (sensu Durham, 1953b). The New World species are unlikely to be Scutella, and some of the European taxa will undoubtedly turn out to be Parascutella or Parmulechinus (sensu Durham, 1953b). Of the “Scutella” listed by Kier and Lawson (1971:66–67), seven were described after Durham's (1953b) attempt to unravel the complicated taxonomy of this group. The placement of these taxa may or may not reflect Durham's recommendations.

DISTRIBUTION.—Malta, Austria, Italy, Hungary.

STRATIGRAPHIC OCCURRENCE.—Upper Oligocene or Lower Miocene.

FIGURES.—Figure 9f, basicoronal plates; Figure 26f, oral surface plate pattern; Figure 27c, aboral surface; Durham et al. (1966:U476), aboral and oral surfaces, food grooves, side view.

Scutellaster Cragin, 1895

TYPE SPECIES.—Scutella interlineata Stimpson, 1856.

NUMBER OF SPECIES.—Approximately ten (Wagner, 1974:118).

DISTRIBUTION.—Western North America, central California to Oregon; Alaska; Sakhalin(?).

STRATIGRAPHIC OCCURRENCE.—Upper Miocene to Pleistocene (Steven C. Beadle, pers. comm.)

FIGURES.—Figure 25d, food grooves; Durham et al. (1966:U481), aboral surface, oral surface plate pattern, food grooves.

Scutellina L. Agassiz, 1841

TYPE SPECIES.—Scutella lenticularis Lamarck, 1816.

NUMBER OF SPECIES.—Mortensen (1948:229) suggests that there are at least twelve species of Scutellina, all fossil.

DISTRIBUTION.—Europe, northern Africa.

STRATIGRAPHIC OCCURRENCE.—Middle Eocene. Engel's (1976:55–56) intimation that his new species of Scutellina is from the Cretaceous is suspect, and he himself suggests that the specimens “may belong to remaniated Eocene material.”

FIGURES.—Figure 7f, plate pattern adjacent to apical system; Figure 21d, internal buttress system; Figure 26c, oral surface plate pattern; Durham et al. (1966:U470), aboral and oral surfaces.

Scutulum Tournouer, 1869

TYPE SPECIES.—Scutulum parisiense Tournouer, 1869.

NUMBER OF SPECIES.—Only the type species is known.

DISTRIBUTION.—Europe, France.

STRATIGRAPHIC OCCURRENCE.—Oligocene.

FIGURES.—Durham et al. (1966:U489), aboral surface. Not in key.

Sinaechinocyamus Liao, 1979

TYPE SPECIES.—Sinaechinocyamus planus Liao, 1979.

NUMBER OF SPECIES.—Only the type species is presently contained in this genus.

DISTRIBUTION.—Yellow Sea.

STRATIGRAPHIC OCCURRENCE.—Recent.

FIGURES.—Liao (1979, figs. 1–4, pl. 1), periproct, peristome, spines, aboral and oral surfaces.

Taiwanaster Wang, 1984

TYPE SPECIES.—Taiwanaster mai Wang, 1984.

NUMBER OF SPECIES.—Three. The type is the only living species (Wang, 1984b: 134–151).

DISTRIBUTION.—Taiwan.

STRATIGRAPHIC OCCURRENCE.—Early Pliocene to Recent

FIGURES.—Figure 21f, internal buttress system; Wang (1984b, figs. 4–9, pls. 1–8), aboral and oral surfaces, spination, aboral and oral surface plate patterns, internal views of test, lantern supports, apical system, tubercles, podial pores.

Tenuiarachnius Durham, 1955

TYPE SPECIES.—Echinarachnius gabbii kleinpelli Grant and Hertlein, 1938.

NUMBER OF SPECIES.—Apparently only the type species is known.

DISTRIBUTION.—Western USA, California.

STRATIGRAPHIC OCCURRENCE.—Upper Miocene.

FIGURES.—Durham et al. (1966:U484), aboral surface, oral surface plate pattern.

Vaquerosella Durham, 1955

TYPE SPECIES.—Scutella andersoni Twitchell, 1915.

NUMBER OF SPECIES.—Four or five (Durham, 1955:166–167; 1957:630–631).

DISTRIBUTION.—Western USA, California, Oregon; Mexico; Baja California.

STRATIGRAPHIC OCCURRENCE.—Middle Miocene.

FIGURES.—Figure 26j, oral surface plate pattern; Durham et al. (1966:U484), aboral surface, crai surface plate pattern.

Illustrated Glossary

To find a diagram that illustrates a given term, look up that term on the alphabetical list below. The figure(s) indicated show(s) the structure referred to by the term. Although not all of the terms in the following list occur in the text of the key, they are illustrated in the glossary (Figures 31–34) for general reference. In Figure 31, comparisons are made between the general test morphology of a typical clypeasteroid, and that of a related form, an echinolampadid cassiduloid.

Aboral surface (Figure 31)

Accessory podium (Figure 33a)

Adradial suture (Figure 31)

Ambitus (Figure 31)

Ambulacral buttress (Figure 34b)

Ambulacral pillar (Figure 34b)

Ambulacrum (Figure 31)

Anus (Figure 32b)

Apex (Figure 31)

Apical system (Figure 32a)

Aristotle's lantern (Figure 34a)

Auricles (Figure 34b)

Basicoronal plates (Figure 31)

Bourrelet (Figure 31)

Buccal podium (Figure 32b)

Circumferential suture (Figure 31)

Conjugation groove (Figure 32a)

Demiplate (Figure 32a)

Discontinuous interambulacrum (Figure 31)

Epiphysis (Figure 34a)

External wing (Figure 34a)

Food groove (Figures 32b, 33b)

Food groove podium (Figure 32b)

Geniculate field (Figure 33b)

Gonopore (Figure 32a)

Hinge of pedicellaria (Figure 33c)

Inner pore (Figure 32a)

Interambulacral buttress (Figure 34b)

Interambulacrum (Figure 31)

Internal wing (Figure 34a)

Interporiferous zone (Figure 32a)

Interpyramidal muscle (Figure 34a)

Interradial suture (Figure 31)

Keel (Figure 32b)

Labial process (Figure 34a)

Lantern supports (Figure 34b)

Large food groove podium (Figure 32b)

Locomotory field (Figure 33b)

Long barrel-tipped podium (Figure 33b)

Lovén's system (Figure 31)

Lunule (Figure 32b)

Mouth (Figure 32b)

Neck of pedicellaria (Figure 33c)

Ocular plate (Figure 32a)

Oral surface (Figure 31)

Outer pore (Figure 32a)

Pedicellariae:

Bidentate (Figure 33c)

Biphyllous (Figure 33c)

Globiferous (Figure 33c)

Ophicephalous (Figure 33c)

Tridentate (Figure 33c)

Triphyllous (Figure 33c)

Periproct (Figure 31)

Periproctal membrane (Figure 32b)

Peristome (Figure 31)

Peristomial membrane (Figure 32b)

Peristomial point (Figure 32b)

Perradial suture (Figure 31)

Petaloid (Figures 31, 32a)

Phyllode (Figure 31)

Pressure drainage channel (Figure 33b)

Pyramid (Figure 34a)

Respiratory podium (Figure 33a)

Rotule (Figure 34a)

Short barrel-tipped podium (Figure 33b)

Sphaeridial chamber (Figure 32b)

Sphaeridium (Figure 32b)

Spines:

Aboral primary (Figure 33a)

Geniculate (Figure 33b)

Locomotory (Figure 33b)

Miliary (Figure 33a)

Stem of pedicellaria (Figure 33c)

Stereom (Figure 33a)

Supra-alveolar process (Figure 34a)

Test (Figure 33a)

Tooth (Figure 34a)

Tooth slide (Figure 34a)

Tubercle (Figure 33a)

Valve of pedicellaria (Figure 33c)

FIGURE 31.—Terminology associated with plate patterns on aboral and oral surfaces of an echinolampadoid, Echinolampas depressa Gray, and a generalized clypeasteroid. (Paired interambulacral plate columns stippled; paired ambulacral plate columns unshaded; perradial and interradial sutures numbered according to Lovén's system: ambulacra in Roman numerals, interambulacra in Arabic. (In all cases except for profiles, anterior is towards top of page.)

FIGURE 32.—Terminology associated with (a), petaloids and apical systems of a generalized clypeasteroid; and (b), peristomes, periprocts, food grooves, and sphaeridia of a generalized clypeasteroid.

FIGURE 33.—Terminology associated with a), external features on aboral surface of a generalized clypeasteroid; (b), external features on the oral surface of a generalized clypeasteroid; and (c), pedicellariae of a generalized clypeasteroid.

FIGURE 34.—Terminology associated with (a), the Aristotle's lantem of a clypeasteroid, Clypeaster rosaceus Linné, legend at lower left, numbered according to Lovén's system; and (b) lantem supports and internal buttress systems of a generalized clypeasteroid.

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1972b. Upper Miocene Echinoids from the Yorktown Formation of Virginia and Their Environmental Significance. Smithsonian Contributions to Paleobiology. 13:1–41, figures 1–7, plates 1–10.

1980.   The Echinoids of the Middle Eocene Warley Hill Formation, Santee Limestone, and Castle Hayne Limestone of North and South Carolina. Smithsonian Contributions to Paleobiology, 39:1–102, figures 1–26, plates 1–22.

1982.   Rapid Evolution in Echinoids. Palaeontology, 25(1):1–9, figures 1–3, plates 1–2.

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1984.   The Fossil and Recent Shallow Water Irregular Echinoids from Chile. 235 pages. Doctoral dissertation, University of Southern California, Los Angeles, California.

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1979.   A New Genus of Clypeasteroid Sea-urchin from Huang Hai. Oceanologia et Limnologia Sinica, 10(1):67–72, figures 1–4, plate 1.

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