Handbook of Zoology. Volume 2 Echinoidea: with bilateral symmetry. Irregularia 9783110368536, 9783110371697

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Handbook of Zoology Echinodermata Echinoidea Volume 2: Irregularia

Handbook of Zoology Founded by Willy Kükenthal Editor-in-chief Andreas Schmidt-Rhaesa

Echinodermata Edited by Andreas Schmidt-Rhaesa

DE GRUYTER

Echinoidea

Volume 2: Echinoidea with bilateral symmetry. Irregularia Heinke Schultz

DE GRUYTER

Author Heinke Schultz Dorfstr. 32 25485 Hemdingen, Germany Scientific Editor Andreas Schmidt-Rhaesa Centrum für Natukunde (CeNak) Universität Hamburg Zoologisches Museum Martin-Luther-King-Platz 3 20146 Hamburg, Germany

ISBN 978-3-11-037169-7 e-ISBN (PDF) 978-3-11-036853-6 e-ISBN (EPUB) 978-3-11-038600-4 ISSN 2193-2824 Library of Congress Cataloging-in-Publication Data A CIP catalogue record for this book is available from the Library of Congress. Bibliografic information published by the Deutsche ­Nationalbibliothek The Deutsche Nationalbibliothek lists this publication in the Deutsche Nationalbibliografie; detailed bibliographic data are available in the Internet at http://dnb.dnb.de. © 2017 Walter de Gruyter GmbH & Co. KG, Berlin/Boston Typesetting: Compuscript Ltd. Shannon, Ireland Printing and Binding: Hubert & Co. GmbH & Co. KG, Göttingen ∞ Printed on acid-free paper Printed in Germany www.degruyter.com

Foreword The present volume follows and complements volume 1, published in 2015. Whereas volume 1 presented a brief introduction and the “regular” sea urchins, this volume treats the “irregular” sea urchins. Irregular sea urchins fascinate by their bilateral symmetry and their endobenthic mode of life. After a brief introduction and an overview on the classification, species are described and illustrated in systematic order. For a brief introduction to echinoids (sea urchins) and a glossary, the reader is referred to volume 1. The two Echinoidea volumes in the Handbook of Zoology series were generated from a three-volume

book publication, Sea Urchins: A Guide to Worldwide Shallow Water Species (published in 2005 by Heinke & Peter Schultz Partner Scientific Publications), followed by the supplement volumes, Sea Urchins II: Worldwide Irregular Deep Water Species (2009) and Sea Urchins III: Worldwide Regular Deep Water Species (2011). All three volumes had a limited distribution and are out of print. Andreas Schmidt-Rhaesa Chief and Volume Editor

Contents Foreword

v

Abbreviations

3.3.1 viii

1 Introduction 2 Classification

3.3.1.1 3.3.1.2

1 3

3 Taxonomy: systematic description Infraclass Acroechinoidea Smith, 1981 7 3.1 Irregularia Latreille, 1825 7 3.1.1 Order Echinoneoida H.L. Clark, 1925 7 3.1.1.1 Family Echinoneidae L. Agassiz & Desor, 1847 7 3.2 Superorder Neognathostomata A.B. Smith, 1981 9 3.2.0.1 Family Apatopygidae Kier, 1962 10 3.2.1 Order: Cassiduloida Claus, 1880 11 3.2.1.1 Family Cassidulidae L. Agassiz & Desor, 1847 11 3.2.1.2 Family Echinolampadidae Gray, 1851 18 3.2.1.3 Family Neolampadidae Lambert, 1918 29 3.2.2 Order Clypeasteroida L. Agassiz, 1835 34 3.2.2.1 Family Arachnoididae Duncan, 1889 34 3.2.2.2 Family Clypeasteridae L. Agassiz, 1835 40 3.2.2.3 Family Echinocyamidae Lambert & Thiéry, 1914 68 3.2.2.4 Family Fibulariidae Gray, 1855 73 3.2.2.5 Family Laganidae A. Agassiz, 1873 78 3.2.2.6 Family Rotulidae Gray, 1855 88 3.2.2.7 Family Astriclypeidae Stefanini, 1911 95 3.2.2.8 Family Dendrasteridae Lambert, 1889 96 3.2.2.9 Family Echinarachniidae Lambert, 1914 98 3.2.2.10 Family Mellitidae Stefanini, 1911 100 3.2.2.11 Family Scutellidae Gray, 1825 113 3.2.2.12 Family Taiwanasteridae Wang, 1984 115 3.3 Superorder Atelostomata von Zittel, 1879 118

3.3.1.3 3.3.1.4 3.3.1.5 3.3.1.6 3.3.2 3.3.2.1 3.3.2.2 3.3.2.3 3.3.2.4 3.3.2.5 3.3.2.6 3.3.2.7 3.3.2.8 3.3.2.9 3.3.2.10 3.3.2.11 3.3.2.12 3.3.2.13 3.3.2.14 3.3.2.15 3.3.2.16

Order Holasteroida Durham & Melville, 1957 118 Family Calymnidae Mortensen, 1907 119 Family Corystusidae Foster & Philipp, 1978 120 Family Carnarechinidae Mironov, 1993 122 Family Plexechinidae Mooi & David, 1996 122 Family Pourtalesiidae A. Agassiz, 1881 125 Family Urechinidae Duncan, 1889 139 Order Spatangoida L. Agassiz, 1840 145 Family Hemiasteridae H.L. Clark, 1917 147 Family Palaeostomidae Lovén, 1868 148 Family Brissidae Gray, 1855 150 Family Palaeotropidae Lambert, 1896 192 Family Eupatagidae Lambert, 1905 198 Family Eurypatagidae Kroh, 2007 202 Family Loveniidae Lambert, 1905 212 Family Maretiidae Lambert, 1905 236 Family Spatangidae Gray, 1825 252 Family Macropneustidae Lambert, 1895 260 Family Aeropsidae Lambert, 1896 261 Family Micrasteridae Lambert, 1920 263 Family Paleopneustidae A. Agassiz, 1904 266 Family Pericosmidae Lambert, 1905 269 Family Prenasteridae Lambert, 1905 278 Family Schizasteridae Lambert, 1905 281

Paleopneustina incertae sedis A Palaeopneustina incertae sedis B 339

Literature

Acknowledgments Index

349

347

321 322

Abbreviations In the text, the names of the museums or collections are abbreviated. The following abbreviations were used: AWI

Alfred-Wegener-Institut, Helmholtz-Zentrum für Polar- und Meeresforschung, Bremerhaven, Germany

HF

Heidi Friedhoff, Norderstedt, Germany

MfN

Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitäts-Forschung, Berlin, Germany

NHM The Natural History Museum, London, UK NIWA National Institute for Water and Atmospheric Research Ltd., Wellington, New Zealand VT

Hans-Volker Thiel, Düsseldorf, Germany

ZMUC Zoological Museum, Natural History Museum of Denmark, Copenhagen, Denmark ZMH Zoologisches Museum, Centrum für Naturkunde, Hamburg, Germany ZSM

Zoologische Staatssammlung, Munich, Germany

1 Introduction Early diademataceans are the ancestors of the Irregularia. They evolved in the Lower Jurassic occupying a new habitat: they burrow more or less deeply into the sediment, where they live and feed. In adaptation to a life in this “new” environment, the morphology of these sea urchins changed radically. The test developed a bilateral symmetry, which also established the anterior-posterior axis and, simultaneously, unidirectional movement: the irregulars moved forward, with the anterior side (ambulacrum A III) ahead.

The new morphology followed a general trend. The test became flatter and the spines much shorter and more densely distributed, forming a uniform canopy, which implied less resistance to the surrounding substrate. When the animal ploughs through the sand, a mucus shield, created by the miliary spines, prevents particles from falling onto the epithelium of the animal. The tube feet of the upper side of the test were modified into flat, flap-like structures with an enlarged surface area and thin  walls.

A

B

C

D

Fig. 1.1: (A–D) Irregular sea urchins. (A) General trend of evolution in early irregular echinoids: the trend leads from a globose regular sea urchin with long spines living on the sediment to a flatter test with shorter spines and teeth still present to a flat, short-spined specimen without teeth, the periproct separated from the central apical system in an anal groove, and living infaunally (anterior to the left). Modified after A.B. Smith (1984). (B, C) Petals of irregulars: (B) On the upper side, the pore pairs in the ambulacra are concentrated in leaf-shaped densely packed bands. (C) The inner pore is subcircular, the outer one more or less elongated. The associated podia are long lobes with an extremely thin surface specialized for gaseous exchange (Peronella japonica). Scale bar is 5 mm. (D) The periproct is at the posterior end of the test (Brissus unicolor). Scale bar is 10 mm. DOI 10.1515/9783110368536-001

2 

 1 Introduction

This specialization increased the efficiency of respiration, because oxygen in the sediment is not as available as in the free water. The pore pairs in the ambulacra of the upper side are arranged in four or five leaf-shaped patterns (the petals). The periproct shifted to the posterior or even to the oral side. The feces were not released on the top of the test, in the apical shield, but posteriorly, ­clearing the rear of fecal material when burrowing.

This fundamental morphological change since the Early Jurassic (approximately 190 million years ago) generated a multiplicity of forms of echinoids (Fig. 1.1); many of which would later become extincn. Two groups succeed to date, the Clypeasteroida and the Spatangoida (Smith 1984).

2 Classification The taxa included in this volume belong, together with all regular sea urchins except for the subclass Cidaroidea, to the subclass Euechinoidea Bronn, 1860 (see volume 1: Schultz 2015). Order Echinoneoida H.L. Clark, 1925 Family Echinoneidae L. Agassiz & Desor, 1847 Genus Echinoneus Leske, 1778 Genus Koehleraster Lambert & Thiéry, 1921 Genus Micropetalon A. Agassiz & H.L. Clark, 1907 Superorder Neognathostomata Smith, 1984 Family Apatopygidae Kier, 1962 Genus Apatopygus Hawkins, 1920 Genus Porterpygus Baker, 1983 Order Cassiduloida L. Agassiz & Desor, 1847 Family Cassidulidae L. Agassiz & Desor, 1847 Genus Cassidulus Lamarck, 1801 Genus Eurhodia Haime in d’Archiac & Haime, 1853 Genus Oligopodia Duncan, 1869 Genus Rhyncholampas A. Agassiz, 1869 Family Echinolampadidae Gray, 1851 Genus Conolampas A. Agassiz, 1883 Genus Echinolampas Gray, 1825 Family Neolampadidae Lambert, 1918 Genus Anochanus Grube, 1868 Genus Aphanopora de Meijere, 1902, 1903 Genus Nannolampas Mortensen, 1948 Genus Neolampas A. Agassiz, 1869 Genus Studeria Duncan, 1891 Genus Tropholampas H.L. Clark, 1923 Order Clypeasteroida L. Agassiz, 1835 Suborder Clypeasterina L. Agassiz, 1835 Family Arachnoididae Genus Ammotrophus H.L. Clark, 1928 Genus Arachnoides Leske, 1778 Genus Fellaster Durham, 1955 Family Clypeasteridae L. Agassiz, 1835 Genus Clypeaster Lamarck, 1801 Suborder Scutellina Haeckel, 1896 Infraorder Laganiformes Desor, 1847 Family Echinocyamidae Lambert & Thiéry, 1914 Genus Echinocyamus van Phelsum, 1774 Genus Mortonia Gray, 1851

DOI 10.1515/9783110368536-002

Family Fibulariidae Gray, 1855 Genus Fibularia Lamarck, 1816 Genus Fibulariella Mortensen, 1948 Family Laganidae A. Agassiz, 1873 Genus Laganum Link, 1807 Genus Peronella Gray, 1855 Family Rotulidae Gray, 1855 Genus Heliophora L. Agassiz, 1840 Genus Rotula Schumacher, 1817 Infraorder Scutelliformes Haeckel, 1896 Family Astriclypeidae Stefanini, 1912 Genus Astriclypeus Verrill, 1867 Genus Echinodiscus Leske, 1778 Family Dendrasteridae Lambert, 1900 Genus Dendraster L. Agassiz, 1847 Family Echinarachniidae Lambert in Lambert & Thiéry, 1914 Genus Echinarachnius Gray, 1825 Family Mellitidae Stefanini, 1912 Genus Encope L. Agassiz, 1840 Genus Leodia Gray, 1852 Genus Mellita L. Agassiz, 1841 Genus Mellitella Duncan, 1889 Family Scutellidae Gray, 1825 Genus Scaphechinus A. Agassiz, 1864 Family Taiwanasteridae Wang, 1984 Genus Sinaechinocyamus Liao, 1979 Incertae sedis: Marginoproctus Budin, 1980 Superorder Atelostomata von Zittel, 1879 Order Holasteroida Durham & Melville, 1957 Family Calymnidae Mortensen, 1907 Genus Calymne Thomson, 1877 Genus Sternopatagus de Meijere, 1903 Family Carnarechinidae Mironov, 1993 Genus Carnarechinus Mironov, 1978 Family Corystusidae Foster & Philip, 1978 Genus Corystus Pomel, 1883 Family Plexechinidae Mooi & David, 1996 Genus Plexechinus A. Agassiz, 1898 Family Pourtalesiidae A. Agassiz, 1881 Genus Ceratophysa Pomel, 1883 Genus Cystocrepis Mortensen, 1907 Genus Echinocrepis A. Agassiz, 1879 Genus Echinosigra Mortensen, 1907

4 

 2 Classification Genus Helgocystis Mortensen, 1907 Genus Pourtalesia A. Agassiz, 1869 Genus Rictocystis Mironov, 1996 Genus Solenocystis Mironov, 2008 Genus Spatagocystis A. Agassiz, 1879 Family Urechinidae Duncan, 1889 Genus Antrechinus Mooi & David, 1996 Genus Cystechinus A. Agassiz, 1879 Genus Pilematechinus A. Agassiz, 1904 Genus Urechinus A. Agassiz, 1879

Order Spatangoida L. Agassiz, 1840 Family Hemiasteridae H.L. Clark, 1917 Genus Holanthus A. Agassiz, 1848 Genus Kupeia McKnight, 1974 Family Palaeostomatidae Lovén, 1868 Genus Palaeostoma Lovén, in A. Agassiz, 1872 Genus Sarsiaster Mortensen, 1950 Suborder Brissidina Stockley et al., 2005 Family Brissidae Gray, 1855 Genus Anabrissus Mortensen, 1950 Genus Anametalia Mortensen, 1950 Genus Brissalius Coppard, 2008 Genus Brissopsis L. Agassiz in L. Agassiz & Desor, 1847 Genus Brissus Gray, 1825 Genus Cionobrissus A. Agassiz, 1879 Genus Idiobrissus H.L. Clark, 1939 Genus Meoma Gray, 1851 Genus Metalia Gray, 1855 Genus Neopneustes Duncan, 1889 Genus Plagiobrissus Pomel, 1883 Genus Rhynobrissus A. Agassiz, 1872 Genus Taimanawa Henderson & Fell, 1969 Family Palaeotropidae Lambert, 1896 Genus Kermabrissoides Baker, 1998 Genus Palaeobrissus A. Agassiz, 1883 Genus Palaeotropus Lovén, 1874 Genus Paleotrema Koehler, 1914 Genus Scrippsechinus Allison, Durham & Mintz, 1967 Superfamily Spatangoidea Gray, 1825 Family Eupatagidae Lambert, 1905 Genus Eupatagus L. Agassiz in L. Agassiz & Desor, 1847 Family Eurypatagidae Kroh, 2007 Genus Elipneustes Koehler, 1914 Genus Eurypatagus Mortensen, 1948 Genus Linopneustes A. Agassiz, 1881 Genus Paramaretia Mortensen, 1950 Genus Platybrissus Grube, 1866

Family Loveniidae Lambert, 1905 Genus Araeolampas Serafy, 1974 Genus Breynia Desor in L. Agassiz & Desor, 1847 Genus Echinocardium Gray, 1925 Genus Lovenia Desor, in L. Agassiz & Desor, 1847 Genus Pseudolovenia A. Agassiz & H.L. Clark, 1907 Family Maretiidae Lambert, 1905 Genus Granobrissoides Lambert, 1920 Genus Gymnopatagus Döderlein, 1901 Genus Hemimaretia Mortensen, 1950 Genus Homolampas A. Agassiz, 1874 Genus Maretia Gray, 1855 Genus Nacospatangus A. Agassiz, 1873 Genus Pycnolampas A. Agassiz & H.L. Clark, 1907 Genus Spatagobrissus H.L. Clark, 1923 Family Spatangidae Gray, 1825 Genus Plethotaenia H.L. Clark, 1917 Genus Spatangus Gray, 1825 Family Macropneustidae Lambert, 1895 Genus Argopatagus A. Agassiz, 1879 Suborder Micrasterina Fischer, 1966 Family Aeropsidae Lambert, 1896 Genus Aeropsis Mortensen, 1907 Family Micrasteridae Lambert, 1920 Genus Cyclaster Cotteau, 1856 Genus Isopatagus Desor, 1858 Suborder Paleopneustina Markov & Solovjev, 2001 Family Paleopneustidae A. Agassiz, 1904 Genus Paleopneustes A. Agassiz, 1873 Genus Peripatagus Koehler, 1895 Genus Plesiozonus de Meijere, 1903 Family Pericosmidae Lambert, 1905 Genus Faorina Gray, 1851 Genus Pericosmus L. Agassiz in L. Agassiz & Desor, 1847 Family Prenasteridae Lambert, 1905 Genus Agassizia Valenciennes, 1846 Genus Prenaster Desor, 1853 Genus Tripylus Philippi, 1845 Family Schizasteridae Lambert, 1905 Genus Abatus Troschel, 1851 Genus Aceste Thomson, 1877 Genus Brisaster Gray, 1855 Genus Diploporaster Mortensen, 1950 Genus Hypselaster H.L. Clark, 1917 Genus Moira A. Agassiz, 1872 Genus Moiropsis A. Agassiz, 1881 Genus Ova Gray, 1825

2 Classification 

Genus Protenaster Pomel, 1883 Genus Schizaster L. Agassiz, 1835 Genus Tripylaster Mortensen, 1907 Paleopneustina incertae sedis A Genus Schizocosmus Markov, 1990

 5

Paleopneustina incertae sedis B Genus Amphipneustes Koehler, 1900 Genus Brachysternaster Larrain, 1985 Genus Genicopatagus A. Agassiz, 1879 Genus Heterobrissus Manzoni & Mazzetti, 1878 Genus Parapneustes Koehler, 1912

3 Taxonomy: systematic description Infraclass Acroechinoidea Smith, 1981 3.1 Irregularia Latreille, 1825 Today, the taxon Irregularia includes three well differentiated groups: – Order Echinoneoida – Superorder Neognathostomata: one family and two orders: Cassiduloida, Clypeasteroida – Superorder Atelostomata: two orders: Holasteroida, Spatangoida

3.1.1 Order Echinoneoida H.L. Clark, 1925 The members of this order are the most primitive irregular echinoids, widely distributed, and partly very common

A

in the Tertiary. Only three nearly related genera survive today (Fig. 3.1). The extant members of this family have nonpetaloid ambulacra, this means that their pore zones are narrow and undifferentiated. They are a sister group to the Neognathostomata and Atelostomata, which are more advanced clades that have well-developed petals with specialized pore pairs and tube feet. 3.1.1.1 Family Echinoneidae L. Agassiz & Desor, 1847 There are three genera: Echinoneus Leske, 1778. Monospecific; imperforated primary tubercles. Koehleraster Lambert & Thiéry, 1921. Monospecific; perforated primary tubercles.

D

B

C

Fig. 3.1: (A–D) Echinoneids. (A) Koehleraster abnormalis: the primary tubercles are perforated. (B) Echinoneus cyclostomus: the primary tubercles are not perforated. Photos: NHM Wien, courtesy of A. Kroh. (C) E. cyclostomus. Ile de la Réunion, France. Close-up of aboral side: the white primary tubercles are surrounded by many glassy knobs that are not set with spines. Scale bar is 2 mm. (D) E. cyclostomus. The shape is very variable. The brown specimen retained the skin. All specimens from Réunion. The spines are short and stout, and a little longer on the oral side. DOI 10.1515/9783110368536-003

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 3.1 Irregularia Latreille, 1825

Micropetalon A. Agassiz & H.L. Clark, 1907. Monospecific; oblique periproct. Literature: Westergren 1911; Hendler et al. 1995; Kroh 2005.

changed: in the Mediterranean, a part of the former Tethys, the genus went extinct. It survived in two separate regions in the Tropics, the West Indies in the west and the Indo-Pacific in the east. The genus is monospecific (Fig. 3.2).

Genus: Echinoneus Leske, 1778 The only common Recent genus, Echinoneus is reported in the Oligocene and Miocene in the Tethys, the central ocean, to the Caribbean region. After the closure of that ocean during the Miocene, the geographical distribution of Echinoneus

Echinoneus cyclostomus Leske, 1778 Test: Rather solid, very variable shape, slightly oval to greatly elongated, highly domed to flattened aborally, oral side more or less concave; margin rounded; maximum length, 50 mm.

A

C

B

F

D

G

E H

Fig. 3.2: (A–H) Echinoneids: (A, B) Echinoneus cyclostomus. (A) Test length 27 mm; Ile de la Réunion, Indian Ocean. The pore zones are narrow bands. The apical system of fused plates is small. (B) Test length 36 mm; Ile de la Réunion; Indian Ocean. Oral side: the irregularly shaped peristome and the large periproct are covered by plates. (C, D) Koehleraster abnormalis. Length 21 mm; Aldabra Island, Indian Ocean. Photos: NHM Wien, courtesy of Andreas Kroh. Scale bar is 10 mm. (E–H) Micropetalon purpureum. Length 17 mm; Molokai, Hawaii. (E) Aboral side: the ambulacra are nonpetaloid, the simple pores form a straight series. (F) Side view. (G) Oral side: both peristome and periproct are oblique. (H) The apical system is tetrabasal, the genital plates separated. The primary tubercles are not perforated. From Westergren (1911).



Apical system: Small, monobasal with four gonopores; ocular plates very small. Ambulacra: Pore zones very narrow and slightly depressed, three pore pairs per plate; pore pairs forming a narrow band; tube feet 5 to 6 mm, longer than spines, interporiferous zone might be slightly inflated. Peristome: In the center of the oral side, more or less depressed, irregularly triangular; membrane densely covered by small, smooth plates. Periproct: Large, just posterior to the peristome; elongated with sharp or less pointed ends; covered with small, spined plates. Tuberculation: Densely covered by small imperforated tubercles, each surrounded by a ring of miliary tubercles; numerous glassy knobs at the aboral side, often as large as primary tubercles. Spines: Short, stout and simple, with longitudinal wedges. Color: The spines as well as the denuded test may be whitish to light brown. The long tube feet are sometimes a bright red or rich purple, forming 10 conspicuous lines. Distribution: The species is common in the Tropics around the world, in the Caribbean, and in the Indo-­Pacific from East Africa to Hawaii and New Zealand. Bathymetric range: 5 to 570 m, but mostly in shallow water. Biology: Echinoneus has teeth only in early ontogeny. They are resorbed before the mouth opens at a test length of 4 to 5 mm, as are the paired auricles, where the muscles were fixed. These echinoids live in or on coarse sand, feeding selectively on organic material using their oral tube feet and spines, which are not specifically modified. Genus Koehleraster Lambert & Thiery, 1921 The genus is monospecific. Koehleraster abnormalis (de Loriol, 1883) Test: Solid, circumference oval, aboral side domed, oral side more or less concave; margin rounded; length may reach more than 50 mm. Apical system: Slightly anterior, small, monobasal, or tetrabasal with four gonopores; ocular plates very small. Ambulacra: Pore zones narrow, plating simple throughout with a single pore-pair; pore pairs forming a narrow band. Peristome: In the center or slightly anterior, more or less depressed, irregularly triangular; membrane densely covered with small, smooth plates. Periproct: Large, just posterior to peristome; longitudinally elongated; covered with small plates bearing spines. Tuberculation: In contrast to Echinoneus, the small and densely packed primary tubercles are perforated, each surrounded by a ring of miliary tubercles; the glassy knobs are small and inconspicuous.

3.1.1 Order Echinoneoida H.L. Clark, 1925 

 9

Spines: Short, stout and simple, with longitudinal wedges. Color: The spines may be white, or light brown to olive, the denuded test is yellowish to olive. Distribution: The species is recorded from Mauritius in the Indian Ocean to the China Sea and Hawaii; living from the littoral to a depth of 85 m. Genus Micropetalon A. Agassiz & H.L. Clark, 1907 The genus is monospecific. Micropetalon purpureum A. Agassiz & H.L. Clark, 1907 Test: Of medium size; oval in outline, low, aboral and oral side flattened, margin broadly rounded. Apical system: Subcentral, tetrabasal with four gonopores. Ambulacra: Nonpetaloid, pore zones simple and straight, not widened near the peristome; single pores from the apical system to the peristome. Peristome: Subcentral, depressed, large and obliquely triangular. Periproct: Very large, covering most of the area between mouth and posterior margin, obliquely elongated. Phyllodes and bourrelets: Not developed, no buccal pores. Primary tubercles: Not perforated, regularly distributed in a mass of small miliary tubercles. Distribution: The very rare species is recorded from Molokai, Hawaiian Islands, and the Macclesfield Bank in the South China Sea; bathymetric range 45 to 70 m. Remarks: This species is very similar to Echinoneus and Koehleraster, but differs in having an oblique periproct. In the two other genera, only the peristome is oblique. The pore zones are not depressed in Micropetalon as in Echino­ neus and Koehleraster, but flush with the test.

3.2 Superorder Neognathostomata A.B. Smith, 1981 Typical characteristics: the plating of all five interambulacra on the oral side is principally similar. Their apical disc is either tetrabasal or, more advanced, monobasal. The monobasal madreporite is probably genital plate G2, and the other genital plates have not formed (Mooi, personal communication, 2015). The ocular plates are very small (Fig. 3.3). This superorder contains three monophyletic groups: Family Apatopygidae as stem group. Order Cassiduloida with three families. Order Clypeasteroida with 12 families. Literature: Mortensen 1948; Mooi 1990a,b; Suter 1994; Kroh & Smith 2010.

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 3.2 Superorder Neognathostomata A.B. Smith, 1981

A

C

B

D

E

F

Fig. 3.3: (A–F) Neognathostomata. (A, B) Apical systems: (A) Tetrabasal, young Apatopygus, scale bar is 1 mm. (B) Monobasal, Cassidulus caribaearum. Scale bar is 1 mm. After Mooi (1990a). (C–F) Schematic cassiduloid echinoids: upper row aboral side, lower row oral side. (C) Apatopygus. (D) Cassidulus. (E) Echinolampas. (F) Neolampas. The naked areas on the oral side are shaded. Scale bars are 10 mm. Modified after Mooi (1990a).

3.2.0.1 Family Apatopygidae Kier, 1962 The family Apatopygidae is a stem group of the superorder Neognathostomata (Fig. 3.4).

with a shallow groove leading to the posterior margin. The phyllodes are single-pored, forming a simple irregular line in each half column.

Genus Apatopygus Hawkins, 1920 Three species are found in Tertiary layers from South Australia and the Chatham Islands, New Zealand. Two further extant species are known from temperate waters of the same geographic region: Apatopygus occidentalis H.L. Clark, 1938. Southwest Australia. Apatopygus recens Milne-Edwards, 1926. New Zealand.

Apatopygus occidentalis H.L. Clark, 1938. Test: May reach a length of 27 mm. Distribution: This species is endemic to the southwest coast of Australia between Bunbury and Rottnest Island; depth range 18 to 40 m. Remarks: A. occidentalis differs from A. recens by its narrower test, its smaller peristome and periproct and by its distinctly separated geographic range. The anterior pair of petals is much shorter than the posterior pair.

Test: Rather thin; in profile upper side convex, oral side flattened to also convex, but sunken toward the periproct; outline trapezoid, the posterior part broader than the anterior one. Apical system: Tetrabasal in young, monobasal in adults; slightly anterior, with four gonopores. Petals: Narrow, with narrow pore zones of the same length, distally open; anterior pair shorter than posterior pair. Oral side: Sunken toward the large, transversely elongated or lozenge-shaped mouth; posterior interambulacrum may be more or less depressed; inconspicuous bourrelets, phyllodes not widened and no specialized spines. Periproct: In a deep groove between the posterior pair of petals, a shallow furrow leading to the more or less sharp margin. Spines: On the aboral side short, longer on the oral side. Literature: Higgins 1974; Baker 1983; Holmes 1999. The two genera are characterized by having the periproct more or less positioned between the posterior petals

Apatopygus recens Milne-Edwards, 1926. Very similar to A. occidentalis. Length may reach more than 40 mm. Due to the broader test, the anterior pair of petals is only slightly shorter than the posterior one. Color: The living animals are reddish-brown, turning green after death; the bare test is grayish-white. Distribution: A. recens is confined to New Zealand, from Auckland City to the Auckland Islands in the south at depths of between 2 and 150 m. Biology: The species lives buried in coarse, shelly bottoms, approximately 30 mm deep. It seems to be an almost continuous deposit feeder. When placed on the sediment, it is able to burrow completely within 20 min. The animal moves forward 1 cm/h (Higgins 1974).

3.2.1 Order: Cassiduloida Claus, 1880 



A

D

B

E

 11

C

F

G H

Fig. 3.4: (A–H) Apatopygids. (A, B) Apatopygus occidentalis, test length 17 mm, Rottnest Island, Southwest Australia. Photo courtesy of A. Miskelly. (A) Aboral side. (B) Oral side. (C, D) Apatopygus recens. Test length 29 mm; New Zealand. (C) Aboral side: the periproct is positioned on the upper side of the test, deeply depressed. A shallow furrow leads to the posterior margin. The petals are open distally, and the pore zones narrow. (D) Oral side: the peristome is large and transversely lozenge-shaped. (E–H) Porterpygus kieri. Test length 21.9 mm, Three Kings Islands, North Island of New Zealand. After Baker (1983). (E) Aboral side. (F) Oral side. (G) Side view. (H) Apical system. Scale bar is 1 mm.

Genus Porterpygus Baker, 1983 The genus is monospecific. Porterpygus kieri Baker, 1983 Test: Small, low, in outline rounded trapezoid, widest posterior; oral side concave, strongly sunken toward peristome, flatter and angled upwards to a sharp posterior margin; length hardly 22 mm. Apical system: Anterior; tetrabasal with three gonopores. Ambulacra: Petals narrow and short, distally open. Periproct: Supramarginal, oval, slightly oblique. Peristome: Anterior, transversely elongated; phyllodes with single pores in a single series per half column; no buccal pores. Occurrence: Found off the Three Kings Islands, North Island of New Zealand; bathymetric range 90 to 300 m.

3.2.1 Order: Cassiduloida Claus, 1880 The cassiduloids were abundant during the Jurassic, Cretaceous, and Tertiary, but since the Eocene, they have dramatically decreased in number (Fig. 3.5). A total of 30 species survive today, which is only 5% of all known cassiduloids (Suter 1994). They are divided into three families: Cassidulidae L. Agassiz & Désor, 1847 Echinolampadidae Gray, 1851 Neolampadidae Lambert, 1918 Literature: Mortensen 1948; Krau 1954; Kier 1962; Gladfelter 1978; Mooi 1990a,b; Suter 1988, 1994. 3.2.1.1 Family Cassidulidae L. Agassiz & Desor, 1847 The periproct lies on the aboral side just above the ambitus, either on the posterior slope or on the more or less vertical

12 

 3.2 Superorder Neognathostomata A.B. Smith, 1981 slightly inflated; phyllodes mostly with two outer series of single pores, slightly widened toward the peristome; buccal pores present. Periproct: Supramarginal, transversely oval; short, shallow anal groove down to the posterior margin. Spines: Aborally short, on the oral side much longer and supported by much larger tubercles. Four or five species are known: Cassidulus caribaearum Lamarck, 1801. Caribbean. Cassidulus infidus Mortensen, 1948. Bahia, Brazil. Cassidulus malayanus Mortensen, 1948. Kei Islands, ­Indonesia. Cassidulus mitis Krau, 1954. Rio de Janeiro, Brazil. Krau (1960) described an additional species, Cassidu­ lus delectus, from a bay near Rio de Janeiro, a single broken specimen. Thus, there seems to exist three species at the east coast of Brazil, of which only one is fairly common in a very restricted area.

Fig. 3.5: Peristome of a cassiduloid (Echinolampas ovata): Close to the peristome, the ambulacral pores are enlarged, forming the leaf-shaped phyllodes. Each pore is associated with a specialized tube foot. Nearest to the peristome, two more or less enlarged pores are developed, which give rise to the two buccal podia. Alternating with the phyllodes, there are the bourrelets, more or less inflated interambulacral plates carrying specialized spines. Scale bar is 5 mm.

posterior end; posterior interambulacrum on oral side with large granuled naked area, extending more or less into the adoral ambulacra; bourrelets distinct. Three extant genera are included in this family: Genus Cassidulus Lamarck, 1801. Four or five species. Genus Eurhodia Haime in d’Archiac & Haime, 1853. Monospecific. Genus Oligopodia Duncan, 1889. Monospecific. Genus Cassidulus Lamarck, 1801 Test: Small size; rather low; oval, but posteriorly broader; oral side flattened, more or less concave. Apical system: Anterior, monobasal, four gonopores; few madreporic pores. Ambulacra: Petals distinct, rather straight; pore zones narrow and mostly of more or less unequal length in one ambulacrum; single pores beyond petals. Oral side: Distinctly concave along the midline; naked areas in ambulacra and interambulacrum 5 large and finely and irregularly pitted; peristome anterior, transversely pentagonal; bourrelets forming vertical walls, only

Cassidulus caribaearum Lamarck, 1801 Test: Depressed; outline ovate; posterior side with ­projection; length scarcely reaches 25 mm. Apical system: Slightly anterior, four gonopores. Ambulacra: Pore series in the petals of unequal length, less than 60 pore pairs in the anterior petal (Fig. 3.6). Oral side: Flattened and concave along the midline; large, naked median areas with irregular pits; phyllodes slightly widened, fewer than 15 pores in the anterior one. Periproct: On the aboral side below the posterior ­projection. Spines: Aborally short and uniform, much longer on the oral side. Color: In life, the sea urchin is light brown, the bare test is whitish. Distribution: The species is known from Belize to the Bahamas and from the Antilles to Barbados. It lives in very shallow water between less than 2 and 10 m. Biology: C. caribaearum is one of the few brood-protecting species in tropical waters (28°C). After spawning and ­fertilization of the yolk-rich eggs, the female groups them around the apical pole between the spines. Within 3 to 5 days, the modified larva has formed, after 5 to 7 days, metamorphosis has taken place, and 9 to 11 days after spawning, the little sea urchin is set free. The sex ratio of the populations is greater than five females to one male. The populations live in isolated patches in habitats with coarse, moderately sorted sand in agitated, shallow water (Gladfelter 1978). Remarks: C. caribaearum differs from Rhyncholampas pacificus by the smaller size and the less numerous pores in its anterior petals and phyllodes.

3.2.1 Order: Cassiduloida Claus, 1880 

A

C

 13

B

D

E

Fig. 3.6: (A–E) Cassiduloids. (A, B) Cassidulus caribaearum. Test length 23 mm, height 11 mm; St. Thomas, Caribbean. ZMUC. (A) Aboral side. The petals are open distally, the pore zones narrow. This brood-protecting species has not developed any brood-pouches (the hole in front of the peristome was created by a predator, a snail). (B) Oral side: the large naked areas are covered in irregular pits. The five pairs of buccal pores are distinct. (C–E) Cassidulus infidus. Length 25 mm, Brazil (labeled “Bahia”). (C) Aboral side. (D) Oral side. (E) Side view. Scale bar is 10 mm. Modified after Mooi (1990b).

Cassidulus infidus Mortensen, 1948 This species is known from a single specimen with a length of 10 mm labeled “Bahia, Brazil”. It has not been found again (Fig. 3.7). Mortensen found it nearly related to C. caribaearum, but he emphasized the following differences: in C. infidus, the test is narrower and higher with steeper sides, the margin is rounder, the peristome is more sunken, and the periproct is almost round, not transversely elongate as in C. caribaearum.

Cassidulus malayanus Mortensen, 1948 Test: Delicate and fragile, circumference oval, sides parallel, aboral side lowly arched; length up to 25 mm. Apical system: Only slightly anterior, small. Petals: Rather short, of about the same length, posterior pair slightly curved; pore zones of approximately equal length; the single pores beyond the petals very small and inconspicuous. Peristome: Rounded pentagonal, broader than long; phyllodes distinct, bourrelets well-developed.

14 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

A

B

C

D

E

F

Fig. 3.7: (A–F) Cassiduloids. (A–C) Cassidulus malayanus. Test length 25 mm; Kei Islands, Indonesia. ZMUC ECH 236. (A) Aboral side: the periproct is positioned high on the upper side. (B) Oral side. (C) Side view: there is no posterior hook on top of the periproct. (D–F) Cassidulus mitis. Length 30 mm, Baia de Sepetiva, Rio de Janeiro, Brazil. (D) Aboral side. (E) Oral side. (F) Side view. Scale bar is 10 mm. Modified after Mooi (1990a).

Periproct: Elongated oval, positioned rather high on the aboral side in a long, narrow, and deep groove, not overarched by the test. Tuberculation: Fine and uniform on the aboral side partly much larger on the oral side. Color: The test and the spines are white. Distribution: Only two specimens were dredged off the Kei Islands, Indonesia, at a depth of 250 to 290 m.

Cassidulus mitis Krau, 1954 The species is nearly related to C. caribaearum. It differs from that species by its more posterior periproct. The hook is only faintly developed, whereas in C. caribaearum, it is prominent. C. mitis differs from C. infidus by its lower test, the apical system being positioned more centrally. C. mitis lives endemic in some lagoons and along beaches in the vicinity of Rio de Janeiro, Brazil.

3.2.1 Order: Cassiduloida Claus, 1880 

 15

Biology: C. mitis is brood-protecting, keeping its young among the aboral spines like C. caribaearum. Because of the colder water, the development of the juveniles takes more time. Postmetamorphic individuals of an age of 62 days (diameter 0.5 mm) still retain functional teeth. It seems that they use their lantern to graze on bacterial microfilms covering the sand grains. Later in growth, the echinoids lose their lantern completely, although there is no accurate information on the age or stage of the loss. Afterward, the podia encircling the peristome are used for feeding (Contins & Ventura 2011). Literature: Freire et al. 1992; MacCord & Ventura 2004; Contins & Ventura 2011.

Periproct: High on the posterior side under a slightly overhanging hood, shallow vertical subanal sulcus down to the margin. Tubercles: On the oral side distinctly larger than aborally. Distribution: Only two specimens are known from western Suriname and Venezuela, dredged at 57 and 112 m, respectively. Biology: E. relicta lives shallowly buried in fine to coarser sand.

Genus Eurhodia Haime in d’Archiac & Haime, 1853 Approximately 20 species of Eurhodia were established from Paleocene and Eocene layers worldwide when two specimens were found in 1968 from the collection of the USNM, dredged by the RV “Pillsbury”. Mooi (1990) described them as a new species, a “living fossil”. Their nearest relatives were common in the Late Eocene of the southeast USA and the Caribbean, but in later periods, were thought to be extinct.

Oligopodia epigonus v. Martens, 1865 Test: Small and ovoid; aboral side lowly arched, posterior interambulacrum forms a low keel often terminating as a small beak above the periproct; oral side slightly concave and depressed toward the peristome; length up to 17 mm. Apical system: Slightly anterior, monobasal, four gonopores, distinctly larger in females than in males (Fig. 3.9). Ambulacra: Petals distinct, flush with the test and distally open; the two columns almost parallel and of about equal length; plates beyond the petals rather high with single, small pores; pore pairs very oblique. Oral side: Peristome longitudinally elongated, slightly anterior; bourrelets as distinct areas with specialized spines on the almost vertical walls, but not inflated; phyllodal pores arranged in two irregular series, scarcely widened at the peristomial edge; buccal pores distinct; naked areas inconspicuous. Periproct: On the truncated posterior end, more or less deep subanal groove leading down to the ambitus. Primary spines: Very short, approximately 0.5 mm long, smooth and curved. Color: Test and spines are white. Distribution: This species is found from East Africa over the Malayan region to the Bonin Islands, Tonga Islands and New Zealand. Bathymetric range: living specimens were found from 35 to 141 m, dead tests from 5 to 390 m.

Eurhodia relicta Mooi, 1990 Test: Small, more or less highly domed, oral side longitudinally concave; length up to 13.2 mm. Apical system: Slightly anterior, monobasal, madreporite with only five to six hydropores and four gonopores; ocular plates small (Fig. 3.8). Ambulacra: Petals rather short, posterior pair longest; the two columns per ambulacrum of about equal length; interporiferous zone approximately as wide as pore zones; pore pairs conjugate, outer pore round or only slightly elongated; pores beyond petals simple. Oral side: Peristome slightly anterior, longitudinally elongated; phyllodes with 7 to 10 pores in outer series, only two or three in inner series; bourrelets distinct, forming slight bulges covered by more than 25 densely arranged tubercles set with curved spines; large naked area in IA 5 and A III, conspicuously pitted.

A

B

Genus Oligopodia Duncan, 1889 The genus is monospecific.

C

D

Fig. 3.8: (A–D) Cassiduloids. (A–D) Eurhodia relicta. Length 12.5 mm; off Surinam (holotype in USNM). (A) Aboral side. (B) Oral side (naked areas shaded). (C) Side view. (D) Posterior side. Scale bar is 10 mm. Modified after Mooi (1990b).

16 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

A

B

C

D

E

Fig. 3.9: Oligopodia epigonus, female. Test length 17 mm; Jolo, Philippines. ZMUC. (A) Aboral side. (B) Oral side: the peristome is deeply sunken with almost vertical walls. (C) Side view. The periproct is positioned on the posterior side opening into a short anal groove. (D, E) Specimen with spines. Test length 16 mm; Jolo, Philippines. ZMUC. (D) Aboral side: the spines are short and uniform. (E) Oral side: the vertical wall of the peristome is set with specialized spines.

3.2.1 Order: Cassiduloida Claus, 1880 

Genus Rhyncholampas A. Agassiz, 1869 The genus is monospecific. Rhyncholampas pacificus (A. Agassiz, 1863) Test: High convex; outline regularly oval or posteriorly slightly wider; maximum length 70 mm.

A

D

 17

Apical system: Slightly anterior, four gonopores. Ambulacra: Petals long and well-developed, pore zones in paired petals of slightly different lengths; more than 70 pore pairs in anterior petals, more than 20 pores in anterior phyllodes; buccal podia large (Fig. 3.10).

B

C

E

Fig. 3.10: Rhyncholampas pacificus. Test length 43 mm; Acapulco, Mexico. MfNB Berlin. (A) Close-up of peristomial region: the inflated bourrelets in the interambulacra are set with tufts of specialized spines. (B, C) Specimen with spines. Length of test 59 mm, height 27 mm; Acapulco, Mexico. ZMH. (B) Aboral side: the spines, short and fur-like, show scattered greenish spots. Near the posterior margin, the periproct opens below a distinct roof-like projection. (C) Oral side: the spines are much longer than on the upper side. Large naked areas are developed in the posterior interambulacrum and around the phyllodes in the ambulacra. The bourrelets are distinct. (D, E) Specimen without spines. Test length 43 mm; Acapulco, Mexico. MfN Berlin. (D) Aboral side: the petals are long and slender. The anterior side of the test is damaged. (E) Oral side: the pits in the large naked area are distinctly smaller than those of Cassidulus caribaearum.

18 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

Oral side: Large naked areas in the posterior interambulacrum and in the ambulacra around the phyllodes; pits small and round; peristome anterior, broadly pentagonal; anterior bourrelets broad. Periproct: Below a projection on the posterior aboral side. Spines: Short on the aboral side, a little longer at the projection above the periproct, the oral spines are much longer. Color: The spines are brownish, and conspicuous green spots are scattered over the aboral side. The bare test is whitish. Habitat: This species lives gregariously on sandy beaches at a depth of 2 to 130 m. Distribution: R. pacificus is known along the west coast of Central America from Baja California to Panama and the Galapagos Islands. Remarks: R. pacificus differs from C. caribaearum by the larger size and the comparatively more numerous pores in its anterior petals and phyllodes. Literature: Mooi 1990.

3.2.1.2 Family Echinolampadidae Gray, 1851 This family is characterized by having the periproct just below the ambitus on the oral side. The naked area is small and inconspicuous; the bourrelets are more or less developed. The family includes two extant genera: Genus Conolampas A. Agassiz, 1883. Four species. Genus Echinolampas Gray, 1825. Ten species. They differ in the following features: Echinolampas is more elongated in outline than Conolam­ pas and has a slight posterior projection. The tuberculation in Echinolampas is denser than in Conolampas. The oral side of Echinolampas is sunken toward the peristome, in Conolampas it is flattened. Literature: Döderlein 1906; Nisiyama 1968; Thum & Allen 1975; Shigei 1986; Mooi 1990a,b; Rowe & Gates 1995. Genus Conolampas A. Agassiz, 1883 Test: Large and solid, highly vaulted, hemispherically to subconically with sharp margin, oral side flat, very slightly sunken toward peristome; circumference almost circular. Apical system: Subcentral, monobasal with four gonopores. Ambulacra: Long, distally open petals, poriferous zones narrow, pore pairs subequal, more or less circular, conjugate; plates beyond petals with single pores (Fig. 3.11). Oral side: Peristome subcentral and pentagonal, slightly wider than long; phyllodes slightly sunken; single pores,

arranged in three series per half-ambulacrum; buccal pores present; bourrelets distinct. Periproct: On the flat oral side more or less near the posterior edge, transverse. Tuberculation: Scarce but evenly distributed, short spines that are a little longer on the oral side. Color: Yellowish-brown, cleaned test white. Biology: Mooi (1990b) assumed that the species of Cono­ lampas with flat undersides do not live buried on the sediment but feed on fine substrates. Four species are included in the genus. Conolampas diomedeae Mortensen, 1948. Philippines Conolampas malayana Mortensen, 1948. Malayan Conolampas murrayana Mortensen, 1948. Maldives Conolampas sigsbei A. Agassiz, 1878. Caribbean The species are very similar. Conolampas diomedeae Mortensen, 1948 Test: Hemispherical with sharp margin, oral side perfectly flat; maximum length 90 mm. Apical system: Knob-shaped. Petals: Long, more or less unequal in length. Oral side: Naked area in posterior interambulacrum inconspicuous; peristome subcentral to slightly posterior; anterior bourrelets more developed than posterior pairs. Color: In life, the spines are yellowish, the bare test is white. Distribution: The species is reported only from the Philippines; bathymetric range 80 to 300 m. Literature: David & de Ridder 1989; Mooi 1990. Conolampas malayana Mortensen, 1948 Test: Length up to more than 90 mm. This species differs from the two others, also found in the Malayan/Philippine region, by the length of the petals, which reach only halfway down to the ambitus. In C. murrayana and C. diomedeae, the petals extend almost to the ambitus. C. mala­ yana and C. diomedeae both bear an apical knob (Fig. 3.12). Distribution: This rare species was found at the Kei Islands, Indonesia, and off Zamboanga, Philippines; at depths of 230 to 400 m. Conolampas murrayana Mortensen, 1948. The only two known specimens measure 67 and 68 mm. The species is distinguished from C. diomedeae by the broader and more lanceolate petals. Mortensen further mentioned the sparser tuberculation on the proximal oral side, which is denser in C. diomedeae. C. murrayana lacks the apical knob that is present in C. diomedeae and C. malayana. The specimens were found off the Maldives at a depth of 229 m.

3.2.1 Order: Cassiduloida Claus, 1880 

A

B

D

 19

C

E

F

Fig. 3.11: (A–C) Conolampas. Ambulacral plates. (A) Conolampas diomedeae. (B) Conolampas murrayana: the pore pairs are distinctly conjugate. (C) Conolampas sigsbei: the plates are sparsely tuberculated, the pore pairs are distant and nonconjugate. Scale bar in (A) and (B) is 4 mm, in (C) is 3 mm. After Mortensen (1948). (D–F) C. diomedeae. Test length 87 mm, Balicasag Island, Philippines. (D) Aboral side. (E) Oral side. (F) Side view. Photos courtesy of H. van Noordenburg.

Conolampas sigsbei A. Agassiz, 1878. Test: High with steep sides; length may reach 105 mm. Ambulacra: In comparison to the Indo-Pacific species, the pore pairs in the petals are more distant. Oral side: Bourrelets and phyllodes well-developed. Color: In life, yellowish, cleaned test white (Fig. 3.13). Distribution: Recorded throughout the West Indies, along the west coast of Florida, and off the northern coast of Yucatan; bathymetric range approximately 120 to 800 m. Genus Echinolampas Gray, 1825 Test: Medium to large size, more or less elongated with a slight projection posteriorly, aborally domed, sunken toward the peristome.

Apical system: Monobasal with four gonopores, positioned anterior to subcentral. Ambulacra: Five petals well-developed, straight or arched, distally open; pore pairs small, the two columns per petal of more or less unequal length; pores simple beyond petals. Oral side: Naked area in posterior interambulacrum 5 small and inconspicuous. Peristome: Subcentral, wider than long and transverse pentagonal or oval. Phyllodes: Short and more or less conspicuous, two or three single pored series per half-ambulacrum; buccal pores present; bourrelets weakly inflated with vertical sides leading to the mouth opening.

20 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

A

B

C

Fig. 3.12: Conolampas malayana. Test length 84 mm; off Zamboanga, Philippines. ZMUC. (A) Aboral side. (B) Oral side. (C) Side view: the species has a knob-like apex.

Periproct: Flush with the test, inframarginal, transversely elongated; three valve-like periproctal plates. Tubercles: Usually of the same size on aboral and oral sides, spines short. The following 10 extant species are known: Echinolampas alexandri de Loriol, 1876. Red Sea to Malayan Archipelago, Coral Sea. Echinolampas chuni Döderlein, 1905. Indonesia. Echinolampas crassa Bell, 1880. South Africa. Echinolampas depressa Gray, 1851. Caribbean. Echinolampas keiensis Mortensen, 1948. Indonesia to Philippines. Echinolampas koreana H.L. Clark, 1925. Korean Strait. Echinolampas ovata (Leske, 1778). Red Sea to Western Australia. Echinolampas rangii Desmoulins, 1837. West Africa. Echinolampas sternopetala A. Agassiz & H.L. Clark, 1907. Japan. Echinolampas sumatrana Döderlein, 1905. Indonesia. A comparison of four species of Echinolampas occurring in the Malayan region is shown in Tab. 3.1. Unfortunately, the width of variation within the species is considerable.

The fifth species, E. keiensis, is distinguished at first sight by its strongly unequal poriferous zones and the welldeveloped bourrelets. Echinolampas alexandri de Loriol, 1876 Test length may reach more than 80 mm. This species is very similar to E. ovata and is not easily distinguished from it. It differs in usually having a flatter, wider and thinner test and less pronounced phyllodes. The peristome is oval, contrary to E. ovata, where it is distinctly pentagonal. The two columns of pore pairs in each petal are unequal (Fig. 3.14). Distribution: Red Sea, Indian Ocean (Mauritius), and Malayan Archipelago; depth 8 to 365 m. The geographic range does not extend to northwest Australia like E. ovata (Rowe & Gates 1995), but two dead tests were found in northeast Australia in the Coral Sea (Miskelly 2002). Echinolampas chuni Döderlein, 1905 Test: Strong, aboral side lowly hemispherical, circumference elongated ovoid, the posterior end being distinctly rostrate; length 75 mm.

3.2.1 Order: Cassiduloida Claus, 1880 

A

 21

B

C

D

Fig. 3.13: Conolampas sigsbei. (A) Specimen with spines, oral side. Test length 79 mm; Cuba. ZMUC. The bourrelets are well developed, set with specialized spines, which spread over the peristome like a fan. (B–D) Specimen without spines. Test length 81 mm; Cuba. ZMUC. (B) Aboral side: the pore pairs are rather distant from each other and not at all conjugate. (C) Oral side: the floscelle is well developed. (D) Side view: the apical system forms a distinct knob. Tab. 3.1: Four species of Echinolampas in comparison. E. alexandri

E. chuni

E. ovata

E. sumatrana

Test rather broad, low, rounded margin

Test elongated, rostrate, low margin

Test elongated, high with rounded margin

Test rather broad, subconical with low margin

Thin, but robust

Very strong

Solid

Very strong

Apical system anterior

Apical system anterior

Apical system anterior

Apical system subcentral

Petals hardly bowed

Petals narrow, parallel-sided

Petals broad, bowed

Petals broad, parallel-sided

Peristome transversely oval

Peristome transversely subpentagonal

Peristome transversely pentagonal

Peristome transverse and lowly pentagonal

22 

A

 3.2 Superorder Neognathostomata A.B. Smith, 1981

B

C

D

E

F

3.2.1 Order: Cassiduloida Claus, 1880 

Apical system: Distinctly anterior. Petals: Parallel sided, shorter and narrower than in E. sumatrana. Oral side: Pore zones depressed; peristome subpentagonal; phyllodes inconspicuous, bourrelets rudimentary. Tuberculation: Coarser than in E. sumatrana. Remarks: During the “Valdivia” Expedition 1898/1899, the only two specimens of Echinolampas chuni were dredged in one and the same haul together with a single specimen of Echinolampas sumatrana at Station 192, west of Sumatra from a depth of 371 m. Because they differed conspicuously, Döderlein (1906) described them as two species.

A

 23

The specimens were old, naked tests, rather worn and set with encrusting organisms. No spines, podia, or pedicellariae were left. Echinolampas crassa Bell, 1880. Test: Strong; high and subconical with rather sharp edges; length up to 125 mm. Apical system: Subcentral (Fig. 3.15). Petals: Long; the pore zones in a petal being of subequal length. Oral side: Peristome pentagonal, bourrelets and phyllodes distinct.

B

C

D

Fig. 3.15: Echinolampas crassa. Length of test 121 mm, South Africa. ZMUC. (A) Aboral side: the posterior end may sometimes be slightly rostrate. (B) Side view: the test is high and subconical. (C) Oral side: the bourrelets and phyllodes are well developed. (D) Ambulacral plates. Scale bar is 4 mm; after Mortensen (1948). ◂ Fig. 3.14: (A–F) Species of Echinolampas. Echinolampas alexandri. Test length 55 mm, Philippines. Photos courtesy of H. van Noordenburg. (A) Aboral side. (B) Oral side: the peristome is transversely oval. (C) Side view. The test is lower and the upper surface is more flattened than in Echinolampas ovata. (D–F) Echinolampas chuni. Test length 75 mm, Siberut Strait, west of Sumatra, Indonesia. MfN Berlin, Holotype, Ech Nr. 5846. (D) Aboral side: the apical system is anterior. (E) Oral side: the posterior end is rostrate. (F) Side view: the upper side is hemispherically vaulted with a low margin.

24 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

Distribution: This species is restricted to South Africa; bathymetric range 12 to 500 m. Biology: This echinoid feeds on detritus by lifting particles into the mouth using the oral podia and the bourrelet spines. The size of the particles is limited, larger grains are rejected because of the fixed size of the peristome, and smaller ones are obviously difficult to handle. The species lives in ripple beds, burrowing during feeding about 30 to 50 mm deep. As the estimated growth rate per year is approximately 5 mm, this sea  urchin is likely to be long living (Thum & Allen 1975). Echinolampas depressa Gray, 1851 Test: High, upper side more or less flattened; length may reach 50 mm. Petals: Short, pore zones of strongly different length. Oral side: Phyllodes hardly widened, bourrelets inconspicuous. Distribution: The species is found in eastern Central America, southeast USA, and the West-Indies to French Guyana; living between 30 and 310 m (Fig. 3.16).

A

Echinolampas keiensis Mortensen, 1948. Test: Broadly ovoid, aboral side subconical, oral side flat; length up to 83 mm. Apical system: Slightly anterior (Fig. 3.17). Ambulacra: Pore zones in petals very unequal and curved; pore pairs conjugate. Oral side: Peristome pentagonal and sunken; phyllodes well-developed, bourrelets distinctly inflated. Distribution: Kei Islands, Indonesia and off Zamboanga, Philippines; 245 to 400 m. Remarks: Echinolampas keiensis and E. sternopetala are very similar in having strongly unequal pore zones. However, in the former, the pore pairs are conjugate but not in the latter. Echinolampas koreana H.L. Clark, 1925. Test: High, length of holotype 52 mm. Petals: Moderately short, pore zones unequal in length. Peristome: Small and sunken. Occurrence: Korean Strait and West Japan at a depth of 73 m. Remarks: This species has a higher test with broader petals than E. sternopetala.

B

C

Fig. 3.16: Echinolampas depressa. Test length 37.5 mm, height 22 mm; Florida, USA. ZMUC. (A) Aboral side and (B) oral side. The petals are short and the peristome transversely oval with poorly developed phyllodes and bourrelets. (C) Side view: the test is highly convex.

3.2.1 Order: Cassiduloida Claus, 1880 

A

B

C

D

 25

E

Fig. 3.17: (A–C) Echinolampas keiensis. Test length 62 mm; Kei Islands, Indonesia. ZMUC. (A) Aboral side of test: one pore zone has half the length of the other. (B) Oral side of test: phyllodes and bourrelets are well developed. (C) Side view: the test is subconical. (D, E) Echinolampas: ambulacral plates: (D) Echinolampas sternopetala with very few primary tubercles, pore pairs not conjugate. (E) Echinolampas keiensis with distinctly more primary tubercles, pore pairs conjugate. Scale bar is 3 mm. After Mortensen (1948).

Echinolampas ovata (Leske, 1778) Test: Strong, highly domed with thick margin; circumference egg-shaped; maximum length about 80 mm. Apical system: Distinctly anterior. Ambulacra: Petals well-developed, narrow pore zones, pore zones in the paired petals of different lengths (Fig. 3.18). Oral side: Deeply depressed toward the large, pentagonal mouth, bourrelets vertical and rather inconspicuous, phyllodes small. Periproct: Transversely elongated, just beneath the posterior margin. Spines: Uniformly short on the aboral side, longer and thicker on the oral side. Color: In life, light to chestnut brown, denuded test grayish. After death, the color of the sea urchin turns a bright green, which fades over time.

Distribution: This species is known in the tropical Indian Ocean from the Red Sea to Ceylon, Mauritius, and Western Australia at depths between 9 and 75 m. Echinolampas rangii Desmoulins, 1837 Test: Broadly ovoid, aboral side moderately vaulted; margin rounded; maximum length 103 mm. Apical system: Slightly anterior, slightly knob-shaped (Fig. 3.19). Petals: Broad, narrowing distally, but not closed; pore  zones of unequal length, especially in the anterior  pair; pores unequal, outer pore slit-like, inner one round. Oral side: Sunken toward the pentagonal peristome; phyllodes and bourrelets well-developed. Tuberculation: Aborally dense and uniform.

26 

A

 3.2 Superorder Neognathostomata A.B. Smith, 1981

B

C

D

E

F

3.2.1 Order: Cassiduloida Claus, 1880 

A

C

 27

B

D

Fig. 3.19: (A–C) Echinolampas rangii. Test length 79 mm; Dakar, Senegal. ZMUC. (A) Aboral side: the petals are broad and rather bowed, distally open. (B) Oral side: Phyllodes and bourrelets are distinct. (C) Side view: in profile, the test is subconical. (D) Ambulacral plates of E. rangii. The petals are broad, the interporiferous zone densely set with primary tubercles. The pore pairs are strongly conjugate. Scale bar is 4 mm. After Mortensen (1948).

Color: Test and spines are light brownish, cleaned tests are whitish. Distribution: This species is recorded from the Cape Verde region and Senegal, West Africa. The bathymetric range reaches from 1,570 to 1,670 m, but Mortensen (1948) also reported findings from “shallow waters”. Echinolampas sternopetala A. Agassiz & H.L. Clark, 1907. Test: Fairly strong, ovoid in outline, slightly tapering toward the posterior end; length may reach up to 79 mm.

Apical system: Anterior to subcentral (Fig. 3.20). Petals: Narrow, pore zones very unequal in length and curved, anterior ambulacrum shortest; pore pairs not conjugate. Oral side: Nearly flat, sunken toward the transversely oval peristome; phyllodes short and not widened, bourrelets rudimentary. Periproct: Broader than the peristome, positioned obliquely below the posterior end, the distal edge nearly straight. Primary tuberculation: Rather sparse.

◂ Fig. 3.18: Echinolampas ovata. (A–C) Test length 65 mm; Dahab, Gulf of Aqaba, Red Sea. (A) Aboral side: in the paired petals, the pore zones are of different lengths. The specimen has been drilled by a predating snail and died. (B) Oral side: the pentagonal mouth is deeply depressed. (C) Side view of test. (D, E) Test length 56 mm; Dahab, Gulf of Aqaba, Red Sea. (D) Aboral side: the test is covered by short and dense spines. (E) Oral side. (F) The periproct is built by three plates. Scale bar is 1 mm. After Mortensen (1948).

28 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

A

B

C

D

E

F

3.2.1 Order: Cassiduloida Claus, 1880 

Color: In living specimens, the test and spines have a yellow tinge. Distribution: This species is endemic to Japan, known only from Sagami Bay southward to the Kagoshima Gulf; bathymetric range: 100 to 500 m. Echinolampas sumatrana Döderlein, 1905 The only known specimen is a rather worn, encrusted, naked test (see also E. chuni). Test: Strong, aboral side rounded conical; outline broadly ovoid, tapering to the posterior end. Apical system: Subcentral. Petals: Longer than in E. chuni and broader, parallel-sided; the pore zones only slightly unequal in each petal (Fig. 3.21). Oral side: Depressed, peristome transversely pentagonal, distinct phyllodes, bourrelets hardly inflated; periproct below the posterior margin.

A

 29

Occurrence: A single specimen found off the west coast of Sumatra, Indonesia, at a depth of 371 m together with two specimens of E. chuni. 3.2.1.3 Family Neolampadidae Lambert, 1918 The members of this family are characterized by their apetaloid ambulacra. The plates are simple, each perforated by a single, small pore, which may be in some species aborally rudimentary or even entirely lacking. All species display sexual dimorphism, the females having either large gonopores or a marsupium. Literature: Mortensen 1948; McKnight 1968; Baker & Rowe 1990; Mooi 1990b; Suter 1994, 1996; Holmes 1995. Six genera are included in this family, each with a single extant species (Tab. 3.2): Genus Anochanus Grube, 1868. China Sea. Genus Aphanopora de Meijere, 1903. Sulu and Tasman Seas.

B

C

Fig. 3.21: Echinolampas sumatrana. Test length 75 mm, Siberut Strait, west of Sumatra, Indonesia. MfN Berlin Holotype, Ech Nr. 5845. (A) Aboral side. (B) Oral side: the phyllodes are conspicuous. (C) Side view.

◂ Fig. 3.20: Echinolampas sternopetala. (A–C) Test length 68 mm, Sagami Bay, Japan. ZMUC. (A) Aboral side: the narrow ambulacra with strongly unequal pore zones widen conspicuously toward the margin. (B) Oral side: the peristome is deeply sunken. (C) Side view. (D) Monobasal apical system: there is a single large, fused plate with hydropores, four gonopores, and five small ocular plates. Scale bar is 2 mm; after Mortensen (1948). (E, F) Specimen with spines. Test length 71 mm, Sagami Bay, Japan. ZMUC. (E) Aboral side: the spination is dense and uniform. (F) Oral side.

30 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

Tab. 3.2: Six genera from the family Neolampadidae in comparison. Anochanus sinensis

Aphanopora echinobrissoides

Nannolampas tenera

Neolampas rostellata

Tropholampas loveni

?

Apical disc probably tetrabasal

Apical disc tetrabasal

Apical disc monobasal

Apical disc monobasal

?

Two gonopores

Two gonopores

Three gonopores

Four gonopores

Simple pores up to apex

Simple pores up to apex

Simple pores up to apex

Simple pores up to apex

No pores aborally

Female with marsupium

Gonopores large in females, small in males

Gonopores large in females, small in males

Gonopores large in females, small in males

Female with marsupium

Periproct aborally in groove

Periproct aborally in groove

Periproct on posterior face Periproct on posterior face Periproct on posterior face

?

Weak bourrelets

No bourrelets

Genus Nannolampas Mortensen, 1948. Timor Sea. Genus Neolampas A. Agassiz, 1869. West Indies to the East Atlantic, Mediterranean. Genus Studeria Duncan, 1891. Indonesia to Tasman Sea. Genus Tropholampas H.L. Clark, 1923. South Africa. Genus Anochanus Grube, 1868 The genus is monospecific. Anochanus sinensis Grube, 1868 The holotype and only known specimen reached a length of 9.5 mm. It was poorly described and never completely figured, until it was subsequently lost. Description according to Mortensen (1948): Test: Small form, in shape resembling A. recens. Apical system: Nearly central, forming a deep marsupium containing embryos. Ambulacra: Apparently with simple plates, not petaloid aborally; small, simple pores continue from the peristome to the apical system. Peristome: Slightly anterior. Periproct: Aborally in a groove? Occurrence: A single specimen has been dredged in the China Sea, depth not recorded. Remarks: It is a case of speculation if Aphanopora is the male of the marsupiate Anochanus. Mortensen and later McKnight (1968) and Mooi (1990b) thought it possible, but Holmes (1995) refutes this suggestion after having found fossil specimens of Aphanopora in southern Australia. In his material, there are females with large gonopores and males with distinctly smaller ones. That means the sexual dimorphism appears to be expressed in the different size of the gonopores, not in the presence or absence of marsupia. Genus Aphanopora de Meijere, 1902 The genus is monospecific.

Weak bourrelets

No bourrelets

Aphanopora echinobrissoides de Meijere, 1903 In shape, resembling A. recens, but differing at first sight by its two gonopores (A. recens has four genital pores). Test: Thin, low arched; maximum recorded length 15 mm. Apical system: Central, probably tetrabasal; two gonopores, one hydropore. Ambulacra: Narrow, on the aboral side nonpetaloid with minute pores at the horizontal sutures, adoral plates with two transversely oval depressions with the pore positioned in the outer one; aboral plates each with a single small, perforate and crenulated primary tubercle (Fig. 3.22). Interambulacra: Several irregularly arranged tubercles. Oral side: Concave, peristome anterior, transversely oval or round; phyllodes and bourrelets very faint. Periproct: On the aboral side at the upper end of a shallow anal groove. Tuberculation: Regular but not dense. Spines: On the upper side curved and smooth. Occurrence: Recorded from the Sulu and Timor Seas at  depths of 350 to 390 m, and from an area north of  Norfolk  Island, Tasman Sea, dredged at a depth of 710 m. Genus Nannolampas Mortensen, 1948 The genus is monospecific. Nannolampas tenera de Meijere, 1903 Test: Small and thin-plated, ovoid in outline with cutoff posterior end; upper side gently arched from the anterior round margin to the truncated posterior side; maximum length 13 mm. Apical system: Tetrabasal, two gonopores and one or more hydropores perforating genital plate G2; gonopores in females larger than in males.

3.2.1 Order: Cassiduloida Claus, 1880 

A

B

 31

C

D E

F

Fig. 3.22: (A, B) Aphanopora echinobrissoides. Male, test length 8 mm; Wanganella Bank north of Norfolk Island. Photos from D.G. McKnight (1968), placed kindly at my disposal by Owen Anderson, NIWA, Wellington. (A) Aboral side: there are two gonopores (in G1 and G4) and one hydropore (in G2) in the apical system. (B) Oral side: the relatively large peristome is circular. (C–F) Nannolampas tenera. Male specimen, test length 10 mm; Timor Sea, Indonesia. ZMUC. (C) Aboral side. In the apical system, two gonopores are developed with a single hydropore. (D) Oral side: the posterior end of the test is truncated. (E) Side view. (F) Posterior side: the periproct opens into a short, deep anal sulcus.

Ambulacra: Nonpetaloid; pores minute on the aboral side, larger around the peristome. Oral side: Depressed; peristome anterior, transversely oval; phyllodes and bourrelets absent. Periproct: At the truncated posterior side in a short, deep groove.

Primary tubercles: Perforate and crenulated. Occurrence: Only known from the Timor Sea at a depth of 390 m. Genus Neolampas A. Agassiz, 1869 The genus is monospecific.

32 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

Neolampas rostellata A. Agassiz, 1869 Test: Small, thin, and fragile; ovoid in outline; aboral side lowly arched, posterior side conspicuously truncated; length may reach 20 mm. Apical system: Slightly anterior; monobasal with three gonopores (the fourth pore in the specimen from the ZMU Copenhagen Fig. 892 is anomalous!), much larger in females than in males. Ambulacra: Nonpetaloid, aborally very small or quite rudimentary pores, orally larger (Fig. 3.23). Oral side: Concave; peristome slightly anterior, elongated, or round; incipient phyllodes, bourrelets more or less developed. Periproct: At the posterior face, rather deeply sunken at the upper end of a long anal groove. Tubercles: Indistinctly perforate and crenulated.

A

Distribution: This species, the most common in the family Neolampadidae, is recorded from the West Indies, particularly along the coast of Florida, eastern Atlantic from Morocco to the Channel Islands, and in the Mediterranean; bathymetric range 145 to 1,260 m. Genus Studeria Duncan, 1891 The genus is monospecific. Studeria recens A. Agassiz, 1879 Test: Ovoid in outline, high; maximum length 44 mm. Apical system: More or less anterior with three gonopores. Petals: Short, slightly bowed. Oral side: Well-developed phyllodes and bourrelets; no distinct naked areas in the posterior interambulacrum; peristome central and pentagonal.

B

C

Fig. 3.23: (A, B) Neolampas rostellata. Test length 9 mm, Talisman Station 15. ZMUC. (A) Aboral side. Note that the fourth gonopore of this female is anomalous; usually, there are only three! (B) Oral side: around the peristome, the ambulacral pores are enlarged, set with conspicuous tube feet. (C) Studeria recens. Test length 34 mm, New South Wales. Australia. (C) Aboral side. (D) Oral side. (E) Side view. Drawn after photos from Baker & Rowe (1990).

3.2.1 Order: Cassiduloida Claus, 1880 



Periproct: At the posterior side below a small hook. Distribution: From the Arafura Sea, Indonesia and northern New South Wales, Australia, over the Tasman Sea to the Kermadec Islands at a depth 135 to 236 m (Baker & Rowe 1990) Genus Tropholampas H.L. Clark, 1923 The genus is monospecific. Tropholampas loveni Studer, 1880 Test: Very small and fragile, aboral side highly vaulted or subconical, oral side flattened, posterior end obliquely truncated; in outline elongated, anterior part rounded, rather abruptly tapering to a truncated posterior end; length may reach only 8 mm. Apical system: Monobasal with four genital pores concealed in a deep, elongated marsupium in females and slightly sunken in males; one hydropore.

A

B

 33

Ambulacra: Nonpetaloid; simple pores on oral side, but not continuing above the ambitus, thus wholly lacking on the aboral side (Fig. 3.24). Oral side: Peristome subcentral, subpentagonal, or round; phyllodes or bourrelets absent. Periproct: At the truncated posterior side. Primary tubercles: Indistinctly perforated and crenulated, forming a dense granulation. Spines: Short and uniform, aborally densely covering the marsupium. Color: In living specimens, test and spines are whitish. Distribution: This species is reported only from South African waters, from off Cape Peninsula to Cape St. Francis in the east, at depths of approximately 135 to 350 m. Biology: Mortensen found up to three embryos in all stages of development in the marsupia of the females. One was almost ready to leave.

C

F D

E

Fig. 3.24: Tropholampas loveni. (A–C) Female, test length 6 mm; off Cape Peninsula, South Africa. ZMUC. (A) Aboral side: there is a deep, elongated marsupium, where the young grow up. (B) Oral side: the periproct is subpentagonal. (C) Side view: the test is highly inflated, the periproct on the lower posterior side. (D) Female with spines. Test length 6.1 mm, off Cape Peninsula, South Africa. ZMUC. The elongated marsupium is overarched by spines. (E, F) Male specimen, test length 6.1 mm; Cape Peninsula, South Africa. ZMUC. (E) Aboral side of test without spines: the apical system with the four gonopores is only slightly depressed. The hydropore is visible in the center. (F) Oral side of test with spines, length 5.9 mm. The subcircular periproct is overarched by specialized spines, which are longer than on the rest of the test.

34 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

3.2.2 Order Clypeasteroida L. Agassiz, 1835 In the Paleocene, approximately 60 million years ago, the clypeasteroids evolved from some forms of Cassiduloida through the extinct Oligopygoida. They are the youngest and most specialized group of irregular echinoids, being highly adapted for living borrowed in the uppermost, detritus-rich layer of sandy sediments, predominantly in shallow and more or less agitated waters. Their teeth function as a milling structure crushing sediment grains. Within 15 to 20 million years, the extremely flat discs expanded throughout all the oceans and they are found as fossils from all continents (Fig. 3.25). Clypeasteroids (common name, sand dollars) are unique in having multiple ambulacral pores associated with tube feet on each ambulacral plate. In all other echinoids, there is only a pore pair or a single pore per ambulacral plate (Fig. 3.26). Test: More or less flattened, strengthened by internal pillars and buttresses. Apical system: Monobasal plate (= genital plate 2) with four or five gonopores and five tiny ocular plates. Ambulacra: Wider or at least as wide as interambulacra at the ambitus.

Fig. 3.25: Main clypeasteroid characteristics: flat test, five leaf-shaped petals with wide pore zones, ambulacra wider than interambulacra at the ambitus; monobasal apical system with small ocular plates (Clypeaster prostratus, length 84 mm).

Podia: Nonrespiratory (or accessory) podia distributed in fields, patches or lines; respiratory podia arranged in petals. Oral side: Food grooves or ambulacral furrows (Fig. 3.27). The order Clypeasteroida includes two suborders: Clypeasterina, which has two families, and Scutellina, which can again be differentiated into two infraorders: Laganiformes with three families and Scutelliformes with six families. Literature: Mortensen 1948a,b; Durham 1955; Kier 1970; A.B. Smith 1984; Hopkins 1988; Mooi 1989, 1990.

Suborder Clypeasterina L. Agassiz, 1835 The members of this group have double lantern supports on the internal side of each pair of ambulacral plates next to the peristome. The respiratory pore pairs lead along sutures of demiplates. Two families are placed in this suborder: Arachnoididae with three genera and Clypeasteridae with a single genus (Fig. 3.28).

3.2.2.1 Family Arachnoididae Duncan, 1889 Kroh & Smith (2010) integrated the arachnoidids into the family Clypeasteridae, which is divided into two subfamilies: Ammotrophinae with the genus Ammotrophus, and Arachnoidinae with the genera Arachnoides and Fellaster. The three genera live buried just beneath the surface of sandy sediments in the intertidal zone, often in very large numbers. They are characterized by having very flat tests with a thin margin, and by the development of socalled “combed areas”, a form of food groove, which is unique in clypeasteroids (Fig. 3.29). Biology: In the “combed areas” on the oral and aboral sides, small tube feet are arranged in parallel series alternating with rows of small tubercles set with spines. These rows lead in an angle of approximately 45° to the broad food grooves along the ambulacral median suture, which extend almost from the apical system around the margin toward the peristome. On the oral side, the parallel rows are directed to the central mouth, on the aboral side, respectively, to the margin. Along these series, the tube feet transport organic particles, which they have picked selectively out of the

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

A

C

 35

B

D

E

F G

Fig. 3.26: (A, B) Internal supports: (A) Clypeaster rosaceus. Cross-section: scale bar is 20 mm, Michelin in Mortensen (1948a,b). The internal surface is more or less covered by calcareous skeleton supports forming pillars from the upper to the lower side, thin lamellae or needle-like structures such as stalagmites and stalactites. The arrangement of these processes may allow the grouping of this large, heterogeneous genus. (B) Horizontal section of the test of Clypeaster eurychorius; length 120 mm, off Natal, South Africa. ZMUC. In very flat clypeasterids (e.g., C. eurychorius or C. subdepressus), the calcareous supports are strongly developed along the inner side of the edge. Additional pillars occur more centrally, leaving space for the loop of the intestine. The median lines of the ambulacra with the radial canals are marked by narrow radial interspaces. (C–G) Monobasal apical plates of clypeasteroids. (C) Clypeaster australasiae. Scale bar is 2 mm. The entire plate is perforated by the hydropores. (D) Echinocyamus pusillus. Scale bar is 0.5 mm. There is only a single hydropore piercing the plate, four gonopores occur. (E) Laganum laganum. Scale bar is 1 mm. The hydropores lie in a sinuous furrow and five gonopores are developed. (C–E) Redrawn after Mooi (1989). (F) Echinodiscus auritus. With four gonopores, scale bar is 5 mm. (G) Encope michelini. With five gonopores. The star-shaped plates are densely perforated by hydropores.

36 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

A

C

B

Fig. 3.27: (A–C) Food grooves on the oral side of clypeasteroids: organic particles are transported by the podia through the furrows toward the mouth incorporated in a mucous string. (A) Simple food grooves in Clypeaster. (B) Bifurcating grooves in Astriclypeus. (C) Finely branched furrows in Dendraster. All scale bars are 10 mm. (D) Food grooves in clypeasteroids. The food grooves evolved independently in diverse families, being of more or less different shapes. Modified after Smith (1984).

sediment above and below the test. The particles are passed from one podium to the next, as in a bucketchain, toward the food grooves. Here, they are laid down and moved by fine cilia, and eventually manipulated around the margin, toward the mouth. The particles are not incorporated into a mucus string (as in the Mellitidae and related families), but dense, arched spines bend over the grooves and prevent the pieces from falling out. The tubercles and spines of the upper side are smaller than the food-gathering tubercles and spines of the oral side; and the aboral rows are more densely arranged than

the oral ones in the combed areas. Alternating with the combed areas of the underside there are zones with larger tubercles and longer spines used for locomotion. Literature: Mooi 1986, 1989; Rowe & Gates 1995; Miskelly 2002. Genus Ammotrophus H.L. Clark, 1928

Three species are described, endemic to Australia

(Fig. 3.30):

Ammotrophus arachnoides H.L. Clark, 1938 Test: Flat with thin margin; upper side slightly convex; circumference subcircular with shallow notches at

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

A

C

B

D

A

C

E

 37

Fig. 3.28: (A, B) Internal lantern supports in clypeasteroids, schematic: (A) Clypeasterina: the double lantern supports are positioned on the internal side of the ambulacral basicoronal plates (Fellaster zelandiae). (B) Scutellina: there are single structures on the internal side of the interambulacral basicoronal plates (Echinarachnius parma). Interambulacral plates shaded. Modified after Mooi (1989). (C–E) Pore pairs on half an ambulacrum in clypeasteroids. (C) Clypeasterina: the pore pairs lead along the sutures of demiplates. (D) Laganiformes: the pores cross the plate sutures. (E) Scutelliformes: the pores lead along the plate sutures. Redrawn after Mooi (1989).

B

D

E

Fig. 3.29: (A, B) Close-up of arachnoid echinoids. (A) Fellaster zelandiae, aboral side. (B) Arachnoides placenta, oral side. Parallel rows of tubercles carrying spines and pore pairs set with tube feet are arranged like the teeth of a comb. These fields are called “combed areas”. On the oral side, the combed areas alternate with plain fields, set with larger tubercles and spines for locomotion. Arrows indicate direction of food transport. Scale bars are 10 mm. (C–E) Plate patterns of the oral side: the interambulacral columns (shaded) are discontinuous between the single small plate in the ring next to the peristome and the margin. The isolated distal plates are usually visible in denuded tests. Redrawn after Mooi (1989). (C) Ammotrophus cyclius: the interambulacra are interrupted by two enlarged ambulacral plates, continuing with four to six plates near the margin. The periproct lies on the oral side. Scale bar is 50 mm. (D) Arachnoides placenta: the interambulacra are interrupted by four to five ambulacral plates, continuing with only two distal plates. The arrangement in Arachnoides tenuis is principally the same, but the first ring of ambulacral plates is larger than the following ones toward the margin. Scale bar is 30 mm. (E) Fellaster zelandiae: the interambulacra are interrupted by two enlarged ambulacral plates, continuing with four to six distal plates. The periproct lies on the aboral side. Scale bar is 50 mm.

38 

A

D

F

 3.2 Superorder Neognathostomata A.B. Smith, 1981

B

C

E

G



ambulacral midpoints, distinctly deepest in the posterior pair; maximum length may reach 150 mm. Apical system: Monobasal with four gonopores. Ambulacra: Petals broad, hardly reaching half of the test radius; interporiferous zones distinctly sunken; on the oral side, a pair of ambulacral plates interrupting the interambulacra is enlarged. Food grooves: Combed area narrow especially on the aboral side, the furrows themselves are broad and unbranched, extending over the entire aboral and oral side. Oral side: Flat or slightly concave; peristome subcentral; there are four to six distal plates in the interambulacra. Periproct: On the oral side, about one-third of the distance from the margin to the peristome. Spines: Very short like velvet on the aboral side, longer and differentiated on the oral side. Color: In life, the test is brownish, the bare test is grayish-white. Distribution: West coast of Australia from Jurien Bay to Cockburn Sound; depth range 3 to 20 m. Ammotrophus cyclius H.L. Clark, 1928 Very similar to A. arachnoides. Differences: With a maximum length of 90 mm much smaller, periproct closer to margin, test thicker, petals larger, mostly characteristics that are best recognizable in direct comparison. Distribution: The species is known from Southern Australia. The bathymetrical range is from 20 to 45 m. Ammotrophus platyterus H.L. Clark, 1928 Test: Flat, distinctly wider than long, no indentations at the margin, holotype 27 mm in length, 29.5 in width. Apical system: Very small with four gonopores. Ambulacra: Petals broad, A II and A IV slightly narrower than the three others. Food grooves: Four furrows are well-marked but indistinct in A III, whereas they are conspicuously curved in A II and A IV. Oral side: Peristome slightly posterior, periproct 3.5 mm distant from the margin, diamond-shaped. Spines: Not known, the holotype is naked. Occurrence: Gulf of St. Vincent, South Australia, based on a single specimen; depth unknown. Remarks: No other specimen has ever been found (personal communication: A. Miskelly 2015) (Fig. 3.31).

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

 39

Genus Arachnoides Leske, 1778 Two very similar species are known. Test: Very flat, lowly conical, sharp margin; circumference round with posterior notch; broad “combed areas” on oral and aboral sides; length up to 80 mm. Apical system: Four gonopores, which are slightly posterior. Ambulacra: Petals very broad, elevated above adjacent interambulacral zones. Food grooves: “Combed area” broad, furrows leading over oral and aboral sides. Oral side: Flat; peristome slightly posterior; two distal interambularal plates; second ambulacral plate from the peristome not enlarged. Periproct: Marginal to supramarginal. Spines: Very short like velvet, those on the oral side longer and differentiated. Color: In life, the test is more or less dark brown, the bare test is whitish. Literature: Rowe & Gates 1995; Miskelly 2002. Arachnoides placenta (Linné, 1758) The species is characterized by a naked furrow along the midline in the posterior interambulacrum on the oral side. The periproct is connected with the posterior margin by a short furrow. Test length up to 80 mm (Fig. 3.32). Distribution: This species is found in Northern Australia, New Guinea and Samoa to the Malayan Archipelago, the Philippines and south China; common in the intertidal zone, rarely down to 50 m. Arachnoides tenuis H.L. Clark, 1938 This species largely resembles Arachnoides placenta. However, it differs in the following features: the periproct is distinctly supramarginal without any notch and, on the oral side, there is no naked median line in the posterior interambulacrum as in A placenta. Distribution: This species is restricted to the littoral zone in Western Australia from Shark Bay to Broome. Genus Fellaster Durham, 1955 The genus is monospecific and characterized by the periproct positioned aborally.

◂ Fig. 3.30: (A–D) Ammotrophus arachnoides: (A) Aboral side of young specimen, test length 54.4 mm. The widely open petals are rather small. (B) Oral side. The periproct is positioned almost a third the distance from the margin to the peristome. (C) Aboral side of small specimen with spines; length 54.4 mm. (D) Oral side of large specimen with spines; length 148.3 mm. The “combed areas” are very distinct. All specimens from the Marmion Marine Park, Western Australia. (E–G) Ammotrophus cyclius. (E) Aboral side of test 83.3 mm; Long Beach, Robe, South Australia. (F) Oral side: the food grooves extend over the entire aboral and oral side. (G) Oral side with spines: length of test 47 mm; Adelaide, South Australia. The “combed areas” are marked by darker zigzag fields. Photos courtesy of A. Miskelly.

40 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

A

B

C

D

Fig. 3.31: (A, B) Ammotrophus platyterus. (A) Aboral side of test, length 27 mm; St. Vincent Gulf, South Australia. (B) Oral side. Photos courtesy of A. Miskelly. (C, D) Same specimen, after the photos from H.L. Clark (1928). Pattern of food grooves.

Fellaster zelandiae Gray, 1855 Test: Flat with thin margin; upper side on the whole more or less convex; circumference round with shallow notches at ambulacral midpoints; length up to 120 mm. Apical system: Monobasal with four gonopores. Ambulacra: Petals very broad and elevated above adjacent interambulacral zones. Food grooves: Broad, extending onto the aboral side. Oral side: Flat or slightly concave; peristome subcentral; four to six distal plates in the interambulacrum; periproct aboral, more than its own diameter away from the margin. Spines: On the aboral side, very short, longer, and differentiated on the oral side.

Color: In life, the test is dark to light brownish or grayishpurple, the bare test is grayish-white. Distribution: This species is reported from New Zealand, around the North Island, and down to Dunedin on the South Island. It is also found in Australia from Queensland, at Flinders Island in the Bass Strait, and Tasmania. It lives in the littoral zone (Fig. 3.33). Literature: McKnight 1969; Rowe & Gates 1995. 3.2.2.2 Family Clypeasteridae L. Agassiz, 1835 Genus Clypeaster Lamarck, 1801 Numerous clypeasterid species are distributed all over the tropical and subtropical world, having evolved diverse shapes with many intermediates. Because the variability

▸ Fig. 3.32: (A–E) Arachnoides placenta. (A) Aboral side of test, length 61 mm; Lavina, Bali, Indonesia. (B) Oral side of test. From the aboral side, the food grooves lead along the ambulacral midlines around the margin and across the oral side to the central peristome. The naked furrow in the posterior interambulacrum is distinct. (C, D) Enlargement of posterior interambulacrum. (C) Aboral side: the periproct opens into a distinct marginal notch (arrow). (D) Oral side: a naked furrow passes along the median suture. Scales in (C, D) are 10 mm. (E) Aboral side with spines, length 39 mm; Krabi, Thailand. The test is very densely covered in short spines. (F, G) Arachnoides tenuis. (F) Aboral side of test, length 29 mm; Broome, Western Australia. (G) In contrast to A. placenta, there is no marginal notch at the periproct and no naked furrow in the posterior interambulacrum on the oral side. The second ambulacral plates from the peristome are enlarged. Photos courtesy of H. van Noordenburg.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

B

A

C

F

D

E

G

 41

42 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

A

B

C

D

Fig. 3.33: Fellaster zelandiae. (A, B) Test length 101 mm; Stanley Bay, North Island, New Zealand. (A) Aboral side: the broad interporiferous zones are sharply elevated above the interambulacra. (B) Oral side: there are four to six distal plates in the interambulacra. Zones with larger tubercles alternate with the “combed areas”. (C, D) Test length 81 mm; Auckland; New Zealand. (C) Aboral side with spines. Along the midlines of the ambulacra, broad food grooves lead to the margin. The spines are club-shaped. (D) Oral side: the “combed areas” with short spines on both sides of the food grooves alternate with zones set with longer spines for locomotion.

of the species is large, they are sometimes difficult to distinguish, especially if the locality is not known. The clypeasterids are found from the littoral zone to depths of more than 300 m. They live mostly infaunally. The genus is characterized as follows: Test: Large to very large, flat to inflated with rounded or rather sharp margins, the circumference is ovate, subcircular, or subpentagonal.

Apical system: Monobasal plate with five gonopores and five small ocular plates. Ambulacra: Five petals of approximately equal shape are well-developed, distally closed, or open. Oral side: Flat or more or less concave or sunken only toward the peristome, with five simple food grooves. Periproct: Positioned on the oral side more or less near the posterior margin.



Spines: Test covered by very short miliary spines and longer primary spines more or less densely distributed over the surface; those on the oral side usually longer and thicker than aborally. Remarks: Mortensen and other authors note the number of primary tubercles on the ridges between the pore pairs as valuable specific differences, but this number depends largely on the ontogenetic stage of the individual. Young specimens of a species naturally have fewer tubercles than adults. Nevertheless, the arrangement or density of the tubercles may be useful for distinction (see the drawings of Mortensen 1948a,b). Literature: de Meijere 1902, 1903; Clark, H.L. 1914, 1925, 1938; Koehler 1922; Ikeda 1935; Durham 1954; Cherbonnier 1959; Endean & Pope 1964; Kier 1970; Serafy 1970, 1971; Phelan 1977; Pawson & Phelan 1979; Shigei 1986; Hopkins 1988; Mooi 1989; Rowe & Gates 1995; Hendler et al. 1995; Mihaliejevic et al. 2011. The genus consists of numerous species. They are listed here first according to the geographical range, with the following descriptions sorted alphabetically: West Atlantic and Caribbean: Clypeaster chesheri Serafy, 1970. Florida, Antilles, Yucatan to French Guiana. Clypeaster cyclopilus H.L. Clark, 1941. Bahama-Cuba region. Clypeaster durandi Cherbonnier, 1959, French Guiana. Clypeaster euclastus H.L. Clark, 1941. West Indies. Clypeaster lamprus H.L. Clark, 1914. West Indies. Clypeaster luetkeni Mortensen, 1948. West Indies. Clypeaster pallidus H.L. Clark, 1914. Barbados, Dominica, and Colombia. Clypeaster prostratus Ravenel, 1848. South Carolina, USA to Gulf of Mexico. Clypeaster ravenelii A. Agassiz, 1869. Southeast USA, West Indies. Clypeaster rosaceus Linné, 1758. South Carolina, USA to Barbados and Curacao. Clypeaster subdepressus Gray, 1825. Florida, USA to Brazil. East Atlantic: Clypeaster rangianus Desmoulins, 1835. West Africa, Cape Verde Islands to Angola. Japan: Clypeaster japonicus Döderlein, 1885. Southern Japan. Australia: Clypeaster australasiae Gray, 1851. East and Southeast Australia to Tasmania.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

 43

Clypeaster telurus H.L. Clark, 1914. West Australia from Fremantle northward to Fraser Island. Clypeaster tumidus Tenison-Woods, 1878. Northeast Australia. Indo-Pacific: Clypeaster amplificatus Koehler 1922. Red Sea. Clypeaster annandalei Koehler, 1922. Northern Indian Ocean to the Philippines. Clypeaster eurychorius H.L. Clark, 1924. East coast of Africa, Mauritius. Clypeaster eurypetalus H.L. Clark, 1925. Pearl and Hermes Reef, Central Pacific. Clypeaster fervens Koehler, 1922. Red Sea to Moluccas. Clypeaster humilis Leske, 1778. Red Sea to Philippines and New Caledonia. Clypeaster kieri Pawson & Phelan, 1979. Clypeaster latissimus Lamarck, 1816. Indo-China to Sunda Strait. Clypeaster leptostracon A. Agassiz & H.L. Clark, 1897. Hawaii. Clypeaster lytopetalus A. Agassiz & H.L. Clark, 1907. Hawaii. Clypeaster miniaceus H.L. Clark, 1925. Indo-West Pacific, Macclesfield Bank to Mauritius. Clypeaster nummus Mortensen, 1948, South Pacific. Clypeaster oshimensis Ikeda, 1935. Clypeaster pateriformis Mortensen, 1948. Malayan Archipelago and Philippines. Clypeaster rarispinus de Meijere, 1902, 1903. Red Sea, Natal to Java Sea, not Philippines. Clypeaster reticulatus Linné, 1758. Red Sea, Natal to Hawaii, New Caledonia. Clypeaster virescens Döderlein, 1885. Southern Japan to Philippines. West coast of America: Clypeaster elongatus H.L. Clark, 1948. Galapagos Islands. Clypeaster europacificus H.L. Clark, 1905. Gulf of California to Ecuador and Galapagos Islands. Clypeaster isolatus Serafy, 1971. San Felix Island, East Pacific. Clypeaster ochrus H.L. Clark, 1914. Baja California to Ecuador and Galapagos Islands. Clypeaster rotundus A. Agassiz, 1863. Gulf of California to Ecuador and Galapagos Islands. Clypeaster speciosus Verrill, 1870. Baja California to Clarion Island. Clypeaster amplificatus Koehler 1922 Koehler described a single very large specimen with a length of 190 mm, height 23 mm.

44 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

Test: Flat, margin thin, only petalous area slightly inflated. It differs from Clypeaster humilis by the broad petals, which leave almost no space for the adapical interambulacral plates. Occurrence: The specimen is labeled “Massaouah”, Red Sea. Clypeaster annandalei Koehler, 1922 Test: In outline, ovoid to rounded pentagonal; length may reach 124 mm; height of petaloid area very variable, may be subconical to hat-shaped; margin rather thin. Petals: Paired petals almost closed, anterior petal open; interporiferous zones narrow; ridges between the pore pairs with three to six primary tubercles and some miliaries; area approximately 70% of test length. Oral side: Flat, more or less deeply sunken toward the peristome; food grooves almost reaching the margin; periproct about its diameter distant from the posterior margin. Color: Spines and test yellow-brown to orange-brown, oral side usually paler. The plate sutures and the food grooves may be lighter. Distribution: The species is known from the northern Indian Ocean over the Malayan Archipelago to the Philippines; bathymetric range 80 to 550 m (Fig. 3.34). Clypeaster australasiae Gray, 1851 Test: Large and thick, circumference oval to subpentagonal; upper side more or less lowly convex; length may reach 135 mm. Apical system: Sometimes raised. Petals: Broad, reaching approximately more than 60% of the aboral side, interporiferous zones slightly inflated, pore zones sunken and distally rather open; anterior petal the least closed. Oral side: Concave, depressed toward the peristome; food grooves recessed; periproct just below the posterior margin. Spination: Short and very dense. Color: In life, the spines are brown or fawn; the bare, well cleaned test is white. Distribution: The species is endemic to Australia, from Bowen, Queensland, along the east coast to Victoria in the south. It lives from the tidal zone to a depth of 75 m (Fig. 3.35). Clypeaster chesheri Serafy, 1970 Test: Very flat with thin margin, conically raised toward the slightly elevated apex; in outline more or less oval, with slight indentations in the posterior lateral interambulacra; width greater than 88% of test length; total length may reach 184 mm.

Petals: Less than 50% of test length, all petals closed distally, poriferous zones narrow; ridge between pore pairs set with nine or more primary tubercles in adults. Oral side: Nearly flat, slightly concave around the peristome. Color: In living specimens, the test and spines are yellowish-brown, lighter along the sutures. Well cleaned tests are white. Distribution: The species is reported from Florida, Gulf of Mexico, southward throughout the entire Antilles and along the coasts of Yucatan, Mexico, to French Guiana. It lives in depths of 20 to more than 100 m. Clypeaster cyclopilus H.L. Clark, 1941 Test: Nearly circular in outline; margin flat; petaloid area inflated, sharp rise to apex; length up to 65 mm. Petals: Short and widely open, slightly bowed, pore zones narrow; ridge between pore pairs with one to two primary tubercles, sometimes none, and a row of miliaries. Periproct: Relatively large, at a distance less than its own diameter from the posterior margin. Distribution: Reported from the Bahama-Cuba region, bathymetric range 170 to 255 m. Remarks: Both C. cyclopilus and C. euclastus have broad, widely open petals and scattered tubercles among a dense coat of miliaries. However, the former seems to stay, with a maximum length of 65 mm, rather small whereas the latter grows to a much larger size (Fig. 3.36). Clypeaster durandi Cherbonnier, 1959 Test: Large, center more or less inflated; length up to 134 mm. Petals: Broad, curved distally and closed at the end; three to four primary tubercles in a series on the ridges between the pore pairs. Distribution: Found at the coast of French Guiana, depth not reported. Clypeaster elongatus H.L. Clark, 1948 Test: Distinctly elongated, width less than 82% of length; length of holotype 111 mm, width 89 mm, elevating from the margin to a height of 27 mm at the apex, in outline shallowly indented laterally. Petals: Approximately 70% of test length, anterior longest, lateral ones shortest; all virtually closed distally. Oral side: Sunken toward peristome, periproct less than its own diameter distant from the margin. Occurrence: Galapagos Islands, shallow water. Clypeaster euclastus H.L. Clark, 1941 Test: Rounded pentagonal with more or less concave sides, may be slightly wider than long; petaloid area inflated, adapically flattened; test distally quite flat with thin margin; maximum length 184 mm.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

A

B

D

 45

C

E

F

Fig. 3.34: (A) Clypeaster amplificatus. Aboral side of test, length 190 mm, Red Sea (photo from Koehler 1922). (B–F) Clypeaster annandalei. (B) Aboral side with spines; length 61 mm; Cebu, Philippines. (C) Oral side. (D) Aboral side of test, length 54 mm; Madras, India. ZMUC. (E) Oral side. The food grooves extend around the margin to the aboral side. (F) Test length 56 mm, Madras, India, ZMUC. Side view: the height is very variable from conically raised to strongly elevated like a hat.

Petals: Petaloid area about half the test length, petals broad and widely open, not at all turning inwards; few primary tubercles on ridges between pore pairs, not much larger than miliary tubercles. Oral side: Quite flat, peristome only slightly sunken; periproct very close to the posterior margin. Color: Aborally dull pink with darker sutures, the oral side is more yellowish.

Distribution: The species is known from the Bahamas and off the Orinoco River. It most probably occurs in the entire West Indies; known bathymetrical range 86 to 325 m. Clypeaster europacificus H.L. Clark, 1905 Test: Thin; in profile very low, slightly raised toward the apical system; circumference pentagonal with concave sides; maximum length more than 200 mm.

46 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

B

A

C

D

E

F Fig. 3.35: (A–C) Clypeaster australasiae. (A) Aboral side of test; length 125 mm, height 31 mm; New South Wales, Australia. The petals are broad and distally open. (B) Oral side: the food grooves are deeply recessed, the peristome is sunken. (C) Profile of test: the stippled line indicates the oral concavity toward the peristome. Scale bar is 50 mm. (D–F) Clypeaster chesheri. (D) Aboral side with spines; length 157 mm; Key West, Florida, USA. The petals are short. (E) Oral side. (F) Side view: the test is very low and conically inflated toward the apex. (D–F, photos by H. van Noordenburg).

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

 47

B

A

C D

E

F

G

Fig. 3.36: (A–C) Clypeaster cyclopilus. (A) Aboral side of test, length 64 mm, Cuba. ZMUC. (B) Side view: the petaloid area is strongly inflated. (C) Part of petal, scale bar is 3 mm, after Mortensen (1948a,b). (D) Clypeaster durandi. Part of petals, scale bar is 2 mm, after Cherbonnier (1959). (E–G) Clypeaster euclastus. (E) Part of petal. Scale bar is 3 mm, after Mortensen (1948a,b). (F) Sketch of aboral side (plate sutures mostly omitted); length 184 mm. (G) Profile. Scale bar is 40 mm, after a photo in Mortensen (1948a,b).

Petals: Inflated, open distally. Oral side: Flat, only slightly depressed toward the peristome, food grooves indented, reaching to the margin, periproct marginal in a distinct notch at the posterior side. Spination: Short and very dense. Color: More or less brownish. Distribution: This species is known along the west coast of America from the Gulf of California to Ecuador and the Galapagos Islands at depths of 12 to 175 m. Remarks: C. europacificus differs from C. speciosus by the deep notch at its posterior margin, into which the periproct opens, and by its thin margin (Fig. 3.37).

Clypeaster eurychorius H.L. Clark, 1924 Test: Moderately solid, outline rounded pentagonal; on aboral side, distal part flat with rounded edges, sometimes slightly thickened; petaloid area rather abruptly and more or less highly raised; length of test up to 145 mm. Petals: Short and flush with the test, open; pore zones remarkably narrow, interporiferous zones about three times the width of pore zones. Oral side: Distally flat, sunken only near the peristome; food grooves reaching the margin or even the aboral side; periproct close to the posterior edge. Primary tubercles: Rather sparse, miliary tubercles much smaller than primaries.

48 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

A

B

C D

F

E

G

H



Distribution: The species is known from the Natal coast, South Africa, northward to the Pemba Strait, and off Mauritius, at 125 to 320 m. Clypeaster eurypetalus H.L. Clark, 1925 Clark described only a single specimen with “amazingly wide petals”. Length of the test 112 mm. Occurrence: Pearl and Hermes Reef, Central Pacific, found at the beach (Fig. 3.38). Clypeaster fervens Koehler, 1922 Test: Oval, often more or less with concave sides; upper side distinctly inflated rising straight to the flattened apex; test length may reach 90 mm. Petals: Approximately two-thirds of test length; broad, paired ones strongly curved and closed, anterior petal open, interporiferous zone inflated; two to five primary tubercles on ridges between pore pairs, connecting shallow furrows. Oral side: Deeply depressed toward peristome; periproct only half the size of peristome; food grooves reaching the margin. Color: In living specimens, the test and spines are reddish-brown. Distribution: Red Sea and East Africa to the Malayan Archipelago, at a depth of 13 to 110 m. Clypeaster humilis (Leske, 1778) Test: Distinctly longer than wide, pentagonal, sides more or less concave, margin more or less thick, petaloid area raised; length up to 160 mm. Petals: More or less broad, distally closed; area not more than half the width of test. Oral side: May be concave; food grooves long and deeply indented. Spines: Very short, primaries barely longer than miliary spines. Color: In life, the specimens are greenish-gray to brownish, the petals usually darker; the denuded test is beige to yellowish. Distribution: This species is reported from the Red Sea and the Gulf of Iran to the Malayan Archipelago, northeast Australia, and New Caledonia, it is not known from East Africa, living at 0 to 216 m.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

 49

Remarks: C. humilis differs from C. reticulatus by its deeply recessed food grooves and by the lack of a ring-shaped depression around the petaloid area. It differs from C. fervens by its smaller petals and its sharper margin (Fig. 3.39). Clypeaster isolatus Serafy, 1971 Test: Oval to rounded pentagonal, moderately high, rising uniformly from the margin to the apex; length up to 56.6 mm. Petals: Very broad with more or less curved ends, distally open; three to six pore pairs in the ridge between the pore pairs. Oral side: Concave, gradually sloping toward the peristome; periproct about its own diameter distant from the margin. Occurrence: Only dead tests were found off San Felix Island, west of Chile, at a depth of 75 m. Clypeaster japonicus Döderlein, 1885 Test: Large, strong, and heavy with thick margin; outline elongated pentagonal; upper side gradually rising from the margin to the apex, more or less high; maximum length more than 120 mm. Petals: Petaloid area two-thirds to three-quarters of test length; paired petals closed distally, anterior one open to varying degrees. Oral side: Concave toward the peristome, sometimes rather flat and abruptly depressed; food grooves conspicuously indented; periproct near the posterior margin. Spines: Primaries longer than the densely distributed miliaries. Color: In life, the spines are uniformly brown with darker petals, the bare test is grayish to brownish. Distribution: C. japonicus is restricted to Japanese waters from northern Honshu to southern Kyushu at depths of 2 to 50 m. Clypeaster kieri Pawson & Phelan, 1979 Test: Rounded pentagonal; slightly longer than broad; flattened margin, central part gradually raised; length up to 55 mm, may reach 85 mm. Petals: Paired ones curved and closed, anterior one curved distally and more or less open; ridges between pore pairs with four to five scattered primary tubercles.

◂ Fig. 3.37: (A–C) Clypeaster europacificus. (A) Aboral side of test, length 121 mm, width 115 mm, height 16 mm; Costa Rica. ZMH. The petals are inflated. The outline shows concave sides. (B) Oral side: the underside is flat, only slightly depressed toward the peristome. The periproct lies marginal in a conspicuous notch. In the interior of the test, the wings of the lantern are visible. (C) Profile of test: the margin is very thin, the apical region evenly inflated. Scale bar is 50 mm. (D–H) Clypeaster eurychorius. (D) Aboral side of test, length 130 mm; Pemba Strait, East Africa. ZMUC. (E) Oral side. (F) Side view. (G) Part of petal: the primary tubercles are sparse. Scale bar is 4 mm, after Mortensen (1948a,b). (H) Aboral side of spined specimen, length 81 mm, coast of Transkei, South Africa.

50 

A

 3.2 Superorder Neognathostomata A.B. Smith, 1981

B

C

D

E

G

F

3.2.2 Order Clypeasteroida L. Agassiz, 1835 



A

D

F

B

 51

C

E

G

Oral side: Periproct laying its own diameter distant to the posterior margin; food grooves reaching to the margin. Occurrence: Southwest of Mumbai in the Arabian Sea, at depths of 84 to 106 m (Fig. 3.40). Clypeaster lamprus H.L. Clark, 1914 Test: Oval to ovoid, oral side flat, slightly sunken near the peristome; petaloid area raised more than half the height of the test; maximum length 108 mm. Petals: Relatively short, paired ones half open, anterior broadly open; ridge between pore pairs with very few primary tubercles; adapical primary tubercles conspicuously large; on the oral side especially along the median part of interambulacra.

Fig. 3.39: (A–D) Clypeaster humilis. (A) Aboral side, test length 91 mm; Safaga, Egypt, Red Sea. The petals are inflated. (B) Oral side: the food grooves are sharply indented. (C) Aboral side with spines, length 71 mm; Negros, Philippines. The primary spines are barely longer than the very densely distributed miliary spines. (D) Profile: the upper side is convex. Scale bar is 50 mm. (E–G) Clypeaster japonicus. (E) Profile of test (stippled line marks the oral cavity). The peristome is deeply sunken, and the short food grooves are distinctly recessed. (F) Aboral side of test with spines, length 102 mm; Nanao Bay, Ishikawa Prefecture, Japan. (G) Oral side without spines, test length 101 mm; Sagami Bay, Kanagawa Prefecture, Japan. The petaloid area takes approximately two-thirds to three-fourths of the test length.

Periproct: Its own diameter distant from margin. Color: In living specimens, yellowish-green, the dry test is light brown. Distribution: Known in the West Indies at depths of 160 to 220 m. Clypeaster latissimus (Lamarck, 1816) Test: Very flat with thin margin; outline ovoid, posterior side somewhat truncated; very slightly raised toward the center; sometimes, the petaloid area is very slightly domed; length up to 248 mm. Apical system: Small. Petals: Short and narrow, anterior one longest, all petals firmly closed; in large specimens, outer

◂ Fig. 3.38: (A) Clypeaster eurypetalus. Test length 112 mm, Pearl and Hermes Reef. After the photos in H.L. Clark (1925). (B–G) Clypeaster fervens. (B) Aboral side of test; length 77 mm, southeast coast of Arabia. ZMUC. (C) Oral side. The peristome is deeply sunken. (D) Side view. (E) Part of petal: the furrows connecting the pore pairs are shallow. Scale bar is 3 mm, after Mortensen (1948a,b). (F) Aboral side with spines, test length 70 mm, Mauritius. ZMUC. (G) Side view: the test is conically inflated.

52 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

A

B

C

D

E

Fig. 3.40: (A, B) Clypeaster kieri. (A) Aboral side of holotype, length of test 50 mm, Arabian Sea. (B) Profile. Sketches after photos in Pawson & Phelan (1979). (C–E) Clypeaster lamprus. (C) Sketch of aboral side, length of test 92 mm. (D) Profile. Scale bar is 20 mm. Sketches after photos in Mortensen (1948a,b). (E) Part of petal: there are only few primary tubercles between the pore pairs. Scale bar is 3 mm, after Mortensen (1948a,b).

pores round; ridge between pore pairs set with a dense series of primary tubercles, miliaries very small; petaloid area approximately half the test length. Oral side: Perfectly flat, slightly sunken near the small peristome, periproct about three to four times its diameter distant from the margin.

Primary tubercles: Small and densely distributed; primary spines on the aboral side club-shaped. Color: Spines and test are dark brown, well cleaned tests are white. Distribution: This species is recorded from Indo-China and in the entire Malayan region to northern Australia, from the sublittoral to a depth of 45 m (Fig. 3.41).

▸ Fig. 3.41: (A–D) Clypeaster latissimus. (A) Aboral side of test, length 121 mm; Sunda Strait. ZMUC. The apical system is very small. (B) Oral side. (C) Profile of test. (D) Part of petal: the outer pores are round in large specimens. Scale bar is 2 mm. After Mortensen (1948a,b). (E–H) Clypeaster luetkeni. (E) Aboral side of test, length 18 mm, Haiti. ZMUC Holotype ECH 200. (F) Oral side. (G) Profile. The margin is rounded. Sketch after the photo in Mortensen (1948a,b).

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

A

B

C D

E

F

G

 53

54 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

Clypeaster leptostracon A. Agassiz & H.L. Clark, 1897 Holotype 38 mm. In shape very much like C. miniaceus, but the color (in alcohol) is different: it varies from bright yellow or reddish yellow to dirty purplish white, sometimes with indistinct dusky blotches aborally. Distribution: Hawaii Islands; depth range between 200 and 600 m. Clypeaster luetkeni Mortensen, 1948 Test: Relatively small, maximum recorded length 47.3 mm, height 10.2 mm; ovoid and pentagonal, broadest at end of posterior petals; aboral side raised, sloping regularly toward the anterior, toward the posterior with abrupt dip behind the posterior petals. Apical system: Small. Petals: Slightly inflated; posterior pair closed, the three others half open; ridges between pore pairs set with three to five primary tubercles, connecting furrow rather broad; petaloid area a little more than half the test length. Oral side: Concave around the peristome, distally flat; margin rounded and thickened (approximately two-thirds of height), anteriorly a little thicker than posteriorly; peristome sunken, periproct its own diameter distant from posterior end. Primary tubercles: Not much larger than miliaries, all not numerous. Color: Light pink to red purple, plate sutures may be darker. Distribution: Antilles, sublittoral. Remarks: Mortensen described the new species on the base of a specimen of 18 mm, which already showed gonopores (for comparison, the gonopores of C. subdepressus appear at a length of between 44 and 65 mm according to Mortensen) C. luetkeni may reach a considerably larger size, but remain relatively small compared with other species of Clypeaster. Literature: Serafy 1970; Hopkins 1988. Clypeaster lytopetalus A. Agassiz & H.L. Clark, 1907 Mortensen (1948b) thinks this species is synonymous with C. reticulatus. Found at Laysan, Hawaiian Islands, at a depth of 30 to 235 m. Clypeaster miniaceus H.L. Clark, 1925 Test: Rather delicate, outline ovoid to rounded pentagonal, sides sometimes slightly concave; margin rounded; petaloid area slightly raised; distal part of test flat; length may reach 88 mm. Petals: Area less than half the test length; all petals may be more or less widely open, or the paired petals may be almost

or totally closed; anterior petal the longest; ridge between pore pairs with one to three primary tubercles; pore zones narrow, inner and outer pores of almost equal size. Oral side: More or less concave, food grooves shallow but distinct, extending to the margin; peristome slightly sunken; periproct about its own diameter distant from the posterior edge. Tuberculation: On the whole, sparse. Color: The center of the aboral plate is often of a striking red color, but it also may be faded out. On the oral side, the color is always less distinct, or the plates are even white. Distribution: This species is known from the South China Sea and Mauritius, it probably occurs more widely in the Indian Ocean and perhaps in the Philippines and Malayan region; bathymetric range 70 to 230 m (Fig. 3.42). Clypeaster nummus Mortensen, 1948 Mortensen described a single young specimen without gonopores, measuring 37 mm. Test: Broadly oval to subcircular, regularly lowly conical, oral side convex. Petaloid area A little more than half the test length. Occurrence: South Pacific, type found at Tahiti. Clypeaster ochrus H.L. Clark, 1914 Test: Rounded pentagonal, sides slightly concave; upper side raised with rounded margin, deeply sunken toward the peristome (may be also flattened); length up to 112 mm. Petals: All half open, widest at the posterior pair, pore zones strongly turning inwards; ridge between pore pairs usually with four to five primary tubercles, numerous miliary tubercles; petaloid area approximately two-thirds of test length. Oral side: Peristome sunken, periproct less than its own diameter distant to the margin. Color: Very variable, living specimens are dull brown to almost black, or reddish-white. The oral primary spines may have an indistinct dark band. The denuded test is whitish and more or less mottled with brown. Distribution: Recorded at the west coast of America from Baja California to Ecuador and Galapagos Islands, littoral to 90 m. Clypeaster oshimensis Ikeda, 1935 Ikeda (1935) described two dead specimens of 85 and 77 mm length from Kagoshima, southern Japan. They are characterized by unusually formed interambulacral plates at the peristome, resembling the bourrelets in the Cassiduloida.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 



 55

B

A

C

D

E

G

F Fig. 3.42: (A–C) Clypeaster miniaceus. (A) Aboral side of test, length 59 mm, Mauritius. (B) Oral side. (C) Part of petal: the relatively few primary tubercles occur within scattered miliary tubercles. Scale bar is 3 mm, after Mortensen (1948a,b). (D–G) Clypeaster ochrus. (D) Aboral side of test, length 90 mm; Taboga Island, Gulf of Panama. ZMUC. (E) Oral side. (F) Side view. (G) Aboral side with spines, length 100 mm, Taboga Island, Gulf of Panama. ZMUC.

56 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

Test: Outline rounded pentagonal, thick margin rising continuously to the apex. Petals: Slightly elevated, unpaired petal broadly open, paired ones closed; ridges between pore pairs set with one to three small primary tubercles. Oral side: Deeply indented food grooves alternating with “swollen” interambulacral plates at the edge of the peristome. Color: The dead tests are more or less yellow. Remarks: Neither Mortensen (1948a,b) nor Shigei (several articles between 1970 and 1986) mention this species. Perhaps these tests may have been done on abnormal specimen of C. japonicus, which has deeply recessed food grooves as well, albeit no strongly inflated interambulacral plates (Fig. 3.43A, C).

Clypeaster pallidus H.L. Clark, 1914 This species is similar to C. rosaceus. It differs by the less elevated interporiferous zones and the less numerous pore pairs in the petals. Mortensen (1948a,b) doubted the validity of the species. Distribution: Barbados, Dominica, and Colombia; depth range of the two described specimens approximately 100 m (Fig. 3.43B, D). Clypeaster pateriformis Mortensen, 1948 Test: Nearly circular, very low, oral side markedly concave, resembling a shallow bowl; holotype 70 mm long, only 9 mm high. Petals: Broad adapically, but pointed and closed distally; ridges between pore pairs carry four to seven primary

A

B

C

D

Fig. 3.43: (A, C) Clypeaster oshimensis. Sketch of oral side of test, length 55 mm, Kagoshima, Kyushu, Japan. (C) Profile. Sketches after photos in Ikeda (1935). (B, D) Clypeaster pallidus. (B) Aboral side of holotype, length of test 108 mm, Barbados, West Indies. (D) Side view. Photos in H.L. Clark (1914).

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

tubercles; petaloid area only a little more than one-third of length. Peristome: Small, transversely oval; periproct relatively distant from the margin. Color: Uniformly brown. Distribution: Malayan region and Philippines, 30 to 90 m. Clypeaster prostratus Ravenel, 1845 Test: Elongated pentagonal, sides concave; lowly domed, maximum length 115 mm. Petals: Paired ones almost closed, anterior one open, interporiferous zone flush; ridges between the pore pairs

A

 57

set with a dense series of primary tubercles; petaloid area approximately two-thirds of test length. Oral side: Totally flat, only immediately around the peristome steeply sunken; furrows distinct to edge; periproct its own diameter distant from margin. Primary spines: On aboral side club-shaped (as in C. subdepressus). Distribution: From South Carolina, USA to the West Indies; depth 55 to 75 mm (Fig. 3.44). Clypeaster rangianus Desmoulins, 1835 Test: Heavy and strong; upper side rising more or less straight from the rounded margin to the rounded apex;

B

D

C

Fig. 3.44: Clypeaster prostratus. (A) Aboral side of test, length 84 mm, width 75 mm; Gulf of Mexico. ZMUC. (B) Oral side. (C) Side view. (D) Part of petal: the ridges between the pore pairs are densely set with a row of primary tubercles. Scale bar is 3 mm, after Mortensen (1948a,b).

58 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

outline oval to elongated pentagonal, the sides may be concave or convex; length more than 115 mm. Petals: Broad, ovoid, more or less widely open distally; pore zones narrow; interporiferous zones inflated; ridges between pore pairs carry four to six primary tubercles. Oral side: Flat distally but sunken toward the pentagonal peristome. Periproct: Its own diameter distant from the margin, food grooves reaching the edge. Color: Reddish to yellowish-brown. Distribution: The species has been recorded in West Africa from the Cape Verde Islands to Angola, sublittoral (Fig. 3.45). Clypeaster rarispinus de Meijere, 1903 Test: Solid; conically raised; in outline rounded pentagonal, usually slightly longer than wide, with slightly concave sides, posterior side convex, often with small prominence in the middle of the posterior side, margin rounded or distinctly thickened, but it may also be thin and flat; length of test up to 108 mm. Petals: Relatively narrow, lanceolate; paired petals firmly closed distally, ridge between pore pairs with only one primary tubercle close to the inner side of the pore zone; petaloid area half the test length or a little more. Oral side: Flat, sunken toward the peristome; food grooves extend to the edge; periproct less than its own diameter close to the posterior margin. Primary tubercles: Sparse. Color: On the oral side, a darker zigzag band often occurs along the food grooves. Distribution: The species is known in the Indian Ocean, in East Africa from the Red Sea to Natal, from the Persian Gulf over India to the Malayan Archipelago. Bathymetric range, from the littoral to a depth of approximately 100 m (Fig. 3.46). Clypeaster ravenelii A. Agassiz, 1869 Test: Distinctly pentagonal in outline, all sides concave, margin conspicuously thickened; center more or less domed; length up to 133 mm. Petals: All five widely open, pore zones rather straight; interporiferous zone flush; ridge between pore pairs set with several primary tubercles among densely distributed miliary tubercles; petaloid area about half the test length. Oral side: Flat, sunken around the peristome; food grooves deep becoming faint toward the margin.

Periproct: Close to the posterior edge. Distribution: This species is reported from Cape Hatteras, USA, southward through the Antilles to Grenada, and in the Gulf of Mexico from Florida to Texas and off Yucatan. Bathymetric range 50 to 230 m. Remarks: This species is unique among the Caribbean species in having a distinctly inflated margin (Fig. 3.47). Clypeaster reticulatus (Linné, 1758) Test: Of moderate size, solid, slightly raised toward the apex; margin more or less thick, inside the margin a distinct ring-shaped depression; circumference elongated to pentagonal; maximum length 75 mm; shape sometimes very irregular with strongly arched tests, being concave in transverse or long axis on the oral side. Petals: Long and inflated, anterior one the longest, anterior pair shortest; petaloid area more than two-thirds width of test, petals inflated. Oral side: Mostly concave like a round bowl; food grooves shallow and indistinct; periproct near the posterior margin. Spines: Short and coarse. Color: In life, variable, mostly gray-yellowish, often with darker spots, the bare test is whitish. Distribution: Widespread across the entire Indo-West Pacific from the Red Sea to Hawaii and the South Sea, from Taiwan to New Caledonia (not in Australia), living from the littoral zone to a depth of 125 m. Clypeaster rosaceus (Linné, 1758) Test: Generally inflated with thick margin, anteriorly thicker than posteriorly, circumference ovate to slightly pentagonal; maximum length 200 mm. Petals: Broad and laterally of equal length, anterior one longer; interporiferous zone very broad, distinctly raised. Oral side: With deep depression toward the peristome, food grooves indented; periproct near the posterior margin. Spines: Short and coarse. Color: In life, the sea urchin is covered by dark brown spines, the denuded test is white. Biology: Clypeaster rosaceus lives epifaunally in sea grass beds, feeding on dead pieces of plants or algae. It covers its test with fragments of shells or plants, holding them using its aboral tube feet. It is probable that the urchin drops these pieces each night, when it is feeding in the dark, picking up a new supply of local objects each day shortly after sunrise. By grinding large sediment

▸ Fig. 3.45: Clypeaster rangianus. (A) Aboral side with spines, length 89 mm, Rolas Island, San Tomè, Bay of Guinea. ZMH. (B) Oral side. (C) Side view. (D) Aboral side of naked test, length 97 mm, Miamia, Ghana. (E) Oral side. (F) Side view. Photos courtesy of H. van Noordenburg. (G) Part of petal: the primary tubercles are scattered within densely set miliaries. Scale bar is 4 mm, after Mortensen (1948b).

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

A

B

C

D

E

G F

 59

60 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

A

B

C

D

E

Fig. 3.46: Clypeaster rarispinus. (A) Aboral side of test: the petals are relatively narrow. Test length 66 mm, Persian Gulf. ZMUC. The margin is distinctly thickened. However, specimens with thin and flat edges are also found as well as all intermediates. (B) Oral side. (C) Side view. (D) Aboral side of test, length 56 mm; off Natal, South Africa. ZMUC. The margin is strongly thickened. (E) Part of petal: the ridges between the pore pairs carry only one primary tubercle.

particles, C. rosaceus may play an important role in the production of fine reef sediment. A single individual is able to transform 5.5 kg of coarse sand into fine sand each year. Distribution: This species is known from South Carolina, USA through the West Indies to Barbados and from Texas to the coast of Venezuela. It prefers shallow

water from the littoral zone to a depth of approximately 50 m. Remarks: Colloquially known as “sea biscuit”, C. rosaceus is inflated in contrast to the flattened and much larger C. subdepressus living in the same region. Literature: Kier & Grant 1965; Serafy 1979; Kampfer & Tertschnig 1992; Hendler et al. 1995 (Fig. 3.48).

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

A

 61

B

C D

E

F

G Fig. 3.47: (A–C) Clypeaster ravenelii. (A) Aboral side of test, length 80 mm; Gulf of Mexico. MfN Berlin. The margin is thickened. (B) Oral side. (C) Side view. (D–H) Clypeaster reticulatus. (D) Aboral side of test, length 53 mm; Hawaii. The thickened margin and the ring-shaped depression is distinct. (E) The oral side is depressed like a round bowl. (F) Aboral side with spines, test length 54 mm; Ile de la Réunion, Indian Ocean. The spines are often spotted. (G, H) Two profiles: the margin may be more or less thick. Scale bar is 30 mm. Stippled lines indicate oral depressions.

Clypeaster rotundus A. Agassiz, 1863 Test: Usually slightly oval, sometimes circular with slightly concave interambulacral edges; apex domed; margin thin and rather sharp; aboral side of test strong, in contradiction to the very fragile oral side; length up to 185 mm. Petals: Broad, pore zones rather narrow, petals more or less open; ridges between pore pairs densely set with a

row of primary tubercles extending almost over the entire height; petaloid area a little more than half the test length. Oral side: In the main flat, sometimes concave, abruptly sunken toward the peristome; periproct close to the posterior margin. Tuberculation: Primary tubercles small and very densely distributed, leaving little space for scattered miliary tubercles.

62 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

B

A

C

D

E

F

G Fig. 3.48: (A–C) Clypeaster rosaceus. (A) Aboral side of test, length 132 mm, width 92 mm, height 46 mm; Caribbean. The petals are broad and strongly inflated. (B) Oral side: the peristome is deeply sunken, the food grooves sharply indented. (C) Profile: the oral depression (stippled line) is very deep. Scale bar is 50 mm. (D–G) Clypeaster rotundus. (D) Aboral side of test with spines, length 135 mm, Port Culebra, Costa Rica, Pacific. ZMUC. (E) Oral side. (F) Aboral side of test, length 97 mm, Panama, Pacific. ZMUC. (G) Profile of same specimen, anterior side to the right. Scale bar is 3 cm.

Color: In living specimens, test and spines are more or less dark brown to almost black. On denuded tests, the plate sutures may be lighter. Distribution: Gulf of California to Ecuador and Galapagos Islands, littoral to a depth of 90 m.

Clypeaster speciosus Verrill, 1870 Test: Strong and very low, slightly raised toward the apical system, sometimes with a central peak; outline elongated pentagonal; at the posterior end shallowly notched; length reaches 120 mm occasionally.



Petals: Lancet-shaped, only slightly opened, faintly inflated. Oral side: Totally flat except for a small, abrupt depression toward the mouth; food grooves indented; periproct near the posterior margin in a small, but distinct depression. Spines: Short and densely distributed. Color: In life, the species is dark purplish, almost black, young specimens are lighter in a reddish-violet. The denuded and well cleaned test is whitish. Distribution: C. speciosus is typical for the Gulf of California on the west coast of Mexico, where it is rather common. It is also recorded from the Clarion Island from the tide zone to a depth of 90 m. Remarks: This species differs from C. europacificus by its thicker margin and by the test being on the whole stronger. A third species occurs along the American west coast: Clypeaster rotundus A. Agassiz, 1863; with a broadly oval or rounded test, the oral side flat or slightly concave, abruptly sunken toward the peristome (Fig. 3.49). Clypeaster subdepressus (Gray, 1825) Test: Very large and flattened, only central area raised; margin thin; circumference ovate or elongated pentagonal; maximum length 300 mm. Petals: Broad and of equal shape, distally closed and pointed, only slightly inflated; petaloid area a little more than half the width of the test. Oral side: Almost flattened, only the peristome is depressed, food grooves distinct; periproct close to the posterior margin. Canopy of spines very dense. Color: In life, the test is a yellow to dark tan, the bare test is whitish. Biology: The species lives in sandy or shelly sediment in which no sea grass grows. The flat sea urchins burrow just beneath the surface of the sand. After excavation, they are able to rebury within 6 to 12 min. The helmet conch Cassis tuberosa is a major predator. Large individuals that are turned upside down need approximately 1 h to right again (Hendler et al. 1995). Distribution: The species is found from North Carolina, USA, through the Caribbean, and along the east coast of Central and South America to Rio de Janeiro, Brazil. Remarks: C. subdepressus is easily distinguished from C. rosaceus by its very large and flattened test. Clypeaster telurus H.L. Clark, 1914 Test: Elongated and more or less rounded pentagonal; aboral side gradually rising from the margin toward the convex apex, height variable; length up to 157 mm. Petals: Ovoid and more or less open, interporiferous zone slightly inflated; petaloid area about half the test length.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

 63

Oral side: On the whole concave, slightly sunken toward the peristome. Periproct: More than its own length distant from posterior margin. Tuberculation: Primary tubercles small and densely distributed; miliaries very small. Distribution: Northern part of Australia from Fremantle in the West to Fraser Island on the east coast; living in shallow water and down to 40 m (Fig. 3.50). Remarks: Two of the Australian species of Clypeaster are rather similar, and the variation in both is high. In Clypeaster telurus the petals are usually shorter than in Clypeaster australasiae and the periproct is farther from the posterior edge. With the straight and open petals Clypeaster tumidus looks quite different. Clypeaster tumidus Tenison-Woods, 1878 Test: High and regularly arched, oral side deeply concave; outline slightly ovoid; may reach a length of 150 mm. Petals: Quite widely open, pore zones very long and straight, diverging distally. Oral side: Food grooves deeply depressed, periproct marginal. Primary spines: Short and very numerous. Distribution: It is found in the tropical east coast of Australia and in the Tasman Sea, from the sublittoral to a depth of 500 m. Literature: Rowe & Gates 1998; Miskelly 2002; van Noordenburg 2008. Clypeaster cf. tumidus Test: Almost circular, aboral side rounded conical, more or less high; margin relatively thin; rather large form. Apical system: Inflated, forming a knob. Petals: Very long and widely open, anterior petal the shortest, pore zones broad and only very slightly bowed; ridge between pore pairs set with only a few or no primary tubercles. Oral side: Flat, conspicuously sunken toward peristome; food grooves deeply depressed, extending almost to the margin. Primary spines: Aborally sparse. Distribution: Found in the Philippines, at deeper water. Clypeaster virescens Döderlein, 1885 Test: Longer than wide, ovoid to rounded pentagonal; margin thin but rounded; more or less highly domed or subconically raised; length up to 135 mm. Petals: Broad, paired petals remain slightly open, anterior one widely open; pore zones turning inward distally; ridges between pore pairs set with three to four primary

64 

A

 3.2 Superorder Neognathostomata A.B. Smith, 1981

B

C

D

E

F

Fig. 3.49: (A–C) Clypeaster speciosus. (A) Aboral side of test, length 118 mm, width 108 mm, height 25 mm; Baja California, Mexico. The test has been damaged (left side) and healed again. (B) Oral side: the flat lower surface is abruptly sunken toward the peristome. The food grooves are long, reaching the margin. (C) Profile: the apical peak may be unusual; generally, the aboral side is low vaulted (interrupted line). Scale bar is 50 mm. (D–F) Clypeaster subdepressus. (D) Aboral side, test length 168 mm, height 36 mm; Gulf of Mexico, Florida, USA. The petals are distally closed and pointed. (E) Oral side: the surface is almost perfectly flat with only a small depression toward the peristome. (F) Profile of test: the central area of the test is raised, the margin is rather thin. Stippled line indicates the oral depression. Scale bar is 50 mm.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

 65

B

A

C D F

G

E

H

I

Fig. 3.50: (A–C) Clypeaster telurus. (A) Aboral side of test, length 101 mm; Abrolhos Islands. Western Australia. (B) Oral side. (C) Side view: the test is evenly domed. (D) Clypeaster tumidus, oblique view of aboral side, length of test 71.9 mm, width 64.0 mm; off Stylaster and Juneaux Seamounts, south of Norfolk Ridge, Tasman Sea. Photo courtesy of A. Miskelly. (E–G) Clypeaster cf. tumidus. (E) Aboral side of test, length 99 mm, Palawan, Philippines. The long petals are quite open with almost straight pore zones. (F) Oral side. The grooves are deeply recessed. (G) Side view. Photos courtesy of H. van Noordenburg.

66 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

tubercles; interporiferous zones inflated in adults, in young specimens flush with the test; petaloid area about half the test length. Oral side: Deeply sunken toward the peristome; food grooves almost reaching the margin; periproct close to the posterior margin.

A

Tuberculation: Scattered primary tubercles within a dense coat of miliaries. Color: Test and spines are yellow-brown. Distribution: The species is recorded from southern Japan to the Malayan Archipelago, east Australia and New Zealand; at a depth of 100 to 350 m (Fig. 3.51).

B

D C

E

Fig. 3.51: Clypeaster virescens. (A) Aboral side of test, length 122 mm, Kei Islands, Indonesia. ZMUC. The paired petals are distally bent inwards, the anterior petal is widely open. (B) Oral side. (C) Side view: the test is subconically raised. In other specimens, the test may be considerably higher. (D) Aboral side of a young specimen with spines, length 52 mm, Bohol, Philippines. (E) Part of petal: the ridge between the pore pairs is set with a few primary tubercles. Scale bar is 3 mm, after Mortensen (1948a,b).

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

Suborder Scutellina Haeckel, 1896 The members of this taxon are provided with one fused lantern support on the internal side of the single interambulacral plate next to the peristome (Fig. 3.52). This suborder is divided into two infraorders: infraorder Laganiformes with four families (Echinocyamidae, Fibulariidae, Laganidae, and Rotulidae) and infraorder Scutelliformes with six families (Astriclypeidae, Dendrasteridae, Echinarachniidae, Mellitidae, Scutellidae, and Taiwanasteridae). Literature: Clark, H.L. 1914, 1938; de Meijere 1902, 1903; Mortensen 1907, 1948; Durham 1955; A. Clark & Rowe 1971; Telford et al. 1983; Shigei 1986; Mooi 1986b; 1990; Rowe & Gates 1995; Miskelly 2002; Kroh & Smith 2010.

Infraorder Laganiformes Desor, 1847 Characteristics: The aboral interambulacra ends with one or more single plates at the apical system. In the

A

D

ambulacra, the pore pairs of the respiratory podia cross the suture. The tips of the miliary spines are crownshaped. The infraorder is composed of four families with species of rather different shapes. Family Echinocyamidae: Two genera, worldwide. Family Fibulariidae: Four genera, worldwide. Family Laganidae: Two genera in the Indo-Pacific. Family Rotulidae: Two genera, restricted to West Africa. The placement of the Rotulidae in Laganiformes or in Scutelliformes is the subject of discussion (Mooi 1990). According to Mooi (confirmed in 2015), there are important characteristics that the rotulids share with the laganiforms, despite their superficial resemblance with the scutelliforms in having holes and notches. In particular, the arrangement of the respiratory podia pores, which cross the plate suture, is a strong synapomorphy unique to the laganiforms including the rotulids. Furthermore, in both taxa, the interambulacra ends with a single plate at the apical system. In the scutelliforms, the respiratory podia pores are positioned directly along the sutures, and

C

B

E

 67

F

G

Fig. 3.52: Upper side of Scutellina with interambulacrum IA 2 and adjacent ambulacra: (A,B) Laganiformes: (A) Laganum laganum. The interambulacral column ends in one plate at the apical disc. (B) Rotula deciesdigitata. The interambulacral column ends in a few single plates at the apical disc. (C) Scutelliformes: the interambulacral column ends with two plates at the apical disc. Encope grandis. (A, B) Scales 10 mm, interambulacra shaded. (A–C after Durham 1955). (D–G) Internal buttresses of Laganiformes: (D) Fibulariella, 6.6 mm. In Fibularia, there are also no buttresses. (E) Echinocyamus megapetalus, 7.0 mm (F) Echinocyamus platytatus, 6.0 mm. (D–F from H.L. Clark 1914). (G) Mortonia australis, 14 mm. Two buttresses only in the posterior interambulacrum.

68 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

the ambulacra are double-columned. This discussion is definitely not finished. Remarks: There are no differences between Fibularia, Fibulariella, and Echinocyamus in the test structure itself except for the interior. As one finds predominantly naked tests without periproctal or peristomial membranes or plates, the distinction may be very difficult.

3.2.2.3 Family Echinocyamidae Lambert & Thiéry, 1914 The family consists of two genera, small to very small forms of rather similar appearance: Echinocyamus van Phelsum, 1774, with many extant species worldwide, and Mortonia Gray, 1851, with two species, central and southwest Pacific. The two genera differ by the plate pattern and size of the periproct: in Echinocyamus, the periproct is covered by several radially arranged naked plates and it is smaller than the peristome. In Mortonia, the periproct is enclosed by numerous small platelets set with miliary spines and it is at least as large as but often larger than the peristome. Biology: These small sea urchins live in rather coarse sand. They nestle between larger sand grains, and cover the free parts of their tests with other particles. They mostly move at night, selectively taking individual sand grains and grazing the organic coating. Echinocyamus does not use any specialized tube feet or spines for feeding as do the typical clypeasteroids, and no food grooves are developed. In contrast, the free margin of the thickened peristomial membrane serves as a set of mobile lips, which hold the substrate particles in place while the lantern teeth strip away the encrusting diatoms. These species are an important link in the food net; for example, large numbers of Echinocyamus pusillus are eaten by Spatangus purpureus. The accessory podia of Echinocyamus and Mortonia with their suckered discs do not serve for food gathering as in the Rotulidae and the Scutellina but they move substrate particles over both the oral and aboral surfaces, when the animals are exposed. They retain the complete canopy of small sediment grains held firmly against the tips of the spines, even when they are totally buried (Telford et al. 1983). Genus Echinocyamus van Phelsum, 1774 Test: Small to very small, oval to subcircular in outline, thick margin, aboral and oral side more or less flattened. Apical system: Four gonopores and a single or only a small number of hydropores. Ambulacra: Petals small, but usually distinct; open distally; accessory pores with tube feet in bands along the depressed horizontal sutures of ambulacra, extending onto the interambulacra.

Oral side: More or less sunken toward the peristome; peristomial membrane naked; periproct smaller than peristome; covered by several radiating, naked plates. Ten simple, radially arranged internal buttresses along the interambulacral margin. Primary tubercles: Uniformly covering the test, below the petals in horizontal series; numerous miliary tubercles. Spines: Short and usually simple. Mortensen (1948a,b) listed 15 species, and one was added from the southeast Pacific by Mironov & Sagaidachny (1984). All are very similar and difficult to distinguish: Echinocyamus apicatus Mortensen, 1948 Maximum length 6 mm; test oval, raised toward the apical disc; posterior edge truncate or even concave; aboral side regularly arched to the anterior margin, sloping flatter to the posterior side; genital pores in females much larger than in males; oral side sunken; petals with five to six pore pairs. Southeast Australia; 100 m (Fig. 3.53A). Echinocyamus convergens Mortensen, 1948 Only a single naked test of 9.5 mm known; elongate ovoid in outline, flattened above and below, very slightly concave between peristome and periproct, periproct slightly nearer to posterior end than to mouth; petals short, but well-developed, posterior pair converging. Providence Island, Seychelles, west Indian Ocean; 128 m (Fig. 3.53C). Echinocyamus crispus Mazzetti, 1893 Test: Small, aboral side flattened or with a distinct vertex; maximum length 10.5 mm. Apical system: A single hydropore. Ambulacra: Petaloid area approximately two-thirds of the test length; petals well-developed but rather short, broad with slightly curved pore zones, distally open; up to 10 pore pairs; accessory pores in rows along the depressed horizontal sutures. Oral side: Deeply sunken similar to a bowl reaching to the periproct and including a small portion in front of the peristome; periproct about halfway between mouth and posterior end. A few very small primary tubercles among a dense cover of miliary tubercles. Color: Spines and test are yellow-white. Distribution: The species is reported from East Africa and the Red Sea over the Malayan region and the Philippines to Japan and the Hawaii Islands; bathymetric range 20 to 124 m. Remarks: E. crispus is similar to Echinocyamus megapetalus in having a deep bowl-like depression on the oral side. However, that species differs from E. crispus mainly by its

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

A

B

E

C

 69

D

F

G

Fig. 3.53: (A–D) Petaloid area of: (A) Echinocyamus apicatus, female. (B) Echinocyamus crispus, (C) Echinocyamus convergens, (D) Echinocyamus elegans. All scale bars are 2 mm, after Mortensen (1948a,b). (E–G) E. crispus. (E) Aboral side of test, length 7.5 mm; Tubod, Philippines. The petals are broad and short, the pore zones slightly curved. (F) Oral side: there is a bowl-shaped depression toward the peristome. (G) Side view: there is a distinct vertex.

broad and long petals, which have nearly straight pore zones. In E. crispus, the petals are broad, but short with slightly curved pore zones (Fig. 3.53E). Echinocyamus elegans Mazzetti, 1893 Maximum length 6 mm; test subcircular, rounded pentagonal or slightly elongate, aboral side convex, oral side sunken around peristome; petals well-developed with rather large pores; peristome pentagonal, periproct much

smaller, close to the posterior edge. Red Sea and Iranian Gulf southward to Natal; 110 to 275 m (Fig. 3.53D). Echinocyamus grandiporus Mortensen, 1907 Test in outline subcircular; madreporite slightly inflated; ocular pores at least as large as genital pores; petals poorly developed; periproct transversely elongated near the posterior edge. Florida and West Indies to Brazil, Azores; 150 to 1500 m (Fig. 3.54).

70 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

A

D

F

C

B

E

G



Echinocyamus grandis H.L. Clark, 1925 Test oval, maximum 14 mm; aborally evenly arched; genital pores larger than petaloid pores; petals welldeveloped, 11 to 14 pore pairs per column; oral side concave, the peristome deeply and abruptly sunken; peristome round, periproct nearer to mouth than to edge. Seychelles, Indian Ocean. Echinocyamus incertus H.L. Clark, 1914 Test 6 mm, ovoid, aboral side flat, oral side not concave, petals poorly formed, four pore pairs; periproct closer to posterior end than to mouth; likely to have sexual dimorphism, the single specimen having larger gonopores than petaloid pores. Hawaiian Islands. Echinocyamus insularis Mironov & Sagaidachny, 1984 Easter Islands and Sala-y-Gomez, Southeast Pacific; 60 to 80 m. Echinocyamus macrostomus Mortensen, 1907 Test in outline slightly elongated; madreporite elevated, ocular pores not large; petals scarcely developed; peristome much larger than periproct, periproct itself small, very close to posterior edge, transversely elongated. West Indies, Azores, and Cape Verde Islands; 1600 to 2286 m. Echinocyamus megapetalus H.L. Clark, 1914 Maximum length 11 mm, test thin and fragile, gonopores in females larger than in males; petals well-developed with up to 12 pore pairs, broad with nearly straight pore zones and widely open distally, reaching almost to the margin; oral side distinctly sunken toward elongated pentagonal peristome; periproct subcircular, nearer to the mouth than to the posterior end. From Réunion over the Malayan region to the Philippines and Hawaii; 2 to 30 m. Echinocyamus planissimus H.L. Clark, 1938 Maximum test length 8 mm; upper side flattened, petaloid area hardly elevated; several hydropores; periproct separated from margin by more than its own width. From Broome around the South of Australia to New South Wales; 10 to 100 m (Fig. 3.55B). Echinocyamus platytatus H.L. Clark, 1914 Maximum length 9.5 mm; test low and broad, flattened above and below, not concave orally; genital pores in

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

 71

females large, also in males larger than petaloid pores; petals with a few very small pore pairs; periproct round and close to the posterior edge; glassy granules not very conspicuous. New South Wales to South Australia and Tasmania; 20 to 365 m (Fig. 3.55A). Echinocyamus provectus de Meijere, 1903, 1914 Maximum length 8.5 mm; periproct elongate, midway between mouth and posterior margin, conspicuous glassy protuberances. Southeast Africa to Malayan region; 75 to 390 m (Fig. 3.55C). Echinocyamus pusillus O.F. Müller 1776 Test small, reaching up to 12 mm; shape varying; one hydropore in the apical system; pores of petals small; oral side slightly depressed toward peristome; periproct small, midway between peristome and margin. European Atlantic from northern Norway and Iceland to Cape Bojador, West Africa and the Azores, in the North Sea and western Baltic Sea, and the Mediterranean; from the littoral zone to a depth of 1250 m. Echinocyamus scaber de Meijere, 1902, 1903 Length may reach 8 mm; outline ovoid, aboral side lowly convex with apical disc inflated; petals well-developed with four to five pore pairs; oral side sunken toward the peristome; periproct small and round, halfway between mouth and posterior end; glassy granules are scattered on aboral and oral side, distinctly higher than the primary tubercles. Widely distributed from East Africa to the Philippines, eastern Australia and Hawaii; 200 to 1886 m. Echinocyamus sollers Koehler, 1922 Test nearly round, oral side nearly flat; petals broad and rather long, pore zones straight and parallel; accessory pores indistinct, depressed sutures as in E. crispus. Off Ceylon and the Malabar Coast; 260 to 880 m (Fig. 3.55H). Genus Mortonia Gray, 1851 Two very similar species belong to this genus. In Mortonia australis, the internal buttresses are restricted to two partitions bordering the periproct; in Mortonia polyporus, they are only rudimentary. Test: Small, fairly strong; aboral side lowly vaulted, the anterior side with thick, rounded margin, sloping posteriorly toward a thinner end; test length up to 13.5 mm.

◂ Fig. 3.54: (A–C) Echinocyamus grandiporus. (A) Oral side of test, length 7.5 mm. (B) Aboral side. (C) Internal view. (D,E) Echinocyamus macrostomus. Young specimen, length 4.75 mm. (D) Aboral side. (E) Oral side. Both surfaces show the lines of accessory podia. (A–E) from Mortensen (1907). (F,G) Echinocyamus megapetalus. (F) Aboral side of test, length 7.5 mm; Philippines. (G) Oral side. Photos courtesy of A. Miskelly.

72 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

A

B

F

C

E

D

G

H

Fig. 3.55: (A) Echinocyamus platytatus, female, length 6 mm, Victoria, southeast Australia. From left to right: aboral side, side view, oral side. From H.L. Clark (1914). (B,C) Apical system of (B) Echinocyamus planissimus and (C) Echinocyamus provectus. Scale bars are 2 mm; after Mortensen (1948a,b). (D–G) Echinocyamus pusillus. Test with spines, length 6 mm; Mediterranean, Turkey. (D) On the aboral side, the spines are short. (E) On the oral side, they are longer, serving for locomotion. (F) Aboral side of test, length 11 mm; Helgoland Island, North Sea, Germany. The pore pairs of the petals are small. Only one hydropore is developed in the madreporite. (G) Oral side: the peristome is slightly sunken, the periproct is much smaller than the peristome. (H) Echinocyamus sollers. Aboral side of test, length 8 mm. Malabar Coast, India. The petals are long and broad, the pore zones parallel, distal most slightly diverging. From Koehler (1922).

Apical system: Four large gonopores in both sexes and a single hydropore. Petals: Long, pore zones almost parallel, distally turning slightly outwards. Oral side: Sunken at the round peristome, rising gently toward the round periproct and to the posterior edge; anterior part convex continuing to the rounded margin;

peristome and periproct of same size, periproct with small spined platelets. Tubercles: Small and uniform. The genus differs from Echinocyamus by the size of the periproct: in Echinocyamus, the periproct is smaller than the peristome; in Mortonia, it is as large as the peristome, often considerably larger. The second difference lies in the plating



of the periproct: in Mortonia, many irregular platelets set with small miliary spines are developed. In Echinocyamus, several larger, smooth plates are radially arranged. Mortonia australis (Desmoulins, 1837) Length of test may reach 15 mm. Distribution: The species is widely known in the IndoPacific, from Mauritius to the central Pacific. Biology: The sea urchins live shallowly buried in very clean coral sand. Many accessory tube feet are developed in the ambulacra. It is likely that they hold by means of their suckered discs tiny sediment particles over the aboral and oral surfaces just above the tips of the spines. Below this dense canopy, a water current flows over the surface to provide all the appendages, such as the spines, podia, and pedicellariae, with oxygen that they absorb through their skins. It is the same mode as in E. pusillus (Telford et al. 1983) (Fig. 3.56). Mortonia polyporus Mortensen, 1921 Internal buttresses rudimentary, restricted to 10 shallow thickenings inside the very edge, posteriorly more distinct than anteriorly. Distribution: The species is recorded from New Zealand (Cook Strait) and the Kermadec Islands, over the Tasman Sea to New South Wales, southeast Australia; bathymetric distribution 9 to 536 m. 3.2.2.4 Family Fibulariidae Gray, 1855 The species of this family are miniaturized forms, not more than 20 mm long and with a more or less inflated test. No or only simple, radial, internal buttresses are developed, and the food grooves are either absent or restricted to a single, small furrow in the anterior ambulacrum. They live epifaunally among the sand grains of the surface or shallowly buried in coarse, clean sediment. They manipulate individual grains grazing on their organic coating. Occurring in large numbers, they are preyed upon by many other animals. There are two genera: Fibularia and Fibulariella. Genus Fibularia Lamarck, 1816 Test: Small to very small forms; more or less inflated; outline oval to ovoid. Apical system: Four gonopores, few hydropores, sometimes in a groove. Petals: Small, but distinct; accessory podia hardly discernable. Oral side: More or less convex, peristome slightly sunken, membrane naked; periproct covered by five valve-like, naked plates. No internal buttresses. Spines: Short and uniform.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

 73

Six extant species are described, which are restricted to the Indo-West Pacific. Some of the species show sexual dimorphism: the female having distinctly larger gonopores than the male. Females in these species emerge from larger yolk-rich eggs, which develop directly without a larval stage or metamorphosis. Fibularia nutriens is a brood-protecting species with a deep brood pouch on the aboral side. Fibularia cribellum H.L Clark, 1928. Test: Very small, solid, outline oval to slightly ovoid, aboral and oral side slightly convex, margin thick and round; length up to 10 mm. Apical system: Subcentral, the two hydropores lie in a groove; four gonopores, in the female much larger than in the male. Petals: With large, depressed pores, rather distant from each other, the pairs are difficult to recognize; petaloid area almost covering the whole aboral side (it looks like a sieve); below the petals there may be depressions with accessory pores. Oral side: Slightly sunken toward the large, round peristome, periproct subcircular, smaller than and nearer to peristome. Primary tubercles: Very small and irregularly distribu­ ted, also between the pores; numerous, glassy miliary tubercles. Primary spines: Short and smooth. Color: The test and the spines are white or pale yellow to brown. Distribution: The species is recorded from the Malayan region and the Philippines, at a depth of 15 to 400 m (Fig. 3.57). Fibularia japonica Shigei, 1982 Test: Vaulted; barely sunken toward the peristome; aboral side flattened; length up to 10 mm. Apical system: Female with larger gonopores than the male; hydropores in an elongated furrow. Petals: With only two to three pore pairs. Oral side: Periproct at mid-length between peristome and posterior margin, smaller than peristome. Color: Whitish to light brown. Distribution: This species is not endemic to Japan, but is also reported in the Philippines (Mooi, personal communication). The bathymetrical range reaches from 30 to 100 m. Remarks: The similar Fibularia cribellum, found in the same region, is flatter and has larger pores (Fig. 3.58). Fibularia nutriens H.L. Clark, 1909 Test: Extremely small, upper side convex, flattened at the apical system; oral side flat; outline oval; length only up to 3.3 mm.

74 

A

C

 3.2 Superorder Neognathostomata A.B. Smith, 1981

B

D

E

Fig. 3.56: (A, B) Mortonia australis. (A) Aboral side of test, length 11 mm; Hawaii. (B) Oral side: the fields with the accessory podia are visible. (C–E) Mortonia polyporus. Aboral side of test, length 12 mm, Cook Strait, New Zealand. ZUMC. (C) Aboral side: the petals are long with almost parallel pore zones. (D) Oral side: the periproct is as large as the peristome. (E) Side view.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

A

B

D

 75

C

E

Fig. 3.57: Fibularia cribellum. (A) Aboral side of male, length of test 5.9 mm and (B) aboral side of female, length 6.1 mm; Siquijor, Philippines. The upper surface looks like a sieve. The males have very small gonopores; in females, the gonopores are much larger being only slightly smaller than the petaloid pores. (C) Oral side of male. (D) Side view of male. Upper and underside are slightly convex. (E) Petaloid area: there are only two pore pairs in a column, some pores remain single. The pores forming a pair are connected by a gray line. Scale bar is 2 mm, after Mortensen (1948a,b).

Apical system: Females have a deep, horseshoe-shaped brood pouch containing the apical system with four gonopores; no petaloid pore pairs for respiratory podia; in males, sometimes there are five depressions around the apical system, petals rudimentary. Oral side: Peristome round to pentagonal, periproct round or oval, midway between peristome and posterior margin. Spines: Short, about the same length on oral and aboral sides. Color: In life, the spines are light brown, the denuded test is whitish. Distribution: This species is restricted to southeast Australia from Newcastle, NSW to Victoria. It lives at a depth of 5 to 170 m. Biology: The female shelters the juveniles in its brood chamber. From the yolk-rich egg, the young develop directly into little sea urchins without a planktonic larval stage.

Fibularia ovulum Lamarck, 1816. Test: Small; highly inflated; upper side slightly flattened, oral side convex; outline round or slightly oval; length may reach 10 mm. Apical system: With four gonopores, few hydropores in a short, longitudinal furrow. Petals: Short, only a few pore pairs in each series. Oral side: Peristome round, not at all depressed; periproct round or transversely oval. Color: The short spines and the denuded test are whitish. Distribution: F. ovulum is a very common species and is widely distributed in the Indo-West Pacific from the Red Sea and East Africa to the Malayan Archipelago, the Philippines and northeast Australia from the intertidal zone to a depth of 385 m. Remarks: The highly inflated test distinguishes this species from other fibulariids (Fig. 3.59).

76 

A

 3.2 Superorder Neognathostomata A.B. Smith, 1981

C

B

D

apical disc

peristome

E

F

periproct

G

Fig. 3.58: (A–D) Fibularia japonica. (A) Aboral side of test, length of female 8.9 mm; Shizuoka, central Japan. (B) Aboral side of male, test length 9 mm. (C) Oral side of male. The four gonopores of the female are much larger than in the male. The hydropores open into an elongated groove. (D) Profile of the test 1 mm. (E–G) Fibularia nutriens. (E) Aboral side of female, test length 3.4 mm; Southeast Australia. The deep brood pouch contains the large gonopores and there are no respiratory podia arranged in petals. (F) Oral side of female: the peristome is pentagonal, the periproct lies midway between the peristome and the posterior margin. (G) Aboral side of male, test length 3.9 mm; Cronulla, New South Wales, Australia. Four small gonopores and the hydropore are present in the apical system. The large hole has been drilled by a predator, probably by a snail. There are five shallow depressions around the apical disc.

Fibularia plateia H.L. Clark, 1928 Test: Low and flattened; posterior end somewhat truncated; test length up to 9 mm. Apical system: Female with larger gonopores than the male; one hydropore in the madreporite. Periproct: Distinctly square. Distribution: This species is known only in Australia from Queensland around the south coast to Bunbury, Western Australia, at a depth between 5 and 235 m. Fibularia volva L. Agassiz & Desor, 1841 Test: Slightly flattened, somewhat pointed at each end; length reaching 10 mm. Apical system: Several hydropores. Distribution: Indo-West Pacific, shallow water to 50 m. Genus Fibulariella Mortensen, 1948 The genus is characterized by the lack of internal butresses. The peristomial membrane is set with plates, and the

periproct is covered by small and numerous naked platelets. That means the genus differs from Fibularia through characteristics that are not associated with the test itself. Naked tests are difficult to distinguish. There are three species. Fibulariella acuta (Yoshiwara, 1898) Test: Small, aboral side convex and of variable height; outline oval, anterior and posterior end usually pointed; test length up to 15 mm. Apical system: Slightly anterior, six to eight hydropores scattered over the madreporite. Ambulacra: Paired petals short with four to six pore pairs, anterior one longer with seven or eight in each row. Oral side: Depressed toward peristome; food groove like a simple short trunk only in anterior ambulacrum; periproct more or less elongated, closer to the peristome than to the posterior margin. Spines: Less than 1 mm in length; slightly longer on the oral side.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

A

B

C

apical disc

peristome

D

 77

periproct

E

Fig. 3.59: (A–C) Fibularia ovulum. (A) Aboral side of test, length 8.9 mm; South Malé Atoll, Maldives. There are only a few pore pairs on the petals. The round hole at the posterior right side is a drill hole made by a predatory snail. (B) Oral side: the periproct lies mid-length between the peristome and the posterior margin. (C) Profile: the test is highly inflated. Scale bar is 2 mm. (D–E) Fibularia plateia. (D) Aboral side of test, length 7 mm; Queensland, Australia. The petals are short but distinct. (E) Oral side: the periproct is more or less square.

Color: The spines are grayish to whitish, the denuded test is white. Distribution: This species is known from Japan and the Korean Strait to China, the Philippines and the Gulf of Siam, to the Banda Sea and northeast Australia at depths from 5 to 90 m. Biology: F. acuta lives in coarse, clean sand in calm water, covering itself completely with sand grains. It feeds selectively on organic particles in the substrate.

Remarks: The usually pointed test and the single food groove distinguish F. acuta from other fibulariids (Fig. 3.60). Fibulariella angulipora Mortensen, 1948 Test: Oval, rounded pointed at both ends; in profile highly inflated but flattened above and convex below, anterior side with very thick, rounded margin, a little lower toward the posterior end; maximum length 9.5 mm.

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 3.2 Superorder Neognathostomata A.B. Smith, 1981

Apical system: Subcentral, gonopores small, of same size as petaloid pores, six to eight hydropores scattered on the madreporite near the anterior gonopores. Ambulacra: Petals short, three to four pore pairs in the paired petals, five to six anteriorly; pores large and distinctly triangular. Oral side: Convex, peristome subcentral, subcircular, membrane set with a varying number of plates, densely packed to scattered; a short, simple food groove may be present; periproct slightly elongated, closer to the peristome than to the posterior. Spines: Primary and miliary spines form a dense, uniform coat. Distribution: Gulf of Siam to the Moluccan Sea; 15 m.

A

E

B

Fibulariella oblonga (Gray, 1851) Shape of the test similar to F. ovulum, ovoid and highly inflated, but the periproct is elongate-oval, not round or transversely ovate. Distribution: East Indo-West Pacific; probably 24 to 100 m. 3.2.2.5 Family Laganidae A. Agassiz, 1873 The species from the Laganidae family include small to large forms of flat sand dollars. The internal buttresses are composed of branched radial partitions and circumferential walls. Five, more or less long, simple, radial food grooves are present on the oral side. The genera differ in the number of gonopores and in the position of the periproct. The laganids are widely distributed in the

D

C

F

G

H

Fig. 3.60: (A–D) Fibulariella acuta. (A) Aboral side with spines, length 9 mm. The short petals are marked by darker podia. (B) Oral side of specimen with a length of 7.9 mm. The spines are longest around the peristome. (C) Aboral side of naked test, length 7.1 mm. The petals are short and set with few pore pairs, several hydropores are scattered over the madreporite. (D) Oral side of naked test, length 8 mm: there is a short, simple food groove in the anterior ambulacrum. All specimens from Sydney, New South Wales, Australia. (E–H) Fibulariella angulipora. (E) Aboral side of test, length 7.5 mm; Gulf of Siam. (F) Oral side. (G) Side view: the test is strongly inflated and flattened aborally. (H) Petaloid area: the anterior petal is the shortest. The pores forming a pair are connected by a gray line. Scale bar is 2 mm, after Mortensen (1948a,b).

3.2.2 Order Clypeasteroida L. Agassiz, 1835 



Indo-West Pacific, living mostly infaunally in fine sediments (Fig. 3.61). The family consists of two genera, which differ in the pattern of the apical system: Laganum Link, 1807 with 10 species. Peronella Gray, 1855 with 15 species. Literature: Döderlein 1885; Mortensen 1921; Koehler 1922; McKnight 1969; Shigei 1986; Rowe & Gates 1995. Genus Laganum Link, 1807 Test: Pentagonal to oval in outline; flattened; petaloid area more or less inflated; oral side flat or slightly concave. Apical system: Five gonopores, hydropores in a more or less long, sinuous groove. Petals: Distinct, more or less closed distally. Oral side: Peristome subcentral, periproct between peristome and posterior edge; food grooves distinct, straight and simple. Tuberculation: Dense and uniform, spines short, on the oral side longer. Ten species are described, and they strongly resemble each other. Laganum boninensis Mortensen, 1948 Test: Round to elliptical, center raised, distally flattened; recorded length up to 37 mm, but probably larger. Apical system: Slightly anterior; hydropores in an irregularly sunken space. Petals: Petaloid area less than half the test length; pore zones narrow. Oral side: Flat or slightly concave, food grooves distinct in larger specimens; peristome slightly anterior, pentagonal; periproct round or transverse oval, almost as large as the peristome, plates rather densely set with spines. Color: Yellow to brown in life, cleaned tests white. Occurrence: Restricted to the Bonin Islands, West Pacific; depth 120 to 220 m.

A

B

 79

Laganum centrale H.L. Clark, 1925 Test: Margin thin; length of holotype 46 mm. Petals: Petaloid area about half the test length, petals narrow and well open distally. Oral side: Periproct midway between peristome and posterior margin. Occurrence: Tonga Islands. Laganum decagonale (Blainville, 1827) Test: Strong, flat with raised apical system; outline with concave margins in the interambulacra; length may reach 56 mm. Apical system: Hydropores in a short furrow. Petals: Short and rather broad, distally closed. Oral side: Flat, trunks of food grooves short and broad; periproct round, closer to the posterior margin than to the peristome. Spines: Aboral primaries short and scattered over the surface, longer on oral side. Color: In life, the species is light brownish, sometimes with a reddish tint in the tube feet; the cleaned test is whitish. Distribution: This species is reported from the Malayan region and the Philippines, in the China Sea northward to Macao, in the Bay of Bengal, and in northern Australia and the Admiralty Islands at depths between 5 and 301 m (Fig. 3.62). Laganum depressum L. Agassiz, 1841 (syn. Jacksonaster depressum (L. Agassiz, 1841)) Test: Often very strong; flat with more or less thick, but always inflated margin, usually with ring-shaped depression inside the margin; center with apical system raised; outline ovate to pentagonal, maximum length usually 50 mm, exceptionally up to 85 mm. Apical system: Five gonopores, hydropores in a sinuous, branched groove. Fig. 3.61: Apical systems of laganids: (A) Laganum fudsiyama: very small ocular plates are developed at the sides of the monobasal plate, which is pierced by five genital pores. The hydropores open into a short furrow. The primary tubercles are large and surrounded by miliaries. (B) Peronella japonica: there are four gonopores, the scattered hydropores are restricted to the anterior right part of the madreporite. The primary tubercles are small. Scale bars are 2 mm, modified after Mortensen (1948a,b). Mortensen (pages 278 and 341) showed the ocular plates incorporated into the monobasal/ genital plate. That was one of his few errors.

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 3.2 Superorder Neognathostomata A.B. Smith, 1981

B

A

C

D

E

F Fig. 3.62: (A,B) Laganum decagonale. (A) Aboral side of test, length 44 mm, width 42 mm; Macao, China. MfN Berlin. The petals are broad and conspicuously short. (B) Oral side: the periproct is round and is closer to the posterior margin than to the peristome. (C–F) Laganum depressum. (C) Aboral side of test, length 33 mm; Pangani, Tanzania. A ring-shaped depression lies around the petaloid area. (D) Oral side: the periproct is slightly transverse oval. (E) Aboral side with spines, length of test 78 mm, Queensland, Australia. The aboral side is densely covered by short spines, the oral side has longer spines. The petals generally appear as darker bands. (F) Profile of C: the apex is distinctly raised.

Petals: Narrow and long, reaching 60% to 80% of the aboral surface; more or less closed distally. Oral side: Flat, trunks of food grooves broad; periproct transversely elongate; closer to the posterior margin than to the peristome. Spines: Short, longer on the oral side. Color: In life, the spines are light grayish-brown or grayish-­olive, often with darker pore zones, the bare test is whitish.

Distribution: L. depressum is known from the Red Sea and East Africa to the Tonga and Marshall Islands, from Macao, China, to the northern coast of Australia, living from the intertidal zone to a depth of 85 m. Remarks: L. depressum is distinguished from L. laganum by its transversely elongated periproct, in L. laganum it is longitudinally elongated. Some authors made of L. depressum a separate genus, Jacksonaster Lambert in Lambert & Thiéry, 1914, with the species J. depressum.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

Laganum fudsiyama Döderlein, 1885 Test: Rather fragile, flat with regularly raised center (name from Mount Fujiyama in Japan), more or less high, outline slightly oval, circular or angular, margin rounded; length up to 73 mm. Apical system: Small, subcentral, hydropores in curved groove. Petals: Narrow, straight or flexed, distally open or closed; petaloid area hardly half the length of test. Oral side: Flat to slightly concave; periproct round, plates naked. Primary tubercles in a matrix of miliary tubercles, orally longer. Color: Yellow to light brown or pale pink, often after death brightly green (as all clypeasterids). Distribution: East Africa to Malayan region, Philippines, southern Japan (as far north as Sagami Bay), and Hawaii; 50 to 645 m (Fig. 3.63).

A

 81

Laganum joubini Koehler, 1922 Test: Oval, center slightly raised, recorded length up to 47 mm. Petals: Rather short, tapering distally and almost closed. Oral side: Flat, peristome slightly anterior; periproctal plates with spines. Color: May be reddish or pale yellow. Distribution: Red Sea, East Africa, Mauritius, at a depth of 10 to 15 m. Laganum laganum (Leske, 1778) Test: flat with more or less thick margin; often slightly raised toward apical system; outline rounded pentagonally; test length up to 65 mm. Apical system: Hydropores in an irregularly sinuous groove. Petals: Well-developed, distally more or less closed.

B

D

C

Fig.3.63: (A–C) Laganum fudsiyama. (A) Aboral side of test, length 77 mm, off Taiwan, China Sea. The petals are narrow. (B) Oral side. (C) Side view: the test rises gently to the apex. (D) Laganum joubini. Aboral side of young specimen, length 24 mm, Mauritius. ZMUC.

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 3.2 Superorder Neognathostomata A.B. Smith, 1981

Oral side: Peristome central; five broad unbranched food grooves; periproct longitudinally elongate and opening midway between peristome and posterior margin. Color: In life, the echinoid is greenish-olive to light grayish, sometimes with irregular spots. The bare test is yellowishwhite. Distribution: This common species is reported in the Malayan region and the Philippines, to New Guinea, Australia and Tasmania in depths of between 0 and 34 m. Biology: L. laganum lives in the littoral shallowly buried in fine sand, and feeds on organic particles (Fig. 3.64).

A

Laganum putnami (A. Agassiz, 1863) Test round; length of holotype 28 mm; gonopores outside the apical system, petals lanceolate and pointed; food grooves long and conspicuous. Distribution: Japan. Laganum retinens Koehler, 1922 Test: Round to slightly elliptical, flat in profile with thin margin, regularly rising to the apical disc; length up to 32 mm. Apical system: Slightly anterior, hydropores in an irregular groove.

C

B

D E

F

Fig. 3.64: (A–D) Laganum laganum. (A) Aboral side of test, length 34 mm, height 6 mm; Negros, Philippines. At the apical system, the hydropores lie in a sinuous groove. There are five gonopores. (B) Oral side: the trunks of the food grooves are short and broad. The periproct is longitudinally oval. (C) Aboral side of test with spines; length 45 mm; Kei Islands, Indonesia. The spines are as short as velvet. (D) Profile of A: the test is only very faintly raised toward the apical disc. (E–F) Laganum retinens. (E) Aboral side of test, length 32 mm; Mactan Island, Philippines. The plate sutures are well visible. (F) Oral side. Photos courtesy of H. van Noordenburg.



Petals: Long, lyriform, distally diverging, pore pairs rather large; petaloid area approximately 65% of test length. Oral side: Slightly concave, peristome round, periproct small and angular, approximately 25% of distance to the mouth, plates naked. Color: Cream to yellowish, sometimes sutures darker, showing plate pattern. Distribution: From Mauritius to the Philippines; 110 to 120 m. Laganum versatile Koehler, 1922 Test slightly elliptical; length of holotype 27.5 mm; margin rather thick; conically raised petaloid area large, petals about three-fourths in length of radius, straight and open distally. Occurrence: Andaman Islands; depth 145 m. Genus Peronella Gray, 1855 Fifteen species are described, mostly from the Tropics. Test: Small to large, outline elliptical, round or polygonal; usually flat and raised only toward the apical system; margin may be thickened or thin. Apical system: Central, four gonopores; hydropores scattered over the madreporite. Petals: Well-developed, tapering distally, pore zones relatively narrow. Oral side: Short, simple food grooves; peristome central, position of periproct variable from close to the peristome to close to the posterior edge; plates naked or with spines. Remarks: Mortensen (1948a,b) emphasized that the main differentiation among these species is whether the periproct is naked or set with spines. However, he added that, exceptionally, there may be a few spines on the periproctal plates in species usually with a naked periproct or in those with fully spined plates. However, one finds predominantly naked tests without any remaining periproctal plates. Peronella affinis (MClelland, 1840) Petaloid area less than half the test length, posterior side slightly restricted; periproctal plates naked; length up to 64 mm. Distribution: Indian Ocean. Peronella analis de Meijere, 1903 Seems to be not longer than 17 to 18 mm, slightly ovate, conically raised, petals more than half the test length; periproct elongated and naked. Distribution: Malayan region; 22 to 141 m. Peronella hinemoae Mortensen, 1921 Test: Thin, outline nearly circular or polygonal, distal part flat, center gently raised, apex rounded; maximum length approximately 60 mm.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

 83

Apical system: Slightly anterior of center. Petals: Short and fairly broad, pore zones slightly curved and distally turning outward; petaloid area less than half the test length. Oral side: Slightly and regularly concave, peristome subcentral and rounded pentagonal, periproct round, positioned halfway between peristome and posterior edge, periproctal plates naked. Food grooves: Indistinct. Tuberculation: The associated spines are fine. Color: Test and spines are orange- to reddish-brown, the well cleaned test is whitish. Distribution: The species is recorded from off southeast Australia, the Tasman Sea and the Norfolk Island to New Zealand and the Kermadec Islands. Bathymetric range 17 to 260 m (Fig. 3.65). Peronella japonica Mortensen, 1948 Test: Rounded polygonal or slightly oval, faintly raised toward the apical system, margin thick, no ring-shaped depression; length up to 70 mm. Apical system: See Fig. 3.66. Petals: Long, almost closed distally, densely set pore pairs. Oral side: Flat or slightly concave, food grooves distinct; periproct closer to the posterior end than to the peristome. Color: In life, the spines are dark reddish to reddishbrown. The bare, cleaned test is whitish. Distribution: P. japonica is restricted to Japanese waters from central Japan to southern Kyushu, living at depths of 3 to 50 m. Peronella keiensis Mortensen, 1948 Test: In outline, almost circular or rounded pentagonal, margin thin, center distinctly inflated; length up to 49 mm. Apical system: Central, numerous hydropores scattered over the anterior and right part of the madreporite. Petals: Paired ones closed distally, anterior petal open and distinctly longer than paired ones. Oral side: Flat, slightly sunken toward the peristome; periproct positioned near the posterior end, round or transverse oval, with naked plates. Food grooves: Short and not well marked. Tuberculation: Aborally dense and fine, small primaries in a very dense coat of miliary tubercles, orally less dense. Color: Test and spines are a light yellowish-brown. Occurrence: Kei Islands, Indonesia; 245 m. Peronella lesueuri (L. Agassiz, 1841) Test: Large and thin with faintly thickened margin; outline oval, round or polygonal; aboral side slightly raised toward the center; maximum length 160 mm. Petals: Narrow and long; more or less closed distally.

84 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

B

A

C D

E

Fig. 3.65: Peronella hinemoae. (A) Aboral side of test, length: 41 mm, off southeast New Zealand. (B) Oral side. (C) Side view. (D) Aboral side with spines, length of test 21 mm, off southeast New Zealand. (E) Oral side with spines.

Oral side: Periproct transversely elongated and near the posterior margin; ambulacral furrows very short and distinct. Color: The live sea urchin is bright orange, turning green after death. The bare test is whitish. Distribution: This species is found from Malaysia and the Philippines to Australia and New Guinea. Biology: P. lesueuri lives not buried, but on the surface, from the intertidal zone to a depth of 70 m (Fig. 3.67).

Peronella macroproctes Koehler, 1922 Test: Regularly oval, sometimes slightly truncated posteriorly, edge slightly thickened; aboral side elevated; length up to 37 mm. Apical system: Hydropores few and inconspicuous. Petals: Broad, distally not quite closed; petaloid area slightly more than half the test length. Oral side: Slightly concave; periproct almost as large as peristome, plates set with spines; distant from the

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

 85

B

A

C

D

E

F

Fig. 3.66: (A–C) Peronella japonica. (A) Aboral side, test length 82 mm, Fukui Prefecture, Japan Sea, central Japan. The petals are long and distally closed. (B) Oral side: the periproct is closer to the posterior margin than to the peristome. In this specimen, a covering of skin remains. The cleaned test is usually whitish. (C) Aboral side with spines, test length 75 mm; Fukui Prefecture, Japan Sea, central Japan. The miliary spines are very densely distributed, the primaries are less dense and much longer than the miliaries. (D–F) Peronella keiensis. (D) Aboral side of test, length 46 mm; Kei Island, Indonesia. ZMUC ECH 191. The paired petals are broad, narrowing distally, the anterior petal being distinctly longer. (E) Oral side: the periproct is small, positioned much closer to the margin than to the peristome. (F) Side view: the test is conically raised.

86 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

B

A

C

D

E

F

Fig. 3.67: (A,B) Peronella lesueuri. (A) Aboral side of test, length 114 mm; Western Australia. The columns of the interambulacral plates taper strongly toward the margin. (B) Oral side: the food grooves are short and distinct. The periproct lies close to the posterior margin (the very edge is damaged). (C–F) Species of Peronella, part of petals: (C) P. macroproctes. (D) P. oblonga. (E) P. rubra. (F) P. keiensis. All scale bars are 2 mm; after Mortensen (1948a,b).

posterior edge, but less than halfway between peristome and rear. Tuberculation: Exceedingly fine, primaries inconspicuous in the dense mass of miliaries. Distribution: Indo-West Pacific from Sri Lanka to the Philippines, the Malayan region and northeast Australia; bathymetric range 5 to 105 m.

Occurrence: Known only from the Mergui Archipelago, Myanmar.

Peronella merguiensis Koehler, 1922 Test in length up to 42 mm; longer than wide and rounded pentagonal; petals short and narrow, lanceolate and pointed; ambulacral furrows short and distinct; periproctal plates set with spines.

Peronella oblonga Mortensen, 1948 Length of holotype 23 mm; margin thickened, but less than in P. minuta; petals rather broad and closed distally; periproct near the posterior margin, furrows indistinct. Occurrence: South India.

Peronella minuta (de Meijere, 1904) Test thick with a high edge, up to 21 mm in length; oral side distinctly concave; a few madreporic pores; petals broad adapically, nearly closed distally. Distribution: Sulu Sea; depth 13 to 35 m (Fig. 3.68).

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

A

BC

C

D

 87

Fig. 3.68: (A,B) Peronella macroproctes. (A) Aboral side of a young specimen, test length 11.2 mm; south coast of Sri Lanka. (B) Oral side: the periproct is almost as large as the peristome. Photo in Koehler (1922). (C,D) Peronella orbicularis. (C) Aboral side of a young specimen, test length 13 mm; Cape Range, Western Australia. The petals are broad and short. (D) Oral side: the food grooves are short and rather indistinct.

Peronella orbicularis (Leske, 1778) Test: Thick with inflated margin; raised toward the apical disc; outline round or slightly oval; oral side concave; maximum length 22 mm. Petals: Broad at the apical system, outward narrowing, distal end almost closed.

Oral side: Faintly concave, periproct close to the posterior margin; food grooves indistinct. Color: In life, the test is light brown, the bare test is white. Distribution: This species is known from the Gulf of Iran, the Malayan region, the Philippines and Taiwan to Australia

88 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

from Kalbarri, Western Australia, around the northern coast to the Capricorn Islands, Queensland; bathymetric range from the intertidal zone to a depth of 70 m. Biology: Peronella orbicularis lives on shell grit or coarse sand bottoms. Dead tests are commonly found at low tide. Peronella pellucida Döderlein, 1885 Test: Extremely thin, outline nearly circular or polygonal, distal part flat, center inflated, apex rounded; test length may reach 50 mm. Apical system: Slightly anterior of center. Petals: Broadest near the base, narrowing toward the open distal end; petaloid area less than half the test length. Oral side: Slightly concave, posterior interambulacrum faintly inflated between peristome and periproct; peristome rounded pentagonal; periproct smaller, positioned about halfway between mouth and posterior margin, plates naked; food grooves indistinct. Color: Spines and skin whitish to yellowish-light brown (Fig. 3.69). Peronella peronii (L. Agassiz, 1841) Test: Thick and strong with slightly inflated margin; slightly raised toward the apical system, oral side concave; outline round to slightly oval; length may reach 42 mm. Apical system: Gonopores lie outside the apical disc in the interambulacra. Petals: Slightly inflated, pore zones gently curved and in the paired ones closed distally, in anterior one half open. Oral side: Shallowly concave, periproct one-third of the distance from the margin to the peristome; food grooves short and inconspicuous. Color: The living sea urchin is light brown or whitish, the bare test is white. Distribution: This species is confined to southern Australia from New South Wales to Rottnest Island, Western Australia; at depths of 3 to 360 m. Peronella rubra Döderlein, 1885 Test: Circular, elliptical, or polygonal, edge thickened, center moderately inflated; length up to 60 mm. Apical system: With numerous hydropores scattered on the right and anterior part of the madreporite. Petals: Moderately long, broad at the base, narrowing to the nearly closed distal end; petaloid area small, about half the test length or less. Oral side: Slightly concave; periproct about halfway from the posterior end to the peristome, irregular in shape, the plates carrying tubercles; food grooves distinct.

Tuberculation: Fine and dense; aboral primary spines faintly thickened at the tip and only slightly longer than the miliaries. Color: The test and spines are dark red fading to pink, the well cleaned test is nearly white. Distribution: The species is known in Japanese waters from northern Honshu to southern Kyushu. It is also recorded from the Java Sea; bathymetric range: 5 to 60 m (Fig. 3.70). Peronella rullandi Koehler, 1922 Only juveniles were found; length of holotype 18 mm; petals short, pore zones rather broad; periproct midway between peristome and rear. Locality: Probably Indian Ocean (Koehler noted only “Rulland”). Peronella strigata A. Agassiz & H.L. Clark, 1907 Petaloid area less than half the test length; elongated naked periproct; light brown to purple, plate sutures darker. Occurrence: Hawaiian Islands; 250 m. Peronella tuberculata Mortensen 1918 Test: Thin and fragile; outline round to decagonal, upper side weakly convex, oral side slightly concave; margin not inflated but thickened rising gently toward the apex; length may reach 40 mm. Petals: Reach over 60% of the aboral surface, distally almost closed. Oral side: Periproct transversely oval; lies one-third along the distance from the rear to the peristome. Distribution: Restricted to Western Australia, living in the ebb zone and to a depth of 15 m.

3.2.2.6 Family Rotulidae Gray, 1855 The species of this family are restricted to the tropical west coast of Africa. They live, often in very large numbers, shallowly buried in lagoons and estuaries with very fine sediment. In this extreme habitat, they evolved quite unique adaptations. The flat test has digitations alternating with deep indentations around the posterior half of the margin. Additionally, in Rotula, more or less closed holes (lunules) have developed on the anterior part of the test. Organic particles are caught by the tube feet and laid into the food grooves, which reach into the distal ends of the “fingers”. Here, the grains are incorporated into a mucous string and transported to the central mouth. The phylogenetic placement of the rotulids is still the subject of discussion. Two extant genera, each with one species, are included in this family.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

 89

B

A

C D

E

F

Fig. 3.69: (A–D) Peronella pellucida. (A) Aboral side of test, length 38 mm, Mactan, Philippines. The petals are small. (B) Oral side: the test is very thin and fragile. (C) Side view. (D) The three anterior petals. (E,F) Peronella peronii. (E) Aboral side of test; length 35 mm, Esperance, Western Australia. The gonopores are positioned in the interambulacra. (F) Oral side: the food grooves are very short and indistinct.

90 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

A

B

C

D

Fig. 3.70: (A,B) Peronella rubra. (A) Aboral side of test, length 55 mm, Noto Peninsula, Japan. (B) Oral side. The food grooves are distinct. Photos courtesy of H. van Noordenburg. (C,D) Peronella tuberculata. (C) Aboral side of test; length 31 mm; Dirk Hartog Island, Western Australia. All five petals are nearly closed. (D) Oral side: the food grooves are slightly indented.

Genus Heliophora L. Agassiz, 1840 The genus is monospecific. Heliophora orbiculus Linné, 1758 Test: Fragile, flat, more or less strongly domed around the apical system; circumference round, posterior part with 11 deep indentations, the plate columns form more or less

slender finger-like structures; maximum length approximately 70 mm. Apical system: Knob-like, with four gonopores. Petals: Short and open, with almost parallel pore zones; anterior one longest. Oral side: Flat or faintly concave; food grooves bifurcating, not reaching the anterior margin but expanding onto



the digitations posteriorly; periproct between peristome and posterior margin. Spines: Primaries slightly club-shaped, miliaries ending in a crown. Color: In life, the animal is brownish to grayish, turning brightly green after death. The denuded and cleaned test is whitish. Distribution: This species is found along the coast of West Africa from Mauretania to Angola in the intertidal zone. Remarks: The specimens can vary from having only shallow indentations around the posterior side to having digits around almost the entire margin. Between these extremes, many intermediates occur (Fig. 3.71). Genus Rotula Schumacher, 1817 The genus is monospecific. Rotula deciesdigitata (Leske, 1778) Test: Thin and flat, center raised, highest point anterior to apical system about half the distance to the anterior margin, sloping evenly toward the posterior; anterior side rounded with two to six more or less open lunules, posterior side with open indentations at each suture, deepest in interambulacra; length may reach 77 mm. Apical system: Star-shaped and slightly elevated; four gonopores. Petals: Long and distally open with broad, nearly parallel pore zones. Oral side: Flat to concave; food grooves expanding to distal end of digitations; poorly branched; periproct close to the posterior indentation. Spines: On the aboral side short, very dense and ending in a knob, longer on the oral side; miliary spines end in a crown. Color: In life, the echinoid is greenish gray to brownish, the bare test is whitish. Distribution: R. deciesdigitata is found along the coast of West Africa from Mauretania to Angola in the intertidal zone. Remarks: The closed lunules are formed by resorption, the posterior notches by marginal indentation.

Infraorder Scutelliformes Haeckel, 1896 Characteristics: The aboral interambulacra ends in two plates at the apical system. The pore pairs lead along the ambulacral sutures. The miliary spines end in a fleshy sac. Unique in echinoids, scutelliform juveniles are capable of storing sand grains in their intestinal diverticula, which

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

 91

serve as weight belts in turbulent water. Mooi & Chen (1996) interpreted this as a result of phylogenetic history, not of adoptive factors. The species have large, flat tests; several of them have closed lunules or open marginal notches in some or all ambulacra, and eventually an anal lunule in the posterior interambulacrum. On the aboral side, the petals are well-developed. The scutelliform echinoids belong to the most specialized sand dollars, having evolved clearly differentiated kinds of spines and tube feet (see also Fig. 3.72) On the aboral side, they are equipped with very short and very densely distributed primary spines ending in a distinct, more or less inflated knob. The tips of the miliary spines are balloon-like, fleshy sacs, which fit flexibly into the interspaces of the primaries. Principally all echinoids that live more or less buried in fine, sandy sediment need to evolve elaborate strategies to keep the test surface free of clogged particles. The enormous number of external appendages and the epithelium obtain their nutrients dissolved from the surrounding seawater. Therefore, a current of fresh water flowing over the surface has to function undisturbed. Each miliary spine bears two bands of fine hairs, the cilia, which maintain this current immediately above the epidermis. The tips of the primary and miliary spines form a dense canopy which on one hand prevent material to fall onto the surface whereas on the other hand they push in a coordinated movement a thin layer of sand grains backward over the aboral test (Mooi 1997) (Fig. 3.73). On the oral side, diverse spines and tube feet are developed in well-defined fields, each differentiated for a special function: Fringe spines around the lunules and along the outer margin, where long sensory podia also occur. Scattered spines in the pressure drainage channels and their tributaries, no tube feet. Long locomotory spines, areas with no tube feet. Small, bent spines protecting the short barrel-tipped tube feet distributed between them. Long barrel-tipped tube feet are set along the borders of the different fields. Enormous numbers of these short and long tube feet with barrel-shaped sucking discs selectively gather organic particles from the underlying sediment (Fig. 3.74). The grains are manipulated from one podia to the next, similar to a bucket-chain, to the food grooves where they are incorporated into a mucous string being moved to the central mouth by very small podia (Telford et al. 1985).

92 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

A

C

E

B

D

F

3.2.2 Order Clypeasteroida L. Agassiz, 1835 



 93

A

B

E

With these special features on the aboral and the oral sides, the scutellines are adopted to their special habitat. They live more or less buried in clean, fine, sandy sediments, mostly in shallow water with considerable wave motion. They are not found in or on silty or stony substrata. Literature: Grant & Hertlein 1938; H.L. Clark 1948; Durham 1955; Timko 1976; Telford 1981, 1988; Telford & Harold 1982; Ellers & Telford 1984; Wang 1984; Telford et al. 1985; Mooi & Telford 1986; Mooi 1986a,b,c 1990, 1997; Mooi & Harold 1994; Mooi & Chen 1996.

Fig. 3.72: Characteristics of scutelliforms: (A) The lunules, holes between the aboral and oral sides, are developed in ambulacra and/or interambulacra. In some species, additional depressed pressure drainage channels with more or less conspicuous accessory tributaries lead into the lunules and to the edge. These features help the flat discs to keep their stability in heavy waves. Distinct food grooves lead to the mouth. They bifurcate close to the peristome and finely subdivide distally being evolved for the transport of food. The sizes of the tubercles vary according to the differentiated spines. (B) Aboral spines of Mellita tenuis: the primary spines end in a skeleton knob. In contrast, the miliary spines between them are equipped with a flexible, fleshy sac. The shaft is set with two ciliary bands. These cilia, fine hairs, generate currents to provide the epiderm of podia and spines with nutrients and to sweep finer particles off the test. Light gray, epiderm in general; moderate gray, skin covering spines; dark gray, skeleton of test and primary spines; black, skeleton rod of miliary spines; white, fleshy sac of miliary spines. Modified after Mooi (1986c).

The suborder consists of six extant families. The placement of some genera is still the subject of discussion, and some questions remain open. Astriclypeidae Stefanini, 1912. Two or four genera are included, reported from the Indo-Pacific. Dendrasteridae Lambert, 1889. A single genus with three species is distributed in the northern Pacific. Echinarachniidae Lambert, 1914. One extant genus is known with one species, restricted to cold and temperate water in the northern hemisphere.

◂ Fig. 3.71: (A–C) Heliophora orbiculus. (A) Aboral side of a young specimen, test length 17 mm; Luanda, Angola; ZMH. There are already shallow indentations at the sutures. (B) Aboral side of test, length 35 mm; Cape Verde Islands. The long “fingers” are very fragile. When they break, they can be regenerated. (C) Oral side of an adult animal, test length 56 mm, Mauretania. The strongly recessed food grooves do not reach the anterior margin but expand onto the posterior digitations. (D–F) Rotula deciesdigitata. (D) Growth series, aboral sides, test length (from left to right) 7, 16, and 24 mm, Lagos, Nigeria (ZMH). First, the three deep indentations in the posterior interambulacra evolve; later, the more shallow notches in the other sutures appear. The anterior holes, called lunules, variably two to six, are formed by resorption of the test. (E) Aboral side of test, partly with spines, length 55 mm, width 53 mm; Lagos, Nigeria. (F) Oral side. The anterior part of the test shows closed lunules, the posterior side is deeply indented. Often, the fragile digitations break, but can be regenerated.

94 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

Fig. 3.73: Mellita tenuis; Florida, USA. Only a very thin layer of sand covers the “keyhole” urchins. The outlines of the animals and the depressions of the lunules are visible. Photo courtesy of H. Moosleitner.

A

B

E

F

Mellitidae Stefanini, 1911. This family comprises four genera. They are distributed along the tropical and subtropical coasts of the Americas on the Atlantic and Pacific sides. Scutellidae Gray, 1825. Beside many fossil genera, a single extant genus, Scaphechinus, is placed in this family,

Fig. 3.74: Oral side with spines. (A) Specialized spines and tube feet are arranged in well-defined fields. (B) ­Halfdiagrammatic transect through the oral side of a mellitid sea urchin: the test is set with diverse spines and tube feet, differentiated for special functions. Modified after Telford et al. 1985.

although with weakly supported relationship (Mooi, personal communication). Taiwanasteridae Wang, 1984 (Sinaechinocyamus). Additionally, there is the genus Marginoproctus Budin, 1980, which has to be kept incertae sedis because

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

 95

A

B

C

D

E

F

Fig. 3.75: (A, B) The lunules in the scutelline families evolved independently, being constructed in a different way: in the Astriclypeidae, the ambulacral plates in the lunules are cross-linked (Echinodiscus truncatus). (B) In the Mellitidae, the plates are festooned (Mellita isometra). All scale bars are 10 mm (redrawn and modified after Mooi 1989). (C–F) Astriclypeus manni; test length 129 mm; Fukui Prefecture, Japan Sea, central Japan. (C) Aboral side: five lunules are developed in the ambulacra, there is no interambulacral anal lunule. (D) Oral side: the main branches of the food grooves pass closely around the lunules. (E, F) Test with spines; length 122 mm; Fukui Prefecture, central Japan. (E) Aboral side: the spination is dense like short fur. (F) Oral side: there are sharply differentiated fields of shorter spines around the food grooves, and longer ones serving for locomotion.

Genus Astriclypeus Verrill, 1867 The genus is monospecific.

Apical system: Five gonopores. Petals: Short, petaloid area less than half of test length. Oral side: Perfectly flat, food grooves bifurcating, the main branches passing close to lunules, strongly developed pressure drainage channel toward proximal end of lunule; varying number of side branches; periproct on oral side, halfway from margin to peristome. Spines: Primaries more or less club-shaped, miliaries with distal fleshy sac; differentiated on oral side. Color: In life, the test is generally brownish, the bare test is whitish. Distribution: The species is reported from central Japan at the Pacific and the Japan Sea to southern China and Vietnam at depths of 6 to 35 m (Fig. 3.75).

Astriclypeus manni Verrill, 1867 Test: Large, strong, and heavy; center domed, flat toward the margin; circumference rounded to pentagonal, posterior side truncated; five narrow lunules in the ambulacra, no anal lunule; maximum length more than 130 mm.

Genus Echinodiscus Leske, 1778 Jansen & Mooi (2011) made an analysis about the species of Echinodiscus, comparing hundreds of specimens from around the Indo-West Pacific and Japan. They found out that there are several clearly different morphotypes related

it cannot be placed in a certain scutelliform family (Kroh & Mooi 2010).

3.2.2.7 Family Astriclypeidae Stefanini, 1911 Two well-differentiated genera are included in this family: Astriclypeus with five lunules and Echinodiscus with two, which may be open or closed. Stara & Sanciu (2014) established two further genera, Paraamphiope and Sculpsitechinus, which were based mainly on small differences in plate architecture and shape of the visceral hollow.

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 3.2 Superorder Neognathostomata A.B. Smith, 1981

to well-limited geographic regions from East Africa, the Red Sea, and the Gulf of Iran to Japan, Indonesia, New Caledonia, and northwest Australia. In the course of their comprehensive studies, they discovered quite unexpectedly unusual pedicellariae associated with certain morphotypes. Further studies will reveal if they turn out to become new species. In 2014, Stara & Sanciu presented a study “Analysis of some astriclypeids” on fossil taxa and extant specimens from Thailand and Indonesia, establishing two new genera with several species, Sculpsitechinus and Paraamphiope (P. raimondii, known only a holotype). They were mainly based on different plate patterns and the shape of the visceral hollow. All resemble Echinodiscus in having two closed or open lunules posteriorly, with the range of variation being very restricted. Test: Rather solid or thin and fragile; of medium to large size; very flat; slightly raised with the highest point anterior to the apical system; outline rounded trapezoid or posteriorly truncated, two posterior lunules or deeply indented notches. Apical system: Four gonopores. Petals: Moderately short, more or less broad. Oral side: Flat with pressure drainage channels in the ambulacra passing to the anterior margin and to the posterior lunules or notches; bifurcating food grooves with distal side branches. Periproct: Between margin and peristome. Spines: Aborally very short and club-shaped, longer on oral side with well-defined fields for locomotory spines (not in Echinodiscus tenuissimus). Color: In life, the tests are brownish to dark purplish; the bare, cleaned tests are whitish. Habitat: The species of Echinodiscus live in shallow, sheltered water half-buried in sandy sediments, from the intertidal zone to a depth of 50 m. The genus consists of four species with several morphotypes. Additionally, Stara & Sanciu (2014) described a species in a very limited locality at the Andaman Sea in Thailand. Echinodiscus auritus Leske, 1778 Test: Posteriorly broader than anteriorly; anterior margin rounded or sinuous, posterior margin truncated and with two deep, open notches; length may reach 183 mm. Petals: About half the width of the test. Distribution: The species is known from the Gulf of Iran, the Red Sea, and East Africa to Thailand, the Philippines, New Guinea, New Caledonia, and northwest Australia at depths between 0 and 5 m. Remarks: Jansen & Mooi made the interesting discovery that the morphotype from west Australia is set with poison gland–bearing globiferous pedicellariae. This type

of pedicellariae is not found on any other species within the order Clypeasteroida. The tridentate pedicellariae of West Australian morphotypes are many times larger than in E. auritus specimens from other regions. Stara & Sanciu (2014) call this species Sculpsitechinus auritus (Leske, 1778). Echinodiscus bisperforatus Leske, 1778 Test: Outline rounded or broadly trapezoid; in the posterior ambulacra, there are two elongated lunules, closed and more or less slit-like; length up to 80 mm. Petals: Short and broadly rounded. Oral side: Flat; food grooves bifurcating, leading closely along the lunules and the strongly developed pressure drainage channels; sharply limited fields of smaller and larger tubercles in ambulacra and interambulacra. Distribution: The species is known from the Gulf of Iran and the Arabian Sea and from the Red Sea southward to Mosselbay in South Africa in shallow water (Fig. 3.76). Echinodiscus tenuissimus (L. Agassiz & Desor, 1847) Test: Thin and fragile, two round or more or less elongated lunules in the posterior ambulacra. Oral side: No sharply limited fields of different-sized tubercles in ambulacra and interambulacra. Distribution: Found from south Japan over the Philippines to New Caledonia in the intertidal zone to a depth of approximately 20 m. Remarks: Stara & Sanciu (2014) named this species Sculpsitechinus tenuissimus (L. Agassiz, 1841). Echinodiscus truncatus L. Agassiz, 1841 Test: Rounded trapezoid with shallow indentations in the anterior ambulacra, associated with strongly developed pressure drainage channels. Lunules: Distinctly shorter and broader than in other species. Distribution: This morphoform occurs from Myanmar over Thailand and the Malayan Archipelago to New Caledonia from the intertidal zone to a depth of 50 m. Remarks: Jansen & Mooi (2011) detected in E. truncatus a very unusual form of ophicephalous pedicellariae, the teeth of which resemble a “meat pounder”, positioned on the inside of the distal part of the valve.

3.2.2.8 Family Dendrasteridae Lambert, 1889 Only one extant genus is included in this family. Genus Dendraster L. Agassiz, 1847 Test: Strong, lowly domed with thin margin, highest point anterior to apical system; outline almost round or trapezoid with the posterior part of test broader.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 



A

B

C

D

E

F

 97

Fig. 3.76: (A, B) Echinodiscus auritus. Test length 74 mm, width 78 mm; Khor Fakkan, Gulf of Oman. (A) Aboral side: the characteristic open notches are deeply indented. (B) Oral side: the main branches of the food grooves pass closely along the notches and the pressure drainage channels in the three anterior ambulacra. There are numerous side branches extending over the surface of the test. (C) Echinodiscus bisperforatus; test length 94 mm, width 104 mm; Pangani, Tanzania. (D, E) Echinodiscus tenuissimus; length 77 mm; Amami Island, Kyushu, Japan. (D) Aboral side and (E) oral side: the lunules are elongated, but not as narrow as in E. bisperforatus. (F) Echinodiscus truncatus; test length 55 mm, width 63 mm; Bali, Indonesia.

Apical system: Strongly eccentric in being distinctly posterior to center, four gonopores. Petals: Well-developed, posterior pair shorter than the three anterior ones and more or less extending to the sides. Oral side: Quite flat, food grooves very conspicuous, bifurcating and branching distally, in some species extending onto the aboral side; absent on anterior part of test; peristome subcentral, periproct on the oral side close to the posterior end. Spines: On the aboral side, short and distally thickened, on the oral side longer and not thickened. Biology: Dendraster is capable of capturing actively swimming prey. In strong water currents, the echinoid straightens itself, the anterior part of the test is inserted into the sediment, and the posterior part is raised above the ­substrate. Often, large numbers of individuals are found “standing” side by side in the heavy flow. The oral side is orientated against the water current. When potential food such as the larvae of crabs or copepods come into contact with the surface, the spines close like a trap, forming a cone with a diameter of 1 to 2 mm (trapping response). Within

a few seconds, large bidentate pedicellariae extend with open jaws and shut vigorously; only rarely do the prey in the trap escape. After snapping on the swag several times, the pedicellariae release it, spines and tube feet move it toward the food grooves and around the margin, which is densely set with podia. Nonmobile particles and items less than 50 µm in diameter are grasped by tube feet and manipulated into the grooves. Then, the catch is transported to the food grooves, eventually around the margin, and incorporated into a mucous string and moved toward the mouth. Nonmobile food particles are caught by the suckered tube feet and manipulated to the grooves. On the way to the mouth, smaller particles are enveloped in the mucous string, larger ones are moved by a coordinated rowing motion of the ambulacral tube feet. The average time from prey capture to ingestion is approximately 15 min, motile prey longer than nonmotile, then the teeth masticate the prey for another 15 min. The particles are selected and inappropriate ones are rejected, moved toward the edges of the test or dropped down. The “last control” occurs at the Y junction near the mouth, where

98 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

A D

B

C

unacceptable material will be pushed away by tube feet. In less strong water currents, Dendraster excentricus feeds by picking up the particles from below the test like other sand dollars (Timko 1976). Individuals of Dendraster living in more turbulent ­higher-energy water tend to have more eccentric apical discs and grow to a larger size (Stanton et al. 1979). The genus contains three extant species, living along the west coast of North America mainly in the littoral. D. excentricus is very common from Alaska to southern Baja California, where the two other species live in well limited areas. That means the species live more or less sympatrically. They are not known in the Gulf of California (Mooi 1997) (Fig. 3.77). Dendraster excentricus (Eschscholtz, 1831) Test: Length may reach approximately 80 mm. Color: In life, the test is brownish, grayish or light purplish, the bare test is grayish-white. Distribution: Known at the west coast of America from southern Alaska to south Baja California, in the ebb zone to a depth of 90 m. Dendraster terminalis Grant & Hertlein, 1938 (synonym: D. laevis H.L. Clark, 1948) Test: Slightly domed or subconically raised, highest point in front of the center, margin thin, length may reach 50 to 60 mm. Apical disc: Slightly posterior to center. Petals: Open distally, the anterior the longest, the posterior pair shortest; petaloid area small.

Fig. 3.77: (A–C) Food grooves of the three extant species of Dendraster in comparison: (A) Dendraster excentricus: the apical system is strongly eccentric; the aboral spines are distinctly club-shaped and the food grooves reach a little onto the aboral side.(B) Dendraster terminalis: the apical system is small and the least eccentric, the aboral spines are slightly thickened and the food grooves do not extend onto the aboral side. (C) Dendraster vizcainoensis: the apical system is slightly eccentric, the aboral spines are thickened with pointed tips and the food grooves reach onto the aboral side as far as into the petals. Tests shown in oral (left) and aboral view (right), Scales of tests 10 mm. Modified after Mooi (1997). (D) D. excentricus is a suspension feeder, able to catch actively swimming prey, which is swept by the water current against its oral side. Modified after Timko (1976).

Oral side: Flat; periproct just below the posterior margin; food grooves not extending onto the aboral side. Spines: Slightly thickened on the aboral side. Distribution: This species is known from southern California and the Channel Islands, USA, and from northern Baja California and Guadeloupe Islands, Mexico; in shallow water. It overlaps a little with D. vizcainoensis toward the south. Dendraster vizcainoensis Grant & Hertlein, 1938 Test: Petaloid area distinctly raised, distal parts of test flat, highest point slightly posterior to center; maximum length 60 mm; fossil specimens of this species may reach 90 mm. Petals: Approximately the same length, distally open. Oral side: Flat; periproct close to the posterior margin; food grooves strongly developed, extending far onto the aboral surface into the petals. Spines: Aborally distinctly thickened but pointed toward the tip. Distribution: This species occurs at the Pacific coast of central Baja California, northward slightly overlapping with D. terminalis.

3.2.2.9 Family Echinarachniidae Lambert, 1914 Several fossil genera were placed in this family. However, it should be restricted to forms with trifurcating food grooves (Mooi 2015, personal communication). Genus Echinarachnius Gray, 1825 The genus is monospecific.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

A

 99

B

F

C

D

E

Fig. 3.78: (A–C) Dendraster excentricus. Test length 84 mm; Bodega Bay, CA, USA. (A) Aboral side. (B) Oral side: the strongly developed food grooves extend onto the aboral side around the posterior, they are lacking on the anterior part of the test. (C) Length with spines 45 mm; San Francisco, CA, USA. Aboral side of test. The food grooves extend around the posterior edge onto the upper side. The spines are clubshaped. (D, E) Dendraster terminalis. Test length 38 mm, Baja California, Mexico. ZMUC ECH 199 as Dendraster laevis. (D) Aboral side: the petaloid area is remarkably small. (E) Oral side.

Echinarachnius parma (Lamarck, 1816) Test: Solid and flat, upper side subconically raised or convex, margin thin; outline round, slightly indented at  the sutures, small posterior notch; test length up to 50 mm. Apical system: Central, four gonopores. Petals: Broad and distally widely open. Oral side: Flat or slightly concave; peristome central; food grooves forming five trunks with few distal side branches; periproct in a small notch at the posterior margin. Spines: Aborally short, longer on the oral side and not differentiated into fields.

Color: In life, it is generally dark brown, the bare test is yellowish. Distribution: This species is known from the northern Pacific from Puget Sound to Alaska, the Kamchatka Peninsula and north Japan; in the northwest Atlantic from Labrador, Canada to Massachusetts, USA. It lives in the littoral zone and down to a depth of approximately 1,500 m (Fig. 3.79). Biology: This species lives locally in high densities (up to 180 individuals per square meter) on moderately sorted fine to medium sand floors. After current activity, these sand dollars are second in importance for bioturbation by reworking surficial sediments (Stanley & James 1971).

100 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

A

B

C

D

Fig. 3.79: Echinarachnius parma. (A, B) Test length 47 mm; Vancouver Island, Canada. (A) Aboral side: the five petals are open and equal in size. (B) Oral side: the food grooves form five broad trunks, which are branched distally. The central branch reaches the margin. The periproct lies marginally in a distinct notch. (C, D) Test with spines, length 59 mm; Vancouver Island, Canada. (C) Aboral side: the test is densely covered by short spines. The posterior notch with the periproct is distinct. (D) Oral side: the spines are longer than on the upper side serving for locomotion.

3.2.2.10 Family Mellitidae Stefanini, 1911 This family contains the typical “keyhole” urchins, the sand dollars with the most specialized lifestyle of echinoids. Their flat tests are provided with marginal notches and/or closed lunules, which serve mainly for keeping stability in wave motions (their lifestyle, see page 93/94).

In juvenile specimens of Mellita and Encope, the ambulacral lunules are formed initially as marginal indentations, which become deeper and eventually fully closed as the test grows. In contrast, the marginal ambulacral lunules of Leodia are built by resorption of the test. First, there is only a small hole between the ambulacral sutures, which grows and, later in ontogeny, the lunule becomes slit-like.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

A

B

C

D

 101

Fig. 3.80: (A, B) Mellita tenuis, juvenile specimen; test length 11.5 mm; Sanibel Island, West Florida, USA. (A) Aboral side: the ambulacral lunules are still open notches, the anterior pair being much smaller than the posterior ones. (B) Oral side: the main branches of the food grooves are already developed, and, relative to the size of test, rather broad. (C) Mellitella stokesii, juvenile specimen; test length 12.5 mm; Peru. ZMH. Aboral side: the ambulacral notches are deep indentations. The dark patches indicate internal cavities in which sand grains are stored forming a weight belt to make the test heavier and more stable in heavy water. (D) Leodia sexiesperforata; juvenile specimen. ZMH. Test length 12 mm, Puerto Colombia, Colombia. Aboral side: the ambulacral lunules are built by resorption of the test, not by marginal indentations. The holes are present already in very small juveniles.

The family consists of four genera, distributed from the tropics to temperate water on both sides of the Americas living in the littoral: Encope L. Agassiz, 1840. Five gonopores, five lunules or notches in the ambulacra, and an additional anal lunule; Atlantic and Pacific. Leodia Gray, 1852. Four gonopores, five ambulacral lunules, and an additional anal lunule; Atlantic.

Mellita L. Agassiz, 1841. Four gonopores, four ambulacral lunules, and an additional anal lunule; Atlantic and Pacific. Mellitella Duncan, 1889. Five gonopores, five deep notches in the ambulacra, sometimes closed, and an additional anal lunule; Pacific (Fig. 3.80). Remarks: Coppard et al. (2013) seemed to reveal, by molecular analyses, that there are 11 species. They include

102 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

six cryptic species. The morphotypes are not compatible to the molecular analysis. Literature: Clark 1946; Clark 1948; Phelan 1972; Ebert & Dexter 1975; Telford & Mooi 1986; Harold & Telford 1990; Hendler et al. 1995; Coppard et al. 2013, Coppard, 2016. Genus Encope L. Agassiz, 1840 Test: Flat, of medium to large size, aboral side may be raised or flat; five ambulacral lunules or open notches, anal lunule more or less between the posterior petals. Apical system: Five gonopores. Petals: Of approximately similar shape. Oral side: Flat, conspicuous food grooves in ambulacra diverging on post-basicoronal plates (in contrast to Leodia); peristome subcentral, periproct close to the peristome in front of the anal lunule. Spines: On the aboral side, very short and thickened, orally longer and more or less differentiated. Seven species are distinguished with several subspecies. Atlantic side of the Americas: Encope aberrans Martens, 1867. Very shallow marginal notches. Encope emarginata Leske, 1778. Anal lunule very long, marginal lunules open or closed. Encope michelini L. Agassiz, 1841. Marginal notches deeply indented, anal lunule strongly elongated. Pacific side of the Americas: Encope grandis L. Agassiz, 1841. Thick, greatly enlarged anal lunule, marginal notches widely open; endemic to the northern part of the Gulf of California. Encope laevis H.L. Clark, 1948. Upper spines form a smooth “pavement”. Encope micropora L. Agassiz, 1841. Anal lunule small and rather marginal. Encope perspectiva L. Agassiz, 1841. Large anal lunule, aboral spines with broad tips that form a sort of “second surface”. Encope wetmorei A.H. Clark, 1946. Upper spines inverted cone-shaped, each with a concave surface. The following descriptions are listed alphabetically: Encope aberrans Martens, 1867 Test: Strong, subcircular, or longer than wide; length up to 150 mm; highest point between apical disc and anal lunule, sometimes formed like a veritable hump. Petals: Very long, posterior pair longest and curved around the anal lunule. Lunules: Posterior notches distinct, rounded or slightly pointed, the three anterior ones only very shallowly indented; anal lunule small.

Oral side: Food grooves remarkably broad; periproct on the anterior slope of the anal lunule. Color: The test in life is dark brown, the denuded test is whitish. Distribution: This species occurs from Cape Hatteras, NC, USA to the Bahamas and throughout the Gulf of Mexico at depths of 12 to 90 m. Biology: E. aberrans lives like the other sand dollars on sand flats and fine shell gravels, shallowly buried. Remarks: Phelan (1972) defined clear differences between E. aberrans and E. michelini: He could not find intermediate specimens between those with only very slight marginal indentations and those with deep marginal notches. Usually, they do not co-occur in the same populations because E. aberrans is likely to feed on larger organic particles than E. michelini, as indicated by their distinctly broader food grooves. However, in The World Echinoidea Database both species are synonymized (Fig. 3.81). Literature: Phelan 1972; Serafy 1979; Hendler et al. 1995. Encope emarginata Leske, 1778 Test: Strong, longer than wide; upper side like a broad plateau; length up to 141 mm. Petals: Posterior pair longest and slightly curved to the midline. Lunules: Marginal notches deep, open or closed, anal lunule large and more or less irregularly elongated, reaching far between the posterior petals, much longer than in the two other Caribbean species of Encope. Distribution: E. emarginata is known along the American east coast from Florida throughout the Gulf of Mexico southward to the mouth of the La Plata, Argentina, from the intertidal zone to a depth of approximately 50 m. Remarks: E. emarginata is distinguished from Leodia sexiesperforata, which also has six lunules, by the stronger test, the longer anal lunule, and by having five gonopores instead of four. Encope grandis L. Agassiz, 1841 Test: Very strong, usually slightly wider than long; margin thick; posterior end truncated; highest point at the anterior edge of the anal lunule; length up to 100 mm. Apical system: Anterior to highest point. Petals: Posterior pair much longer than the three anterior ones, curved around the lunules. Lunules: Marginal notches very widely open; anal lunule extremely large, broadly elongated. Oral side: More or less concave, periproct in depression anterior of anal lunule, distinct food grooves. Color: In life, the animal is brown to reddish-brown, the denuded test is whitish.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

B

A

C

D

 103

E

apical disc

apical lunule

F

G Fig. 3.81: (A–C) Encope aberrans. Test length 127 mm; west coast of Florida, USA. (A) Aboral side: the petals are long, the posterior pair slightly longer than the others. The anterior notches are inconspicuous. (B) Oral side: the food grooves are broad, indicating that relatively large particles are transported to the mouth; especially in comparison to Encope michelini, which has developed finer food grooves. (C) Profile. (D–G) Encope emarginata; test length 138 mm; Isla Margarita, Venezuela. (D) Aboral side: the posterior notches are open, the anterior ones just closed. The anal lunule is extremely long and often irregularly formed. (E) Oral side: the food grooves are strongly branched. (F) Test length 117 mm; Sao Paulo, Brazil. ZMH. Aboral side: the spines are very short. (G) Profile: there is a broad plateau on the aboral side, slightly higher posteriorly than anteriorly. Scale bar is 50 mm.

Distribution: This species is endemic to the northern part of the Gulf of California, Mexico, living from the littoral zone to a depth of 18 m. Biology: This species lives in patches with Mellita grantii moving according to the physical environment like the sand level. In favorite substrata, there may be more than 300 specimens per square meter. The mean life expectancy

is 5 years for E. grandis and only 1.2 years for M. grantii (Ebert & Dexter 1975). Encope laevis H.L. Clark, 1948 Test: Flattened with an almost horizontal upper side from just behind the anterior lunule to a point just in front of the anal lunule.

104 

A

 3.2 Superorder Neognathostomata A.B. Smith, 1981

B apical disc

apical lunule

D C

Fig. 3.82: (A, B) Encope grandis. Test length 72 mm; Gulf of California, Mexico. (A) Aboral side: the anal lunule is extremely large, and the ambulacral notches widely open. (B) Profile: the highest point is just in front of the anal lunule. Scale bar is 30 mm. (C, D) Encope laevis. Length of test 112 mm, width 116 mm; Nicaragua, Pacific side. ZMUC ECH 193. (C) Aboral side: the “pavement” formed by the spines is quite smooth. (D) Oral side: the spines are much longer than on the aboral side and differentiated.

Petals: Very long and with a blunt, broad end. Lunules: Marginal ones small; anal lunule not much larger than marginal lunules, positioned mainly between the distal ends of the posterior petals. Spines: Aborally inversely cone-shaped, base in contact with each other, thus forming a smooth “second surface”; oral spines much longer and differentiated according to their function. Color: Spines olive-brown, denuded test more olivegray. Distribution: Nicaragua, in shallow water (Fig. 3.82).

Encope michelini L. Agassiz, 1841 Test: Strong, subcircular, or slightly elongated, length up to 140 mm; highest point just in front of the anal lunule, forming a more or less raised hump. Petals: Long. Lunules: Marginal notches more or less deeply indented, the posterior pair a little longer than the other three. Anal lunule elongated with parallel sides, positioned in the main part between the posterior petals. Oral side: Food grooves narrow and fine; deep pressure drainage channels lead into the marginal notches.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

A

 105

B

C

apical disc

anal lunule

Fig. 3.83: (A–C) Encope michelini. Test length 116 mm; west coast of Florida, USA. (A) Aboral side: the petals are long with densely arranged pore pairs. The posterior edge of the test has been damaged and subsequently healed. (B) Oral side: the food grooves of E. michelini are finer than those of Encope aberrans. The species may therefore feed on smaller organic particles than the latter. The grooves are highly branched; deep pressure drainage channels lead into the ambulacral notches. The fields alternating with the regions along the food grooves are set with larger tubercles, carrying longer spines for locomotion. (C) Profile: the highest point between apical disc and anal lunule is extremely raised, often formed like a hump. In other specimens, it may not reach that height. Scale bar is 50 mm.

Color: In life, the test is dark brown or reddish-brown, the bare test is whitish. Biology: E. michelini lives on fine to coarse sand, burrowing to a maximum depth of 10 mm. It presumably feeds on small organic particles, as indicated by the narrow food grooves. Distribution: This species is reported from Cape Hatteras, NC, USA, along the coasts of Florida and through the Gulf of Mexico to Cozumel, Yucatan Peninsula, Mexico, but it is not found at the Bahama Islands. Its bathymetric range is 3 to 90 m. Remarks: Usually, E. michelini do not co-occur with E. aberrans, feeding on smaller food particles (Phelan 1972). Encope micropora L. Agassiz, 1841 Test: Usually strong; in profile variable: anterior part of aboral side gently arched to the highest point distinctly in front of the apical system, from where it slopes more or less straight to the posterior edge, or it may have a more or less flattened upper side; outline round or rounded pentagonal, the posterior margin more or less truncated; length may reach 157 mm.

Petals: Of approximately the same size. Lunules: Marginal ones elongated, more or less open; anal lunule larger than the marginal ones, located between the distal ends of the posterior petals, or at least for the most part. Oral side: Flat. Color: In life, the species is brownish, the cleaned test is grayish-white. Distribution: This species is widely reported along the west coast of America: from the Gulf of California and the west side of Baja California, Mexico to Peru, including the Galapagos Islands. It lives in the littoral zone to a depth of 30 m. Remarks: Mortensen edited the Report on the Echini of the Warmer Eastern Pacific Collected by the Velero III after the death of H.L. Clark in 1947. Herein, Clark described a couple of new species of Encope. However, Mortensen thought that this division would go too far, and he labeled these species as subspecies of E. micropora. Two other species were also labeled as subspecies, E. micropora borealis A.H. Clark, 1946 and E. micropora californica Verrill, 1870.

106 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

C

A

B

E

apical disc

anal lunule

D

apical disc

anal lunule

F

apical disc anal lunule

G

100 mm

Encope fragilis Encope laevis Encope perspectiva Fig. 3.84: (A–E) Encope micropora. Test length 86 mm; Gulf of California, Mexico. (A) Aboral side: the rounded notches may be open or closed. The anal lunule lies to the posterior of the distal ends of the posterior petals. At the right side, the test has been damaged and rehealed. (B) Profile: the upper side is flattened. Scale bar is 30 mm. (C, D) Test length 95 mm; Gulf of California, Mexico. (C) Aboral side: the posterior petals extend a short distance beyond the start of the anal lunule. (D) Profile: the highest point is just in front of the anal lunule. Scale bar is 30 mm. (E, F) Test length 94 mm, Santa Teresa, Nicoya Peninsula, Costa Rica. (E) Aboral side. (F) Side view. (G) Profiles of species of Encope. After H.L. Clark (1948). The shaded areas show the positions of the lunules, the highest point of the test is marked by a vertical line.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 



The authors speak today of the micropora group. Encope micropora borealis A.H. Clark, 1846. Endemic to the northern Gulf of California. Encope micropora californica Verrill, 1870. Endemic to the Gulf of California. Encope micropora cocosi H.L. Clark, 1948. Cocos Island. Encope micropora ecuadorensis H.L. Clark, 1948. Ecuador. Encope micropora fragilis H.L. Clark, 1948. Thin test; Mexico. Large test, very small marginal lunules, anal lunule close to posterior margin. Encope micropora insularis H.L. Clark, 1948. Socorro and Clarion Islands, 25 to 82 m. Encope micropora irregularis H.L. Clark, 1948. Outline irregular, Columbia to Costa Rica. Encope micropora micropora L. Agassiz, 1841. Encope micropora tetrapora L. Agassiz, 1841. Endemic to the Galapagos Islands, Pearl Island, Panama; large with high plateaued test; beige colored with bright purple around the edge and lunules (Fig. 3.84). Encope perspectiva L. Agassiz, 1841 Petals: Of about the same length. Lunules: Marginal lunules small, anal lunule distinctly larger, located far between the posterior petals. Remarks: This species strongly resembles E. laevis. Apart from the profile, the main difference lies in the shape of the aboral spines: In E. perspectiva, the tips of the primary spines are formed like oval balls, not in contact with each other. The surface feels rather coarse. In E. laevis, they are like inverse cones, the upper surfaces touching each other, thus forming a smooth plateau.

A

 107

Distribution: The species is recorded from Baja California, Mexico to Costa Rica in shallow water. Encope wetmorei A.H. Clark, 1946 Test: Length may reach 145 mm. Petals: Not very broad. Lunules: Anal lunule high between the posterior petals. Spines: Tips of aboral spines of different length with more or less long, inverted cones, which are loosely in contact. Color: Test olive-gray. Distribution: Pearl Island, Panama, Pacific coast of Mexico; bathymetric range 9 to 54 m (Fig. 3.85). Genus Leodia Gray, 1852 The genus contains a single extant species. Leodia sexiesperforata (Leske, 1778) Test: Flat and fragile with thin margin; circumference subcircular; length up to 100 mm. Apical system: Very slightly posterior with four gonopores. Petals: Short and of equal length. Lunules: All slit-like, anal lunule shorter than the five marginal ones. Oral side: Main branches of food grooves diverge very close to the peristome on the basicoronal plates; pressure drainage channels simple; periproct well separated from the anterior side of the anal lunule. Spines: Upper spines short and club-shaped, oral spines longer and differentiated. Color: Light brown to yellowish, bare test white.

B

Fig. 3.85: Encope perspectiva. Test length 108 mm. CASIZ 87785, photos by R. Mooi, California Academy of Sciences, San Francisco. (A) Aboral side. (B) Close-up of upper surface.

108 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

Distribution: This species is widely reported from Cape Hatteras, North Carolina to Florida, the West Indies, and from the Yucatan Peninsula along the coast of Central and South America to Uruguay. It lives in the littoral zone and down to a depth of 60 m, but mostly in waters less than 25 m deep. Biology: L. sexiesperforata prefers open sand flats without sea grass and algae, and is restricted entirely to biogenic carbonate sediments. It often co-occurs with E. michelini in mixed flocks. The Mellita species, in contrast, occur on siliceous sediments of terrigenic origin with medium particle sizes (100–500 µm) (Harold & Telford 1990).

Mellita quinquiesperforata (Leske, 1778). In the western Gulf of Mexico, the Antilles, and southward to Brazil and Argentina. This species is more closely related to the four Pacific species of Mellita than to the two other Atlantic species. Mellita tenuis H.L. Clark, 1940. Along the west side of Florida to the Mississippi Delta. There was much confusion about the taxonomy of these Atlantic species because they have been included in M. quinquiesperforata for a long time. However, in 1990, Harold & Telford revised the genus and separated the two other species according to shape and geographical distribution.

Genus Mellita L. Agassiz, 1841 Test: Flat, of medium to large size with thin margin, aboral side may be slightly elevated. Apical system: Four gonopores. Petals: Of approximately similar shape. Lunules: Five ambulacral lunules or open notches, anal lunule more or less between the posterior petals. Oral side: Flat, conspicuous food grooves in ambulacra, main branches diverging on post-basicoronal plates (in contrast to Leodia); often with very distinct lateral depressions for pressure drainage; peristome subcentral, periproct close to the peristome in front of the anal lunule. Spines: On the aboral side, very short and thickened, orally longer and more or less differentiated. Seven extant species of Mellita are recorded, living along the tropical to temperate coasts of North and South America in shallow water.

Pacific side: Four species are reported from the west coast; phylogenetically closely related to M. quinquiesperforata. They live more or less sympatric from the Gulf of California to Peru. The northern part of the Gulf is exclusively set by M. grantii: Mellita grantii Mortensen, 1948. Mellita kanakoffi Durham, 1961. Mellita longifissa Michelin, 1858. Mellita notabilis H.L. Clark, 1947. Remark: Lanthonia gen. nov. Coppard 2016 included the former Mellita longifissa and Mellita grantii. Both are morphologically different from the Atlantic Mellita quinquiesperforata and from the other Mellita, Atlantic and Pacific, in having narrow ambulacra between the lunules and the food grooves. Further the new genus is different by the dentation of pedicellaria from all other Mellita. In the following descriptions, the species are listed alphabetically:

Atlantic side: Three species live along the eastern coasts in well limited regions only slightly overlapping at the estuaries of the Mississippi: Mellita isometra Harold & Telford, 1990. From Nantucket, Massachusetts; along the east side of Florida.

A

B

Mellita grantii Mortensen, 1948 Test: Outline is almost round, scarcely wider than long, highest point at or just anterior to apical system; length may reach 50 mm. Petals: Posterior pair curved downward to midline.

C

Fig. 3.86: Leodia sexiesperforata. (A, B) Test length 77 mm; northern coast of Cuba. (A) Aboral side: the test is set with very short clubshaped spines. (B) Oral side: the main food grooves diverge immediately at the peristome on the basicoronal plates (in contrast to the other Mellitids, in which the grooves diverge on the post-basicoronal plates. The pressure drainage channels are distinct. (C) Length of test 71 mm; Florida, USA. Aboral side: the petals are short and of equal size. The five ambulacral lunules are longer than the anal lunule.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 



Lunules: Marginal ones rather small and slit-like; anterior pair extending horizontally; anal lunule longer than the marginal ones, distinctly separated from the nondepressed periproct. Oral side: Main branches of food grooves not divergent or sinuous; pressure drainage channels shallow and inconspicuous; oral spines little differentiated. Distribution: This species is known from the Gulf of California to Panama, settling exclusively in the northern part of the Gulf; more southward co-occurring with M. longifissa and M. kanakoffi. It lives from the intertidal zone to a depth of 3 m.

 109

Mellita isometra Harold & Telford, 1990 Test: Only slightly wider than long, circular or rounded pentagonal, highest point slightly anterior to apical system; length rarely exceeding 100 mm. Petals: Straight, posterior pair equal in length to petal in A III, lateral pair shortest. Lunules: Marginal ones approximately as long as the anal lunule. Oral side: Main branches of food grooves divergent and moderately sinuous; pressure drainage channels with conspicuous tributaries.

A

B

C

D

Fig. 3.87: Mellita grantii. (A–D) Test length 42 mm; Gulf of California, Mexico. (A) Aboral side: the anal lunule is not much longer than the paired lunules. (B) Oral side: the periproct is not depressed at the anterior border of the anal lunule, but is distinctly separated. (C, D) Specimens with spines, test length 33 mm; Gulf of California, Mexico. (C) Aboral side: in some specimens, the plate sutures are marked by lines. It is, however, not a feature of taxonomic importance. (D) Oral side: the oral spines are only slightly differentiated, in contrast to the other species of Mellita.

110 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

Spines: Fringe spines of anal lunule longer than locomotory spines. Distribution: Only reported from the east coast of the USA, from Nantucket Island, Massachusetts to Florida. This species lives in shallow water in 1 to 3 m depth, mostly in patches of 10 to 15 specimens, sometimes more. Mellita kanakoffi Durham, 1961 Test: Wider than long; highest point slightly anterior to apical system; maximum length 107 mm. Petals: Posterior ones as long as in A III. Lunules: Marginal ones of same size, posterior pair curved, anal lunule long. Oral side: Main food grooves widely divergent and sinuous; two branches parallel to the posterior margin; deep pressure drainage channels with conspicuously depressed tributaries, intervening areas distinct, but not hummocky. Spines: Well differentiated. Distribution: This species settles along the west coast of central Mexico southward to Panama. It lives in very shallow water like the other species of Mellita. Remarks: Mellita longifissa was a well-established species since Michelin (1858). However, Durham (1961) recognized that the populations included a second, distinctly different and much more common species, which he described as M. kanakoffi.

A

Mellita longifissa Michelin, 1858 Test: More or less wider than long; highest point just anterior to apical system; length may reach 75 mm. Petals: Posterior ones curved to midline. Lunules: Marginal ones narrow, posterior pair curved; anal lunule very long and narrow, reaching very far between the posterior petals. Oral side: Periproct depressed into anterior border of the anal lunule; main food grooves not divergent and not distinctly sinuous, no branches parallel to posterior margin; pressure drainage channels shallow and inconspicuous, no tributaries leading into lunules. Spines: Well differentiated. Distribution: M. longifissa is found from Baja California and the Gulf of California to Ecuador and the Galapagos Islands. It occurs sporadically in populations of M. kanakoffi or M. notabilis living from the intertidal zone to a depth of 5 m. Mellita notabilis H.L. Clark, 1947 Test: Distinctly wider than long; highest point of test anterior; posterior margin rather truncated; length more than 80 mm. Petals: Posterior ones longer than petal in A III, curved to the midline.

B

C

Fig. 3.88: Mellita isometra. (A) Test length 68 mm, height 7 mm; Palm Beach, east coast of Florida, USA. Aboral side of an adult specimen and of two juveniles, test length of upper specimens 27 mm. The posterior pair of petals is approximately as long as the petal in A III. The lunules of the juveniles are still short, not yet slit-like as in the adult specimen. (B) Oral side: the paired lunules are not much shorter than the anal lunule. (D) Profile of test: the highest point lies anterior to the apical system. Young specimens are often weakly conical, the point slightly anterior.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

A

C

 111

B

D

Fig. 3.89: Mellita kanakoffi. (A) Test length 51 mm; Golf of Guayaquil, Ecuador. Aboral side: the posterior petals are curved to the midline. (B) Oral side: the pressure drainage channels and their tributaries are very distinct. (C, D) Test length 52 mm; León, Nicaragua. (C) Aboral side. (D) Oral side: only the longer spines, here of lighter color, are used for locomotion in the sandy sediment. The other spines are shorter and mostly bent sheltering the tube feet.

Lunules: All lunules irregularly looking; anterior pair of marginal ones mostly directed to the posterior; posterior lunules curved inwards, anal lunule long and narrow. Oral side: Main food grooves widely divergent and strongly sinuous; two branches parallel to the posterior margin; strongly developed pressure drainage channels, very deep tributaries alternating with hummocky-like areas. Spines: Well differentiated. Distribution: M. notabilis is known from the west coast of central Mexico southward to Panama. It is likely to co-occur with M. longifissa, presumably in very shallow water.

Mellita quinquiesperforata (Leske, 1778) Test: Much wider than long, highest point well anterior of apical system; maximum length 100 mm. Petals: Posterior ones rather straight. Lunules: Anterior pair of marginal ones may be transverse or obliquely directed to the posterior; posterior lunules curved toward midline; anal lunule longer than the others. Oral side: Main food grooves moderately divergent; pressure drainage channels with distinct tributaries. Spines: Fringe spines in the lunules conspicuously paddle-shaped.

112 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

A

B

C

D

Fig. 3.90: Mellita longifissa. (A, B) Test length 44 mm, width 53 mm; San Salvador. (A) Aboral side: the posterior petals and the posterior lunules are distinctly curved to the midline. (B) Oral side: the food grooves lead closely along the ambulacral lunules. There are no tributaries of the pressure drainage channels developed along the long sides of the lunules. (C, D) Test length with spines 51 mm, width 59 mm; San Salvador. (C) Aboral side: the fringe spines at the lunules are long. (D) Oral side: the spines are well differentiated. The food grooves closely surround the lunules.

Distribution: This species is known in the western Gulf of Mexico, from the Mississippi Delta along the coasts of Central and South America to southern Brazil. It also occurs at the Greater Antilles, living between 1 and 3 m deep. Mellita tenuis H. L. Clark, 1940 Test: Thin margin, subcircular or even longer than wide, shallow-domed plateau around or posterior to apical system; length may reach 150 mm. Petals: Straight, posterior pair approximately 20% longer than petal in A III.

Lunules: Marginal ones narrowly elongated with parallel sides, anal lunules longer than marginal ones. Oral side: Main branches of food grooves divergent and moderately sinuous; pressure drainage channels with conspicuous tributaries. Spines: Fringe spines of anal lunule longer than locomotory spines. Distribution: This species is restricted to the eastern part of the Gulf of Mexico, from the west coast of South Florida to western Louisiana (Dulac Beach), at the Mississippi Delta slightly overlapping with the distribution of

3.2.2 Order Clypeasteroida L. Agassiz, 1835 



A

B

D

E

 113

C

F

Fig. 3.91: (A–C) Mellita notabilis. Test length 45 mm, León, Nicaragua. (A) Oral side: the medium-sized specimen has already developed the typical hummocky areas alternating with the tributaries of the pressure drainage channels along the long sides of the paired lunules. Test length 81 mm; León, Nicaragua. The test is damaged around the margin. (B) Aboral side: the posterior petals are distinctly longer than the three anterior ones. (C) Oral side: the lunules look irregular because the tributaries of the pressure drainage channels are very deep and the alternating areas formed like hummocks. (D–F) Mellita quinquiesperforata. Test length 51 mm; Vera Cruz, Mexico, Caribbean coast. (A) Aboral side: the sea urchin has been drilled by a predatory snail. (B) Oral side: the pressure drainage channels and their tributaries are distinct. (C) Test length 56 mm; Vera Cruz, Mexico, Caribbean. (C) Aboral side: the fringe spines at the lunules are distinctly paddle-shaped.

M. quinquiesperforata. It lives from the intertidal zone to a depth of 3 m. Genus Mellitella Duncan, 1889 The genus was often put together with Encope because both have four gonopores. However, Coppard confirmed its separated state by morphological and molecular analysis (personal communication). Mellitella stokesii L. Agassiz, 1841 Test: Thin with sharp margin; upper side conically raised, highest point slightly anterior to apical disc, outline round, length up to 75 mm. Petals: Remarkably short; posterior pair shorter than the three others. Lunules: Marginal notches deep and usually open, sometimes closed as lunules; anal lunule small, very near the posterior margin. Oral side: Slightly concave; food grooves fine but distinct.

Spines: Well differentiated. Distribution: M. stokesii is known along the west coast of Central and South America from Mexico to Peru living in the littoral zone.

3.2.2.11 Family Scutellidae Gray, 1825 The family consists traditionally only of fossil species. The integration of Scaphechinus relies on a weakly supported relationship (Mooi 2015, personal communication). Genus Scaphechinus A. Agassiz, 1864 It is still the subject of discussion in which family this genus has to be placed. It is closely related to the pedomorphic genus Sinaechinocyamus, which was listed in the family Taiwanasteridae by Wang, 1984. Both genera should be better put into a family of their own (Mooi 2015, personal communication).

114 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

A

B

C D

E

Fig. 3.92: Mellita tenuis. (A–C) Test length 74 mm; Sanibel Island, west coast of Florida, USA. (A) Aboral side: the posterior petals are distinctly longer than the petal in A III. (B) Oral side: the paired lunules, shorter than the anal lunule, are surrounded by the gently arched main branches of the food grooves. (C) Profile: the test is low domed. Scale bar is approximately 60 mm. (D) Test length 104 mm, Sanibel Island, west coast of Florida, USA. Aboral side: the test is densely covered by very short spines ending in a distinct knob. (E) Juvenile specimens, test length of largest sample 26 mm; Naples, FL, USA. Initially, the ambulacral lunules appear as shallow notches at the margin (left). Later, in ontogeny, they become deeper and finally closed (right).

A

B apical disc

apical lunule

Fig. 3.93: Mellitella stokesii. (A) Test length 42 mm; Bay of Fonseca, El Salvador. ZMH. Aboral side: the outline of the test is circular, the ambulacral notches mostly open, sometimes closed. The anal lunule lies very near the posterior margin. (B) Profile: the test is conically raised. Scale bar is 30 mm.

3.2.2 Order Clypeasteroida L. Agassiz, 1835 



 115

Test: Rather solid; outline round to more or less broadly pentagonal; upper side slightly domed, in large specimens ring-shaped depression around the petaloid area with “channels” at the midline of the interambulacra leading to the thin margin; test length may reach 71 mm. Apical system: Subcentral with four gonopores. Petals: Long and broad, two-thirds the test length, distally slightly open. Oral side: Flat or slightly concave; bifurcating main food grooves with distal side branches. Spines: On the upper side short, distally thickened and curved, longer and also curved orally. Color: In life, the test is uniformly dark purplish; the denuded test is usually grayish-white. Distribution: The species is known in Japanese waters from Hokkaido to Kyushu, the Korean Peninsula, north China, east Russian coast and Kamchatka Peninsula and the Aleutean Islands from the intertidal zone to 125 m. Four very similar species are described.

Distribution: Northern Japan to the coast of Kamchatka at 7 to 10 m.

Scaphechinus brykovi Budin, 1983 Occurrence: From Peter the Great Bay into the Japanese Sea.

3.2.2.12 Family Taiwanasteridae Wang, 1984 Wang (1984) described a new Scutellid and gave it the name Taiwanaster. He included it simultaneously in a new family Taiwanasteridae ignoring the name Sinaechinocyamus established in 1979 by the Continental-Chinese Liao. Mooi (1990a,b,c,d) felt that Scaphechinus and the pedomorphic Sinaechinocyamus are nearly related and should be placed in a family of their own.

Scaphechinus griseus Mortensen, 1927 Characteristics similar to the genus. This species is differentiated from S. mirabilis by its slightly but distinctly supramarginal periproct.

A

Scaphechinus mirabilis A. Agassiz, 1863 Characteristics similar to the genus. Test: In outline rather rounded or pentagonal. Oral side: Periproct in a small notch at the posterior margin. Remarks: Mortensen mentioned that all intermediates in outline between S. mirabilis and S. tenuis occur. Scaphechinus tenuis (Yoshiwara, 1898) Mortensen (1948) described the differences to Scaphechinus mirabilis as follows: Test more fragile and in outline much broader than long. However, he admitted that intermediates are found between rounded pentagonal and very broad specimens. Shigei (1986) did not list S. tenuis as a separate species. Distribution: From Kagoshima to Hokkaido, shallow water.

B

Fig. 3.94: Scaphechinus griseus. (A) Aboral side of test, length 31 mm, Preobrazhenie, Ost-Siberia, Russia. The periproct is positioned on the upper side, just above the margin. (B) Oral side: the bifurcating main food grooves branched again distally.

116 

 3.2 Superorder Neognathostomata A.B. Smith, 1981

A

B

D

E

C

F

Fig. 3.95: Scaphechinus mirabilis. (A, B) Test length 47 mm; Shizuoka, central Japan, Pacific. (A) Aboral side: the test is broadest toward the posterior. The periproct lies in a small notch at the posterior margin. (B) Oral side: the food grooves have several distal branches. (C) Aboral side of an adult specimen. Length 53 mm, Chiba Peninsula, Japan. (D, E) Test length 49 mm; Kanakawa Prefecture, central Japan, Pacific. (D) Aboral side. (E) Oral side: on the upper side the spines are short and slightly club-shaped, on the oral side they are much longer and not thickened. (F) Aboral side of a juvenile specimen. Length 7 mm, Katahama Bay, Japan.

Genus Sinaechinocyamus Liao, 1979 Test: Very small; aboral side lowly domed, margin thin; hardly exceeding 10 mm in length. Apical system: Slightly anterior, four gonopores, hydropores scattered over anterior part of madreporite. Petals: Well-developed, but short, slightly curved and distally open. Oral side: Perfectly flat, food grooves indistinct (not as visible as in the drawing), short and simple; peristome central with conspicuous stereom points entering the opening. Periproct: Positioned on the aboral side just above the margin. Tuberculation: Fine and densely distributed. Biology: In its pedomorphic shape, Sinaechinocyamus strongly resembles a young Scaphechinus mirabilis Lütken, 1864 according to R. Mooi (1990). The gonopores are already developed at a size of only 3 mm, whereas in Scaphechinus, they only open at a size of approximately 20 mm. Thus,

Sinaechinocyamus is mature at the size and with the morphology of a juvenile. This tiny echinoid may be adapted to a life in interstitial habitats, like the also very small Echinocyamus pusillus, which nests among small pebbles. However, the records about the habitat of Sinaechinocyamus mention “muddy sand bottom” or “fine sands”. Further research is needed. Remarks: In miniaturized taxa such as Sinaechinocyamus, a weight belt is present and supports its membership to the Scutellina. Literature: Liao 1979; Wang 1984; Mooi, 1988, 1990c, 1996; Chen & Chen 1994; Mooi & Chen 1996. The genus consists of two species: Sinaechinocyamus mai Wang, 1984 Test: Very small, flat, aboral side slightly domed, highest point just in front of the apical system; in outline anterior part rounded, widening to the sides, then narrowing

3.2.2 Order Clypeasteroida L. Agassiz, 1835 

A

 117

B

C

D

E

Fig. 3.96: (A, B) The two species of Sinaechinocyamus. Scale bar is 5 mm (redrawn after Mooi 1990). (A) Sinaechinocyamus planus. (B) Sinaechinocyamus mai. Aboral side and oral side. (C–E) Sinaechinocyamus mai. Length 9.5 mm; Taiwan. (C) Aboral side: the gonopores are fully developed. The periproct is positioned on the upper side. (D) Oral side. (E) Profile. Scale bar is 2 mm.

again toward a slightly pointed posterior end; length may reach 10 mm. Petaloid area: Little more than half the test length. Petals: Well-developed, paired petals half open, anterior one widely open; pore pairs conjugate. Oral side: Perfectly flat with indistinct food grooves; peristome central, stereom points entering the opening very conspicuously. Periproct: On the aboral side. Distribution: South China Sea, at depths of 30 to more than 70 m. Sinaechinocyamus planus Liao, 1979 Test: In outline, elongated with a slight indentation in each posterior ambulacrum; maximum length 8 mm. Distribution: Yellow Sea.

Incerta sedis Genus Marginoproctus Budin, 1980 Marginoproctus djakonovi Budin, 1980 Test: Small and lentil-shaped; in outline as broad as long or slightly oval; upper side hardly elevated, margins rounded; internal supports in 10 radial buttresses; length 12 mm. Apical system: Four gonopores and one to four hydropores. Petals: Small and poorly developed, straight and widely open. Oral side: Sunken toward the central peristome, periproct marginal or just supramarginal in a small notch; food grooves absent. Remarks: In miniaturized taxa, such as in Marginoproctus, a weight belt is present and supports its membership to the Scutellina.

118 

 3.3 Superorder Atelostomata von Zittel, 1879

3.3 Superorder Atelostomata von Zittel, 1879

In contrast, the spatangoids are able to burrow in fine substrates or even in mud, living infaunally. They have adapted to their habitat through their funnelbuilding podia and the diverse fascioles crossing their test (Fig. 3.97).

This taxon of more or less heart-shaped echinoids includes the orders Holasteroida and Spatangoida. Both arose in the early Cretaceous from the Disasteroida, having lost all traces of teeth. They initially occupied similar paleoenvironments, and they resemble each other in many ways, e.g., they both developed enlarged tube feet around the peristome and the phyllodes, as well as bands of tiny spines termed fascioles, which maintain a fresh water current over the surface of the test. However, by the late Cretaceous, the holasteroids shifted into deeper water settings down the continental slope, whereas the spatangoids remained on the shelf in shallower water. Additionally, the lifestyles of the two taxa differ: the holasteroids prefer living epifaunally or semiburied, burrowing only in porous and coarse sediment. The fascioles are mostly restricted to the subanal area, keeping the test free of feces, and specialized podia are lacking.

3.3.1 Order Holasteroida Durham & Melville, 1957 In the Cretaceous, the holasteroids were very common, but around the Cretaceous–Tertiary boundary, they underwent a major decline, many genera became extinct. A new group, the Urechinina, evolved, and made up the great majority of the extant holasteroids, living exclusively in the deep sea. They are the most successful group of echinoids in that habitat (Note: the density of echinoids in the abyssal is generally low: 0–78 individuals/104 m2 in the Bay of Biscay; David & Sibuet 1985).

A Holasteroida

Spatangoida

peristome

labrum

sternal plate(s) episternal plates

F

periproct

B Holasteroida G3

O III

O III

O II

O IV G4

G1 OV

G3

G2

OI

Spatangoida G2

O IV

O II

G4

G1

OV

OI

Fig. 3.97: Atelostomata: the two extant orders of Atelostomata differ in the arrangement of the interambulacral plates posterior to the peristome that forms the plastron: Holasteroida (left side), generalized: the elongated labrum (1) is connected to a single sternal plate (2) followed by two episternal plates. Modified after Mooi & David (1996). (B) Spatangoida (right side), generalized: the narrow labrum (1) is connected to two enlarged sternal plates (2a/2b) followed by two episternal plates. The plastron carries large tubercles and strong spines specialized for locomotion within the sediment. Scale bar is 20 mm. (B) Both orders also differ in the plate scheme of the apical system: Holasteroida (left side): the arrangement is elongated, the ocular plates OII and OIV touch each other and separate the anterior (G2/G3) and the posterior (G4/G1) genital plates. The ocular plates OI and OV form the posterior end (Corystus relictus; modified after Mooi & David 1996). Spatangoida (right side): the apical system is more compact, the enlarged genital plate G2 (madreporite) expands to the posterior, separating G1 and G4 and the ocular plates OI and OV (Brissus; after Mortensen 1951). Scale bar is 2 mm. Secondly, one or two gonopores may have been lost in some groups.



The deep sea environment is relatively poor in organic material, and the holasteroids have developed special modes of feeding. As deposit feeders, they depend on organic particles that drift down from surface waters and which episodically accumulate at the surface of the substrate in the deep. The Recent pourtalesiids skim the organic particles and channel them by the anterior funnel toward the peristome. They have developed an elongated or, one could even say, bottle-shaped test, and the locomotion is more or less horizontal. The order consists of six extant families: Calymnidae Mortensen, 1907. Corystusidae Foster & Philipp, 1978. The following four families differ distinctly from the former. A.B. Smith compiled them in the Infraorder Urechinina H.L. Clark, 1946: Carnarechinidae Mironov, 1993. Plexechinidae Mooi & David, 1996. Plastron interrupted. Pourtalesiidae A. Agassiz, 1881. Plastron interrupted. Urechinidae Duncan, 1889. Plastron not interrupted. Literature: Agassiz 1881; Mortensen 1907, 1950; Gage 1984; David & Sibuet 1985; David 1984, 1988, 1998; Mooi & David 1996; David & Mooi 2000; Saucède et al. 2003a, 2009; Smith 2004.

3.3.1.1 Family Calymnidae Mortensen, 1907 Smith (2004) grouped two extant genera in this family: Calymne Thomson, 1877. North Atlantic. Sternopatagus de Meijere, 1903. Timor Sea, Indonesia. The two monospecific genera resemble each other in some aspects of plate architecture and they share the presence of a marginal fasciole, but not its location on the test. However, in other important characteristics, they differ significantly: in Calymne, the peristome is flush with the test, whereas in Sternopatagus, it lies at the end of a deep funnel; in Calymne, the plastron is continuous, whereas in Sternopatagus, it is disjunct. Saucède et al. (2009) redescribed the genus Calymne based on newly found and partly intact specimens. He would prefer to keep this genus by itself in the family Calymnidae, as Mortensen (1907) had originally pro­posed. Genus Calymne Thomson, 1877 Calymne relicta Thomson, 1877 The type material of Calymne, two specimens of approximately 23 and 28 mm length, was already fragmented when A. Agassiz drew and redescribed it in 1881. The sole intact adult specimen was found using a 0.25 m2 grab during a Russian cruise in 1983 in the North Atlantic (RV Akademik Kurchatov cruise 37).

3.3.1 Order Holasteroida Durham & Melville, 1957 

 119

Updated description of genus and species: Test: In adults high, in aboral view ovoid, compressed at the sides, ambitus low; in outline anteriorly rounded without frontal notch; posterior side almost vertically truncated forming a wide bulge above the periproct, more conspicuous in younger specimens; test as a whole extremely thin and fragile. Apical system: Stretched along the midline; two genital pores in G2 and G3, anterior and posterior pairs of genital plates separated by ocular plates OIV and OII; a few hydropores in G2. Ambulacra: Flush with the test; unpaired ambulacrum slightly depressed only near peristome; no tendency of forming petals because the aboral plates are as large and similar in size to interambulacral plates; small aboral tube feet associated with tiny single pores, phyllopodia near peristome with rudimentary digits. Oral side: Convex with a wide keel toward the posterior; labrum small, always in contact with the sternal plate; peristome anterior, flush with the test, small and circular. Periproct: High on the posterior side, longitudinally oval; marginal fasciole present. Spines: Miliary spines covering entire test; primary spines rather sparse, shape very diverse according to their position on the test, e.g., spoon-shaped on plastron, shovelshaped around periproct and up to the posterior aboral side; in subanal tuft below fasciole long and slender. Pedicellariae: Rostrate ones present as four types in adults: ophicephalous pedicellariae umbrella-shaped; triphyllous pedicellariae present, but tridentate and globiferous pedicellariae never observed. Biology: The different primary spines of Calymne are unusual for holasteroid echinoids. The newly found specimens allowed a comparison of these peculiar spines for the first time in form and possible function with spatangoid spines, which are relatively well studied. The asymmetry of the areoles permits conclusions about the preferred direction of the power stroke of the spines and hypothetical behavior patterns of Calymne. 1. There are five to six large tubercles just anterior of the apical system (probably bearing long spines). These tubercles do not resemble those associated with the tuft of long food-gathering or funnel-building spines in spatangoids. Because there are no specialized funnel-building tube feet in Calymne, long aboral spines have to be associated with another function like protection for the gonopores. 2. As the primary spines on the sides are sparsely distributed, they do not seem to function in burrowing, as is the case in spatangoids, in which the lateral spines are densely packed.

120 

 3.3 Superorder Atelostomata von Zittel, 1879

3. At the frontal side, above the fasciole, the areoles of the spines, which consist of several different types of about the same length, indicate a downward power stroke, whereas the spines located below the fasciole perform an upward power stroke. 4. Two frills of shovel-shaped spines on the posterior aboral side and below the periproct have asymmetrical areoles, which are enlarged in a direction converging toward the highest point of the posterior bulge. This means that the movement of aboral spines is directed to the posterior, whereas the movement of lateral and posterior spines is directed aborally. This arrangement is probably unique in echinoids. 5. The areoles of the spatulate plastronal primary spines suggest a preferred stroke to the anterior resulting in a backward movement. In contrast, the plastronal spines of spatangoids are constructed for forward locomotion (Saucède et al. 2009). These are five examples that illustrate the diversity of spines in Calymne—as well as the difficulty of inferring their function. In general, Calymne shows several significant morphological features, which partly seem to be incompatible. This includes the high and rather narrow test, the anterior position of the peristome, the relatively high placement of the periproct on the posterior side, the presence of a marginal fasciole and a subanal tuft. Furthermore, the spines on the plastron apparently perform a reverse power stroke, and the frills of shovel-shaped spines above and below the periproct move in opposite directions. On the basis of these characteristics, Saucède et al. (2009) generated a hypothetical scenario: Calymne lives partly buried, with its subanal tuft and the fasciole slightly below the sea floor. The animal has to move forward to reach the sparsely distributed nutrients on the bottom of the deep sea. The antagonistic movements of the spines may cause a rocking movement of the tilted test with the mouth and its food-gathering phyllopodia being near the relatively nutrient-rich upper layer of the sea floor. Saucède et al. (2009) called this behavior “tilted ploughing”. The function of the conspicuous spines around the periproct is difficult to explain. The large shovels might help right overturned specimens, especially as the test of Calymne is rather high and unstable in water currents. However, these are hypotheses. Distribution: Calymne is known from two locations in the Northwest Atlantic: north of Bermuda and in Newfoundland Basin. In the Northeast Atlantic, it was found north/ west of the Amper Seamount off Portugal. Bathymetric range 3,720 to 4,860 m (Fig. 3.98).

Genus Sternopatagus de Meijere, 1903 The genus is monospecific. Sternopatagus sibogae de Meijere, 1903 Only a single specimen is known. Test: Medium size, in outline wider than long (length 64 mm, width 70 mm, height 53 mm), anterior side slightly concave; aboral side highly vaulted, upper part subconical, oral side flattened, deep furrow from beneath the anterior margin to the peristome; posterior end obliquely truncated below the high ambitus; plates exceedingly thin and fragile. Apical system: Slightly anterior with four gonopores; large ocular plates OII and OIV separating the anterior genital plates from the posterior. Ambulacra: Flush with the test, no petals, plates with single pores. Interambulacra: Aboral plates hardly larger than ambulacral plates; narrow labrum separated from the sternal plate by adjoining ambulacral plates. Oral side: Peristome anterior, vertical opening at the end of the deep furrow; rudimentary penicillate tube feet around the peristome. Periproct: On the truncated posterior end. Fasciole: Marginal fasciole running far below the rounded margin from below the periproct along the lateral oral side toward the anterior groove. Tubercles: Small, crenulate, and perforate, scattered all over the test. Spines: Short and simple. Occurrence: The specimen was collected in the Timor Sea, Indonesia, at a depth of 2,050 m (Fig. 3.99). 3.3.1.2 Family Corystusidae Foster & Philipp, 1978 There is only one genus with a single extant species. Genus Corystus Pomel, 1883 Synonym: Stereopneustes de Meijere, 1902 Corystus relictus (de Meijere, 1902) Test: Thick; outline oval with rounded anterior margin; in profile high domed, posterior end truncated; length may reach 90 mm. Apical system: Subcentral, elongated, four gonopores (see Fig. 3.97). Ambulacra: Subpetaloid, flush with the test, pores double in low plates, extending over the whole aboral side. Oral side: Flattened, sunken toward the peristome; few enlarged pores around the mouth; peristome anterior. Periproct: Inframarginal on the undercut posterior side; subanal fasciole present.

3.3.1 Order Holasteroida Durham & Melville, 1957 



A

E

C

B

 121

D

F O III G3 G2 O IV O II G4 G1 OV OI

G

H

I

Fig. 3.98: Calymne relicta. (A) Aboral side: in front of the small gonopores, a few large tubercles occur. (B) Posterior face. (C) Side view (shaded area is damaged). Sketches after photos in Saucède et al. (2009). (D) Posterior face slightly tilted anteriorly: below the fasciole lies a subanal tuft of long and slender spines. Photos courtesy of T. Saucède. (E) Apical system of adult specimen of 30 mm length. (F) Apical system of juvenile of 11 mm length without gonopores (genital plates in red, ocular plates in blue). Scale bar is 1 mm. (G) Plastron: interambulacral plates in green, labrum (1), sternum (2); ambulacral plates in yellow. Scale bar is 5 mm. (H, I) Juvenile specimens. (H) Side view of specimen of 4.5 mm length. Scale bar is 0.5 mm. (I) Side view of specimen of 11 mm length: the posterior bulge is more prominent than in adult specimens (interambulacral plates shaded, fasciole in black). Scale bar is 5 mm. Modified after Saucède et al. (2009).

Spines: Primary ones short and uniform, not enlarged on the aboral side. Color: In life, the echinoid is dark purplish, more rarely lighter brownish. The denuded test is white to light pink.

Distribution: This deep sea species is known in the Malay region and the China Sea from off Halmahera, Indonesia to southern Japan in depths of 250 to 900 m. Remarks: Corystus relictus (syn. Stereopneustes relictus de Meijere, 1902) was considered to be a member of the

122 

 3.3 Superorder Atelostomata von Zittel, 1879

B

A

C

D

E

family Holasteridae even though it was composed entirely of fossils. A.B. Smith included it into the Corystusidae, the family thus extending to modern days, and made it synonymous to Corystus (Fig. 3.100). 3.3.1.3 Family Carnarechinidae Mironov, 1993 Genus Carnarechinus Mironov, 1978 Carnarechinus clypeatus A. Agassiz, 1879 Status: Mooi & David (1996: p. 915) commented, “We do not here consider the taxonomically confused genus Carnarechinus Mironov, 1978 an urechinid. Despite recent additions to the literature on this genus (Mironov 1993), its affinities remain difficult to establish. We will also not consider Cystechinus clypeatus Agassiz, 1879 (the type species of the genus), a species known only from fragments collected by the Challenger. The systematic confusion surrounding these fragments (see Mortensen 1907 for a summary) means that we may never know whether they represent a single species, or several”. Meanwhile, Kroh & Mooi (2010) commented that “Despite the recent progress in echinoid systematics, its relation to other urechinids still remains unclear, but preliminary molecular data indicate that it is a basal member of the clade” (Kroh & Mooi 2010). 3.3.1.4 Family Plexechinidae Mooi & David, 1996 Genus Plexechinus A. Agassiz, 1898 Test: Elongated with thin plates.

Fig.3.99: Sternopatagus sibogae. No scale, modified after de Meijere (1903). (A) Apical system: (genital plates in red, ocular plates in blue, interambulacral plates in gray). (B) Oral plate pattern: the posterior interambulacrum 5 is interrupted between the labrum (1) and the sternum (2) by the posterior ambulacra (yellow). The marginal fasciole (darker gray) leads laterally around the oral side. (C–E) Test length 64 mm. Banda Sea. From de Meijere (1903). (C) Aboral side. (D) Oral side: there is a deep anterior furrow toward the subcentral peristome. (E) Side view: the test is subconically inflated.

Apical system: Genital plates G2 and G3 fused with one or two gonopores. Ambulacra: Apetaloid with only one pore per ambulacral plate except directly adjacent to the peristome. Oral side: Labrum and sternum interrupted by adjoining ambulacral plates. Subanal fasciole: Present. Seven nominal species are described: Plexechinus aoteanus (McKnight, 1974). Off South Island of New Zealand. Plexechinus cinctus A. Agassiz, 1898. Off southern California and south of the Gulf of California. Plexechinus hirsutus Mortensen, 1905. From the Denmark Strait and Iceland to the Caribbean. Plexechinus parvus (Mironov, 1978). Off Western Australia. Plexechinus planus (Mironov, 1978). Weddell Sea, Antarctic Sea and south of Tasmania. Plexechinus spectabilis Mortensen, 1948. Sulu Sea, Philippines. Plexechinus sulcatus David & Mooi, 2000. Southern Indian Ocean. Plexechinus aoteanus, P. parvus, and P. planus are most probably synonymous, according to Mooi & David (1996). They were unable to find distinguishing characteristics except for the geographical distribution (Tab. 3.3). The group is likely to represent a single circum-Antarctic species with populations that range into the Southern Pacific (Fig. 3.101). Literature: McKnight 1974; Mooi & David 1996; David & Mooi 1999.

3.3.1 Order Holasteroida Durham & Melville, 1957 

A

 123

B

D C

Fig. 3.100: Corystus relictus. (A–C) Test length 62 mm; Balicasag Island, Philippines. Photos courtesy of H. v. Noordenburg. (A) Aboral side: four gonopores open in the holasteroid apical system. The ambulacra are subpetaloid and flush with the test. (B) Oral side: the peristome is surrounded by a few larger pores. The subanal fasciole is distinct. (C) Side view: the subpetaloid rows of double pores reach down toward the margin, below which they become much smaller. (D) Test length 61 mm, Philippines, aboral side. Photo NHM Wien, courtesy of A. Kroh.

Plexechinus aoteanus (McKnight, 1974) (Synonymous with P. planus (Mironov, 1978); P. parvus (Mironov, 1978). Test: Lowly domed, in outline rounded anteriorly, attenuated toward the posterior end; usually up to 30 mm in length, exceptionally up to 40 mm. Apical system: Slightly anterior, stretched and not disrupted; one gonopore in fused G2/G3; G1 and G4 each perforated by a gonopore.

Oral side: In the main flat, sunken toward the peristome, sternum inflated and keeled; peristome anterior; interambulacrum 5 interrupted by a pair of ambulacral plates joining at the midline. Periproct: Supramarginal on the truncated posterior side, distinct subanal fasciole encircling the faintly developed anal rostrum. Primary tubercles: Scattered over the aboral side, on oral side much more numerous; dense coat of miliary tubercles.

124 

 3.3 Superorder Atelostomata von Zittel, 1879

Tab. 3.3: Species from the genus Plexechinus. Plexechinus aoteanus/ parvus/planus

Plexechinus cinctus

Plexechinus hirsutus

Plexechinus spectabilis

Plexechinus sulcatus

Apical system not disjunct

Apical system disjunct

Apical system disjunct

Apical system not disjunct

Apical system disjunct

Gonopores present in G1 and G4

Gonopores lacking in G1 and G4

Gonopores lacking in G1 and G4

Gonopores present in G1 and G4

Gonopores lacking in G1 and G4

One gonopore in fused G2/G3

Two gonopores in fused G2/G3

Two gonopores in fused G2/G3

Two gonopores in fused G2/G3

Two gonopores in fused G2/G3

A

E

C

B

F

D

G

H

Fig. 3.101: (A–D) Plexechinus; plate pattern of apical systems. Scale bars are 4 mm (A–C) and 5 mm (D). (A) Plexechinus cinctus: the system is twice interrupted by interambulacral plates; the anterior fused genital plate G2/G3 is perforated by two gonopores, the posterior genital plates G1 and G4 are not perforated. (B) Plexechinus aoteanus/parvus/planus: the apical plates are just contiguous or just not, in G2/G3, only one genital pore is developed, G1 and G4 are each perforated by a gonopore. (C) Plexechinus spectabilis: the apical system is compact, in G2/G3, two genital pores are present and G1 and G4 are also perforated by gonopores. Modified after Mooi & David (1996). (D) Plexechinus sulcatus: the apical system is triply disjunct. The anterior genital plate is perforated by two gonopores, the posterior gonopores are lacking. After David & Mooi (2000). The genital plates are shaded in red, the ocular plates in blue, interambulacral plates in gray. (E–H) Plexechinus aoteanus/parvus/planus. Test length 29 mm; Weddell Sea, Antarctica. ZMH. (E) Aboral side with three gonopores. (F) Oral side. (G) Side view: the plastron is strongly inflated. (H) Specimen with spines. Test length 33 mm; Weddell Sea, Antarctic. ZMH. Aboral side. It is covered by a dense coat of miliary spines and scattered primary spines.



Spines: Short and very fine. Color: The test is dark brown with a purplish tint, primary spines whitish. Distribution: If the three species turn out to be a single one, this one occurs around Antarctica and extends into the Southern Indian and Pacific Ocean. Bathymetric range: 946 to 2,350 m. Plexechinus cinctus A. Agassiz, 1898 Test: Elongated ovoid, upper side lowly domed, highest point just in front of the periproct, maximum length not much more than 20 mm. Apical system: Three times disrupted, the fused genital plate G2/G3 is perforated by two gonopores, in both the posterior genital plates, G1 and G4 gonopores are absent. Oral side: Peristome anterior, round, slightly sunken; labral and sternal plates interrupted, sternum inflated and distinctly keeled. Periproct: Supramarginal, transversely elliptical, positioned in a shallow depression above the broad anal rostrum; subanal fasciole encircling the rostrum very conspicuously. Primary tubercles: Small, covering the test except on the posterior ambulacra on the oral side and flanking the rostrum, coat of minute miliaries. Distribution: This species is known only from off South California and south of the Gulf of California at a depth of 1,235 to 1,390 m (Fig. 3.102). Plexechinus hirsutus Mortensen, 1905 Test: Regularly ovoid, although the anterior side slope straightly or even a little concave; upper side slightly vaulted, anteriorly rounded, posterior side forming a hood above the anal rostrum; maximum length 20 mm. Apical system: Disjunct, two gonopores in fused genital plates G2/G3; posterior gonopores lacking. Ambulacra: Single-pored except near the peristome, where a rudimentary phyllode is developed. Oral side: Peristome sunken, at the end of a shallow depression extending from the anterior margin, round; labral and sternal plate separated, sternum inflated and keeled. Periproct: High on the posterior side below a small hood; conspicuous subanal fasciole encircling the anal rostrum. Primary spines: Approximately 3 mm in length, slender and curved; those on the sternum widened at the point; miliary spines approximately 0.5 mm. Distribution: This species is reported in the North and East Atlantic from the Denmark Strait between Iceland and Greenland to the Caribbean Sea, at depths of 658 to 2,375 m. Plexechinus spectabilis Mortensen, 1948 Test: Rather high and ovoid in outline, distinctly tapering toward the posterior end forming a rostrum; in profile, the

3.3.1 Order Holasteroida Durham & Melville, 1957 

 125

aboral side is lowly vaulted, anteriorly rounded, at the rear sloping steeply toward the fairly conspicuous anal snout. Apical system: Compact with four gonopores. Oral side: Sunken toward the subcircular peristome, labrum and sternum interrupted, plastron inflated and keeled. Periproct: On the posterior side above the rostrum; distinct subanal fasciole. Primary tubercles: Scattered between the coat of miliaries, aborally more sparsely distributed. Primary spines: Not exceeding 3 mm. Color: Light brownish to whitish. Occurrence: Known only from the Sulu Sea, Philippines, at a depth of 4,163 m. Plexechinus sulcatus David & Mooi, 2000 Only two more or less fragmented specimens are known, length approximately 12 and 35 mm. Test: Elongated, anterior side with a distinct sulcus diverging toward the ambitus, posterior side rounded, not attenuated, sides almost parallel; upper side lowly domed. Apical system: Triply disjunct with two gonopores in the fused anterior genital plate. Oral side: IA 5 discontinuous, sternum separated from labrum by one pair of adjoining ambulacral plates. Periproct: Supramarginal above a slight posterior rostrum; anal fasciole present. Color: Probably violet-brown with purplish overtones, mostly on the oral side and at the spine bases, phyllopodia almost black.

3.3.1.5 Family Pourtalesiidae A. Agassiz, 1881 The family contains species with very strangely transformed architectures. The typical features for all species are a deep peristomial groove, a disjunct apical system, and on the oral side, an interrupted interambulacrum 5. The strange shapes are not expressed in juveniles, but become progressively more obvious during ontogeny. Apical system: A single anterior genital plate with one or two gonopores is followed by two ocular plates each with a gonopore; the posterior genital plates are separated and without gonopores; behind them, the posterior oculars in Pourtalesia, again separated.

Plastron: The plastron is orthosternal: that means, it is composed of the labral plate adjacent to the peristome, a single sternal plate and then a pair of episternal plates. Usually, the labrum is separated from the sternum by one or two pairs of ambulacral plates that are in contact on the midline.

As deep-sea dwellers, the pourtalesiids are welladapted to their extreme habitat. They feed by ploughing through the uppermost layer of the substrate, where the detritus, the “sea snow”, is deposed. When the animal

126 

 3.3 Superorder Atelostomata von Zittel, 1879

E

C

A B

D

F

G

H

I

J

Fig. 3.102: (A–D) Plexechinus cinctus. Test length 20 mm, off South California. From Agassiz (1904). (A) Aboral side: the periproct is situated on the aboral side above the rostrum. (B) Oral side: the plates in the posterior ambulacra are predominately naked. (C) Side view: the posterior end forms a distinct rostrum surrounded by the subanal fasciole. (D) Posterior side: the subanal fasciole is broad and high. (E–G) Plexechinus hirsutus. Test length 17 mm, off the Bermuda Islands. ZMUC. (E) Aboral side. (F) Oral side: the subanal fasciole is well-developed. (G) Side view: the posterior end forms a broad rostrum, surrounded by the subanal fasciole. (H–J) Plexechinus sulcatus. Test length 35 mm; northwest of the Kerguelen Islands. Reconstruction of the general appearance by Mooi on the base of large fragments. Modified after David & Mooi (2000).

moves forward, the sediment is pressed into the anterior groove toward the vertical mouth opening. Some of the species live epifaunally, some are adapted to a more endofaunal behavior having evolved a more elongated test with keels on the aboral and oral side, wide subanal fascioles and spatulate spines. It is likely that the anterior upper side lies free, whereas the posterior end with the rostrum is

covered by the substrate, and the fasciole maintains a water current to build a funnel toward the surface. The animals leave deep furrow-shaped moving traces (Mironov 2008). All species are very thin-shelled, living in deeper water, in the bathyal or even in the abyssal below 4,000 m. It is interesting that in the Cretaceous, echinoids with successively elongated necks had already evolved belonging

3.3.1 Order Holasteroida Durham & Melville, 1957 

to the extinct family Cardiasteridae Lambert, 1917. The most modified test existed in Hagenowia rostrata Forbes, 1852. The animal ploughed vertically through the fine chalk with the strongly rostrate apex at the sediment-water interface. The top was protected by a fan of long spines and only a small hole marked the position of the echinoid in the bottom. The food particles in the uppermost layer were transported downward

F

to the peristome in the deep anterior sulcus, isolated from the surrounding sand by a dense arch of spines. This lifestyle resembles that of spatangoids, e.g., in the Recent genera Moira or Echinocardium. However, the strange shape was similar to the modern holasteroids, for example, Pourtalesia. The genus was restricted to the Upper Cretaceous of Western Europe; approximately 80 to 65 million years ago (Fig. 3.103).

B

A

C

 127

E

D

E

F

Fig. 3.103: (A, B) Fossil Hagenowia rostrata. Upper Cretaceous of England and Germany. (A) In life, the test was densely set with spatulate spines for excavation, whereas long sharp spines arose at the apex to deter possible predators. The anterior sulcus was covered by arched spines and the posterior end was set by a tuft for clearing the waste material. Height of test approximately 20 mm. (B) There is a long anterior rostrum (“neck”) the apical system lying at the top. A deep sulcus along the front side leads to the vertical peristome. After Gale & Smith (1982). (C–E) Pourtalesia jeffreysi. After Lovén (1883). (C) Aboral side. (D) Side view. (E) Oral side. (F) The trend goes from Echinocrepis on the sediment over Helgocystis to Echinogutta. After Mironov (2008).

128 

 3.3 Superorder Atelostomata von Zittel, 1879

The family Pourtalesiidae consists of eight genera: Ceratophysa Pomel, 1883. Monospecific; Antarctic and subantarctic. Cystocrepis Mortensen, 1907. Monospecific; pacific Panama. Echinocrepis A. Agassiz, 1879. Two species; Antarctic, Northern Pacific. Echinosigra Mortensen, 1907. Two species, Atlantic, Pacific and Antarctic. Helgocystis Mortensen, 1907. Monospecific; Antarctic and subantarctic. Pourtalesia A. Agassiz, 1869. Eight or more species, worldwide. Solenocystis Mironov, 2008. Monospecific, Northern Atlantic. Spatagocystis A. Agassiz, 1879. Monospecific; Antarctic. Literature: Lovén 1883; Mortensen 1907, 1950; Koehler 1926; Mironov 1973, 1974, 1978, 1995, 1996, 1997, 2008; Gale & Smith 1982; David 1983, 1984, 1987; 1990; Gage 1984; David & Mooi 1999; David et al. 2003; Saucède et al. 2003. Genus Ceratophysa Pomel, 1883 The genus is monospecific. Ceratophysa ceratopyga (A. Agassiz, 1879) Test: Large, outline triangular, anteriorly very broad with a wide, shallow frontal sulcus, sides slightly concave, gradually narrowing toward the posterior end with distinctly differentiated subanal rostrum; aboral side lowly arched, median keel; on the posterior side sloping steeply toward the periproct situated in a notch in front of the slightly inflated rostrum; in length approximately 100 mm, but fragments suggest a length of up to 200 mm. Apical system: Anterior; four genital pores; single fused anterior plate with two gonopores followed by two ocular plates with gonopores; disrupted from the posterior ocular and genital plates. Ambulacra: All similar, uniporous with minute pores. Oral side: Flat with a short peristomial groove, peristome small, facing forward; plate pattern partly unknown; labrum large, probably separated from the sternum by one pair of ambulacral plates. Periproct: In a deep notch between posterior end and subanal rostrum, subanal fasciole broad. Spines: Small, forming a dense, uniform coat. Color: Dark red brown to light violet. Distribution: Antarctic Ocean south of the Indian Ocean and off Chile as far north as Valparaiso; bathymetric range 3,165 to 4,030 m (Fig. 3.104). Genus Cystocrepis Mortensen, 1907 The genus is monospecific

Cystocrepis setigera (A. Agassiz, 1898) Test: Thin-plated; in outline elongated-oval with a slightly pointed posterior end; in profile on the underside concave, upper side pyramidal, anterior slope distinctly steeper than the posterior. Apical system: Anterior; four gonopores. Ambulacra: Microscopic single pores. Oral side: Peristome subcircular, vertical at the end of a peristomial groove; very small labral plate hidden in the groove and separated from the sternal plate by several ambulacral plates. Periproct: Inframarginal below the posterior hook; no subanal rostrum and no fasciole. Spines: Rather long and scattered over the aboral side; denser distributed on the oral side. Color: Dark brown. Occurrence: The species is known only from the East Pacific off the region of Panama to Peru at depths of 2,500 to 3,440 m. Biology: Cystocrepis setigera lives ploughing slowly through the sea bottom often grouped in clusters. Numerous specimens could be observed by video recordings and collected during a submersible survey of the Peru Trench in 1999 at a depth of 2,500 m. On their aboral side, many organisms were found being transported, sheltered, or parasitizing. On an individual of 10 mm, up to three holothurians were attached at the lateral sides. As the same kind of sea cucumbers were not observed on the sea floor, the setting seems to be durable. Furthermore, ophiuroids, copepods, amphipods, cirripedes, and other small organisms were found between and on the primary spines. Usually, the euechinoids have a defense mechanism against fouling and settling as the whole test and all appendages are entirely covered by epidermis. Astonishingly, in Cystocrepis, neither the spines nor the test seemed to be damaged. Obviously, these pourtalesiids represent islets of hard ground on a very soft, muddy sea bottom being a suitable habitat for many ectosymbionts. The different animals constitute a functioning community (David et al. 2003). Genus Echinocrepis A. Agassiz, 1879 Echinocrepis cuneata A. Agassiz, 1879 Test: Large, irregularly triangular in outline, narrowing toward the slightly pointed posterior end; narrow, deep frontal notch; relatively high in profile, pyramidal, anterior slope steeper than posterior. Apical system: Three or four gonopores, anterior and posterior part not disrupted, but juxtaposed. Ambulacra: With microscopic pores. Oral side: Concave, peristome vertical at the end of the narrow peristomial groove; labral plate long and narrow,

3.3.1 Order Holasteroida Durham & Melville, 1957 



D

B

A

C

FF

E

G I

H

K

 129

L

J

N M

O P

Fig. 3.104: (A–E) Ceratophysa ceratopyga. (A) Oral plate pattern of young specimen, length 20 mm. The large, triangular labral plate (1) is separated from the sternal plate (2) by a single pair (probably!) of ambulacral plates. Modified after Mironov (2008). (B–E) Specimen with spines, after Agassiz (1881). (B) Aboral side: the test is broadest anteriorly. (C) Side view: a very prominent rostrum is developed at the posterior end. (D) Posterior side. (E) Anterior side: the deep anterior funnel ends in a vertical mouth. (F–I) Cystocrepis setigera. Test length 89 mm; Challenger Station 3399. From Agassiz (1904). (F) Aboral side. (G) Oral side. (H) Side view. (I) Test length 80 mm. Oral plate pattern: the very small labral plate is usually hidden in the peristomial funnel. It is separated from the sternal plate (2) by two pairs of ambulacral plates, which are positioned one in front of the other along the axis (posterior ambulacra AI and AV in yellow, interambulacrum IA 5 in green). Modified after David (1984). (J–N) Echinocrepis cuneata. (J) Oral plate pattern. Test length 90 mm. The elongated labral plate (1) is separated from the sternal plate (2) by one pair of ambulacral plates in each column. Posterior ambulacra AI and AV in yellow, interambulacrum IA 5 in green. Modified after David (1984). (K–N) Specimens with spines. Test length of both subjects 88 mm. From Agassiz (1881). (K) Aboral side. (L) Posterior side. (M) Side view. (N) Oral side. (O–P) Echinocrepis rostrata. Test length 86.5 mm, Kamchatka Peninsula. (O) Aboral side. (P) Profile. There is a small but distinct rostrum below the depressed periproct. Sketches after the photos in Mironov (1973). Two other specimens do not show this extreme shape, they are narrower anteriorly and the vertex is less pointed.

130 

 3.3 Superorder Atelostomata von Zittel, 1879

sternal plate small and very narrow, both separated by a single pair of ambulacral plates; distinct keel from peristome to periproct. Periproct: Inframarginal below the posterior hook; subanal rostrum and fasciole absent. Spines: Small and scattered over aboral side. Color: Dark violet-brown. Occurrence: The two specimens came from the Antarctic Ocean, near the Crozet Island, found at depths of between 2,925 and 4,616 m. Echinocrepis rostrata Mironov, 1973 Very much like E. cuneata with the large conical test and the two gonopores. Occurrence: North Pacific, from Kamchatka Peninsula to Northwest of the Channel Islands, California, USA; depth 3,315 to 5,020 m. Genus Echinosigra Mortensen, 1907 The test of Echinosigra has a very characteristic shape. It is formed like a bottle and can be differentiated into a “head” with a deep funnel-like peristomial groove, a more or less elongated, slender “neck”, a thickened “body”, and a rostrate “tail”. Sexual dimorphisms occur: in the female, the large gonopores are set with short papillae; in the male, there are small gonopores with long papillae. Two subgenera, separated by the geographical range, and several species and subspecies are established in this genus: Echinosigra (Echinogutta) Mironov, 1974. South Atlantic, Antarctic; Pacific. Four gonopores. Echinosigra (Echinosigra) Thomson, 1873. Northeast Atlantic. Two gonopores. The North Atlantic species Echinosigra paradoxa, established by Mortensen (1905), was synonymized with Echinosigra phiale by Mironov (1974). This conclusion was confirmed by Gage (1984) on the basis of biometrical measurements of different ontogenetic stages: E. paradoxa is the adult stage of E. phiale. Originally, the Antarctic specimens of Echinosigra were thought to belong to the same species (E. phiale) with a wide geographic range from Antarctica to Iceland. Mironov (1974) stated that there were two different species, separating the Antarctic species E. amphora. Mironov (1974, 1997) differentiated some more subspecies, species, and a subgenus of Echinosigra (i.e., Echinogutta). The shape of the specimens ranges between a very slender “body” with a long “neck” and a compact “body” with almost no “neck”. However, as all intermediates occur, it is very difficult to delimit clearly distinguished morphotypes of the new taxa

(David & Mooi 2000; David, personal communication 2008). Because of this confusion, I can only add a summary of their distribution: specimens of the genus Echinosigra were found in the Northern Atlantic and Pacific, in the Antarctic Sea extending into the Southern Atlantic, the Indian Ocean, south of Tasmania and New Zealand, and in the Indo-West Pacific. Bathymetric range: 1,026 to 7,170 m (Mironov 1997). Echinosigra (Echinogutta) antarctica Mironov, 1974 Echinosigra (Echinogutta) fabrefacta Mironov, 1974 Echinosigra (Echinogutta) valvaedentata Mironov, 1974 Echinosigra (Echinogutta) amphora Mironov, 1974 Test: Compact with conspicuous keels along the median line on the aboral and oral side; compressed flanks; prominent posterior rostrum; maximum length may reach 45 mm. Apical system: Very anteriorly; four gonopores; once divided. Oral side: Convex; deep and wide triangular funnel toward the vertical peristome; labral plate: narrow and elongated, separated from the sternal plate by one pair of ambulacral plates in each column. Periproct: Supramarginal just above the rostrum; subanal fasciole surrounding the rostrum. Color: Yellowish-pink, spines darker. Echinosigra (Echinosigra) phiale Thomson, 1873 Test: Very elongated, compressed, distinctly keeled above and below, frontal end functions as roof for deep oral invagination toward the peristome; length may reach 37 mm. Apical system: Very anterior, two gonopores perforating the anterior genital plate, followed by two ocular plates; posterior genitals and oculars separated and far removed to the posterior. Ambulacra: With minute simple pores. Oral side: Peristome subcircular, vertical, at the end of the funnel-like groove; labrum and sternum separated by one pair of very stretched ambulacral plates. Periproct: Supramarginal, above the distinct subanal rostrum; broad subanal fasciole encircling the rostrum. Spines: Small and scattered aborally, more densely distributed along the keels. Distribution: This species is recorded with certainty from the Northeast Atlantic at 1,540 to 3,610 m (Fig. 3.105). Echinosigra (Echinosigra) porrecta Mironov, 1997 Echinosigra (Echinosigra) vityazi Mironov, 1997



Genus Helgocystis Mortensen, 1907 Helgocystis carinata (A. Agassiz, 1879) Test: Subcylindrical, gibbous, with a strongly developed posterior rostrum, length up to 90 to 100 mm. Apical system: Four gonopores: single anterior plate (G2) perforated by two gonopores, following ocular plates OII and OIV each with a gonopore; small posterior genital plates in contact with OI and OV. Ambulacra: With single pores except close to the peristome, where they are double. Oral side: Peristome vertical at the end of deep, funnellike invagination; labrum (1) large, triangular and separated from the rectangular sternum (2) by one pair of ambulacral plates in each column. Periproct: Supramarginal above the very prominent subanal rostrum and below a slight hood. Subanal fasciole: Narrow, encircling the end of the rostrum. Tubercles: Small and scattered aborally. Distribution: Antarctic and Pacific subantarctic waters, at depths of 2,950 to 4,820 m (Fig. 3.106). Genus Pourtalesia A. Agassiz, 1869 The genus is characterized by its relatively compact test with a more or less distinctly differentiated rostrum. In young specimens, the rostrum is usually more prominent than in adults. Test: Flanks more or less compressed, oral side convex. Apical system: Four gonopores, disrupted into three parts. Oral side: Interambulacrum 5 interrupted: labrum and sternum are separated by up to four asymmetrical pairs of ambulacral plates, one plate from AI to AV, respectively, meeting along the midline; peristome vertical at the end of the deep oral groove. Periproct: On the posterior side above the rostrum, more or less overarched by a hook. Eight species are included in this genus: Pourtalesia alcocki Koehler, 1914. Western Indian Ocean. Pourtalesia aurorae Koehler, 1926. Antarctica. Pourtalesia debilis Koehler, 1926. Antarctica. Pourtalesia heptneri Mironov 1978. Banda Trench, Indo­nesia. Pourtalesia hispida A. Agassiz, 1879; Antarctica. Pourtalesia jeffreysi Thomson, 1873. Northern Atlantic. Pourtalesia laguncula A. Agassiz, 1879. Japan Sea and Malaysia. Pourtalesia miranda A. Agassiz, 1869. Northern Atlantic. Pourtalesia tanneri A. Agassiz 1898. East Pacific.

3.3.1 Order Holasteroida Durham & Melville, 1957 

 131

Mironov (1997) listed some more species, but all “species” seemed to range within the width of variation of the existing species. David (1984, 1987) followed the plate architecture during ontogeny and found out that, on the oral side, the ambulacra AI and AV are not interrupted by interambulacral plates—in contradiction to the interpretations of Lovén (1883) and Mortensen (1950c). Instead, AI and AV are in contact at the midline, forming up to four asymmetrical pairs of plates. This ground pattern displays much variation being an area of strong plate movement during ontogeny. Pourtalesia alcocki Koehler, 1914 Main characteristics of the genus: with its subanal rostrum bent upward, this species is very similar to Pourtalesia laguncula; differing in its larger size, up to 50 mm, and by the more prominent plastronal keel on the oral side. Distribution: Western Indian Ocean from the Gulf of Oman to off South Africa, at 1,450 to 2,380 m (Fig. 3.107). Pourtalesia aurorae Koehler, 1926 Main characteristics of the genus: maximum length 57 mm. Test: rather broad, width approximately 65% of length; sides strongly convex, anterior end straight; posterior end with a small conical rostrum encircled by a broad subanal fasciole; oral side flat, without a plastronal keel. Distribution: Antarctic (Indian Ocean), 442 to 1,600 m. Pourtalesia debilis Koehler, 1926 Main characteristics of the genus: maximum length of recorded specimens 26 mm, but still lacking gonopores. Test width less than 50% of length, sides parallel, anterior surface convex; posterior end with a conspicuous, rounded rostrum encircled by a subanal fasciole; primary spines smooth, arranged in short rows along the sides of the test. Distribution: Circum-Antarctic, 192 to 6,290 m. Pourtalesia heptneri Mironov, 1978 The species is likely to be the most deep-sea irregular echinoid found at depths between 7,130 and 7,340 m in the Banda Trench in the Banda Sea, Indonesia, during the voyages of the RV Vityaz. It is characterized by marked individual variation. Pourtalesia hispida A. Agassiz, 1879 Main characteristics of the genus: length up to 60 mm. Test: Very thin, almost transparent; aboral side highly domed, oral side flat; anterior side straight, sides almost

132 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

C

D

E

F

G

H

I

J

K

16.6 mm

L

N

M

20.7 mm

>45 mm

>50 mm



parallel; posterior end pointed toward a small, conical subanal rostrum encircled by a subanal fasciole; primary spines distinctly serrate, arranged in more or less horizontal rows. Distribution: The species lives around the Antarctic at depths of 1,153 to 3,634 m (Fig. 3.108). Pourtalesia jeffreysi Thomson, 1873 Main characteristics of the genus: maximum length is 58 mm. The general shape of the test rather variable; anterior side straight, flanks and underside convex; posterior side tapering to a conspicuous, rounded anal rostrum, encircled by a subanal fasciole; aboral side lowly domed at the posterior end forming a distinct hood overarching the periproct. Labrum: Shows much variation, it may lie within the peristomial invagination, or on the incurved edge, and sometimes as a very narrow plate visible between the ambulacral plates. Primary spines: Of uniform length, about one-third of test length, slightly curved. Distribution: This species is widely spread in the North Atlantic from the Faroer Channel northward to the Norwegian Sea, the Kara Sea, and Baffin Sea at depths of 50  to 2,450 m. It is likely that its bathymetric distribution  is the consequence of temperature and not depth (Fig. 3.109). Pourtalesia laguncula A. Agassiz, 1879 Main characteristics of the genus: small form, maximum length 30 mm. This species differs from the other by its broad subanal rostrum, which is conspicuously bent upwards.

3.3.1 Order Holasteroida Durham & Melville, 1957 

 133

Distribution: The species is rather common in the Japanese Sea, and is also reported from the Malayan ­ region. Bathymetric range: approximately 220 to 1,370 m. Pourtalesia miranda A. Agassiz, 1869 Main characteristics of the genus: length may reach 40 mm. Anterior side of test straight, sides almost parallel, tapering to a more or less differentiated subanal rostrum surrounded by a fasciole; aboral side lowly domed, forming a slight hook toward the posterior side above the periproct; oral side convex; primary tubercles in more or less horizontal, short series. Distribution: This species is recorded from relatively warm waters in the Northern Atlantic, from the Caribbean and the Bay of Biscay to the Davis Strait and south of Iceland at depths of 450 to 5,850 m (Fig. 3.110). Pourtalesia tanneri A. Agassiz, 1898 Main characteristics of the genus: small form, not exceeding 20 mm in length. Distribution: The species occurs off the west coast of America from the Gulf of California and the Galapagos Island to Chile; bathymetric range 1,450 to 2,389 m. Genus Rictocystis Mironov, 1996 The genus is monospecific. Rictocystis jensenae Mironov, 1996 Test: Thin-plated; subcylindrical, posterior rostrum strongly developed. Apical system: Compact or ethmophract; four large gonopores; madreporic pores not reaching between posterior genital plates; posterior ocular plates in contact.

◂ Fig. 3.105: (A–E) Echinosigra amphora. Test length 43 mm, Antarctic. (A) Aboral side: the supramarginal periproct is partly concealed by the posterior slope. (B) Oral side: the labral (1) and sternal plates (2) are separated. Scale bar is 5 mm. Modified after Mironov (1974), as E. (E.) amphora fabrefacta. Ambulacral plates yellow, interambulacral plates green. (C–E) Test length 18 mm; Challenger Station 156, after Agassiz (1881) as E. phiale. (C) Aboral side: there is a distinct keel from the anterior side to the posterior slope. (D) Oral side: a median keel runs from the peristomial funnel to the end of the posterior rostrum. (E) Side view. (F, G) E. phiale. Test length 40 mm, Bay of Biscay. Photos by A. Godon; courtesy of B. David, Biogéosciences Dijon. (F) Aboral side. (G) Oral side. The apical system with the two gonopores is positioned at the extreme anterior edge. There is a distinct keel along the aboral and oral midline. (H–J) E. phiale. Changes of oral plate pattern during ontogeny. (H) Specimen of approximately 6 mm. This early stage closely resembles that of Pourtalesia before its plates become more and more stretched. (I) Specimen of approximately 13 mm. (J) Specimen of approximately 32 mm. The labral plate (1) is separated from the sternal plate (2) by one symmetrical pair of plates, respectively, one plate of the posterior ambulacra AI and AV. There are distinct intraplate lines on the very elongated central ambulacral plates (not to scale). Oral ambulacral plates in AI and AV are yellowish, interambulacrum IA 5 is green. Modified after Gage (1984). (K–N) Development of the apical system during ontogeny. (K) The plates form a continuous, compact disc, the anterior genital plate having only a single hydropore. (L) The anterior genital plate begins to differentiate into two plates, G2 and G3. (M) With the opening of the gonopores, G3 forms a distinct plate with a second gonopore. (N) The posterior part of the apical system, G1 and G4, plus ocular plates OI and OV, shifted to the posterior. The anterior part, containing ocular plate OIII, the two perforated anterior genital plates G2 and G3, and the two ocular plates OII and OIV are now separated from the posterior part. This posterior part being continuous distinguishes Echinosigra from Pourtalesia, in which the posterior genital plates G1 and G4 are also disrupted from the posterior oculars OI and OV. Ocular plates bluish, genital plates reddish, interambulacra gray. Modified after Saucède et al. (2003).

134 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

C

D

H

G F

E

3.1 mm

J

I

3.82 mm

7.1 mm

K

7.78 mm

21.88 mm

8.5 mm

17.8 mm

3.3.1 Order Holasteroida Durham & Melville, 1957 

 135

B

A

C

G

D

E

H I

F

Fig. 3.107: (A, B) Pourtalesia alcocki. Test length approximately 35 mm, Mauritius. Photo A. Godon; courtesy of B. David, Biogéosciences, Dijon. (A) Aboral side. (B) Side view. A distinct keel runs along the upper side continuing toward a hook above the periproct. (C–E) Pourtalesia aurorae. (C) Aboral side. (D) Side view: the highest point is near the truncated anterior side. Sketches after photos in Koehler (1926), 44 to 57 mm. (E) Oral side. Sketch after a photo in David et al. (2000), no size. (F,G) Pourtalesia debilis. Test length 26 mm. (F) Aboral side. The flanks are parallel. (G) Side view. Sketches after photos in Koehler (1926). (H, I) Pourtalesia heptneri. Scale bar is 10 mm. Modified after Mironov (1978). ◂ Fig. 3.106: (A) Helgocystis carinata. Oral plate pattern. Modified after a reconstruction by Mironov (2008) based on four fragments. Intraplate lines are present. (B–D) Helgocystis carinata. Approximately 95 mm. From Agassiz (1881). (B) Aboral side. (C) Side view. (D) Part of oral side with peristomial funnel and triangular labral plate. (E–H) Pourtalesia miranda. Development of the apical system during ontogeny. (E) The apical plates are continuous. There is a single anterior genital plate with a single hydropore. (F) The posterior part of the apical system, G1 and G4, plus ocular plates OI and OV, shifted to the posterior and become separated from the anterior part, containing OIII, G2/G3, and OII + OIV. (G) The posterior ocular plates become disrupted from the posterior genital plates and from each other. (H) The ocular plates OII and OIV turn out to bear the third and fourth gonopores. The triply disjunct apical system distinguishes Pourtalesia from Echinosigra, in which it is only once interrupted. Ocular plates bluish, genital plates reddish, interambulacra gray. Modified after Saucède et al. (2003). (I–K) Pourtalesia: Development of the oral plate pattern during ontogeny: (I) The labral plate 1 is in contact with the sternal plate 2. The pairs of ambulacral plates are positioned one beside the other in each column. (J) The ambulacral plates in columns I to V have, respectively, shifted to the median axis pushing the labral plate 1 anteriorly. (K) In ambulacrum I as well as in ambulacrum V, the plates have shifted to a position in which one is in front of the other along the axis. The labral plate is positioned in the peristome. Labral plate dark green, sternal plate lighter green, episternals still lighter, ambulacra yellow. Modified after David (1987).

136 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

C

D

Fig. 3.108: (A–C) Pourtalesia hispida. Test length 23 mm; Weddell Sea, Antarctic. AWI. (A) Aboral side: the periproct is supramarginal. (B) Oral side: the peristomial funnel is deeply invaginated. (C) Side view: the primary tubercles are arranged in short series. (D) Test with spines, length 42 mm; Weddell Sea, Antarctic. ZMH. Side view: the subanal rostrum is distinctly smaller than in the younger specimen.

3.3.1 Order Holasteroida Durham & Melville, 1957 



A

B

D

 137

C

E

Fig. 3.109: (A,B,D) Pourtalesia jeffreysi. Test length 39 mm; south of Jan Mayen Island. Northeast Atlantic. ZMUC. (A) Aboral side. (B) Oral side. (D) Side view: the hook overarching the periproct on the posterior side and the subanal rostrum is strongly developed. (C, E) Pourtalesia laguncula. Test length 19 mm; Sagami Bay, Japan. ZMUC. (C) Aboral side. (E) Side view: the broad subanal rostrum is typically bent upwards, encircled by the fasciole.

Ambulacra: All similar; single(?) pores close to the peristome; otherwise, only tiny pores. Oral side: Deep funnel from anterior side toward peristome; mouth itself subcircular(?) and facing vertically into a groove; labral plate extremely elongated, other plates insufficiently known. Periproct: Supramarginal, overarched by interambulacral plates. Tubercles: Small, scattered over aboral side. Distribution: Indian Ocean and West Pacific; bathymetric range 3,070 to 4,920 m. Genus Solenocystis Mironov, 2008 The genus is monospecific.

Solenocystis imitans Mironov, 2008 Test: Elongated, bottle-shaped, slightly widening toward the posterior and ending in a thick anal rostrum; aboral surface with a distinct keel along the midline. Apical system: Very anterior, compact, with four gonopores, two in the anterior fused genital plate, the posterior ones perforating the anterior ocular plates OII and OIV; the posterior genital plates carry no gonopores, being in contact with the posterior ocular plates (the interpretation of the “semifused” plates drawn by Mironov is not quite clear; in particular, the anterior ocular plate is only indicated by two small pores). No genital papillae. Ambulacra: Minute simple pores.

138 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

C

D

G

E F

Fig. 3.110: (A–C) Pourtalesia miranda. Test length 31 mm, west of Southern Greenland. ZMUC ECH 397 (as P. wandeli). (A) Aboral side. (B) Oral side. (C) Side view: a slight hook overarches the periproct above the prominent subanal rostrum. (D–G) Pourtalesia tanneri. Test length 17 mm; south of the Gulf of California. From Agassiz (1904). (D) Aboral side. (E) Oral side. (F) Side view. (G) Frontal view, looking into the peristomial groove.



Oral surface: With a distinct median keel, deep peristomial groove toward the mouth; labrum large, separated from the sternum by two symmetrical pairs of ambulacral plates set with intraplate lines. Periproct: Deeply sunken just above the posterior rostrum, which is encircled by a wide subanal fasciole. Tuberculation: Oral keel densely set with large tubercles, the rest of the test rather regularly covered by scattered smaller tubercles among a coat of dense, fine granulation. Distribution: Northern Atlantic, depth range 2,520 to 2,555 (Fig. 3.111). Genus Spatagocystis A. Agassiz, 1879 The genus is monospecific. Spatagocystis challengeri A. Agassiz, 1879 Test: Very fragile, plates thin like paper, outline ovoid; anterior side rounded with shallow indentation, posterior end tapered; upper side lowly vaulted with rounded margins, posterior side overhanging like a hood; length up to 75 mm. Apical system: Slightly anterior; three or four gonopores: single anterior genital plate perforated by one or two gonopores, posterior gonopores in OII and OIV. Ambulacra: With minute single pores. Oral side: Concave; peristome circular, opening vertically into a narrow and deep groove, labrum small, but visible, separated from the very small sternal plate by two pairs of ambulacral plates; following episternal and subanal plates strongly reduced; plastronal keel very distinct continuing as a small anal heel, more prominent in younger specimens than in adults; subanal fasciole also more developed in younger specimens; periproct oral (in adults), depressed above the heel. Tuberculation: Small, uniform, scattered aborally, more densely distributed on oral side, larger only along the oral keel. Color: Mauve. Distribution: The species is recorded from the Southern Indian Ocean and south of New Zealand; bathymetric range 2,950 to 3,607 m.

3.3.1.6 Family Urechinidae Duncan, 1889 The family is characterized by a thin-plated test, an orthosternal plastron with labrum and sternum not disjunct, or only very exceptionally. The apical system is always compact, the anterior genital plate bearing one or two gonopores. The ambulacra are apetaloid with single pores except in phyllodes. The periproct is inframarginal

3.3.1 Order Holasteroida Durham & Melville, 1957 

 139

to supramarginal; in some species, a subanal fasciole is present. The family consists of four genera: Antrechinus Mooi & David, 1996. Three species: sexual dimorphism. North and South Pacific. Cystechinus A. Agassiz, 1879. Three species. West coast of America; subantarctic and Antarctic waters. Pilematechinus A. Agassiz, 1904. Three species. Caribbean, west coast of America, Antarctica. Urechinus A. Agassiz, 1879. Three species. Antarctic waters. Genus Antrechinus Mooi & David, 1996 The genus includes three marsupiate species. They have developed the most advanced mode of brood-protecting in echinoids (Fig. 3.112). Antrechinus drygalskii (Mortensen, 1905) Antrechinus mortenseni David & Mooi, 1996 Antrechinus nordenskjoldi (Mortensen, 1905) Antrechinus drygalskii (Mortensen, 1905) Mironov (1976) established a subspecies A. d. perfidus from the Kurile Islands in the North Pacific. The holotype Urechinus drygalskii Mortensen, 1905 was lost. The geographical range is now from the Antarctic north to the Kurile Island and Alaska. Test: Pointed with a small subanal fasciole. Distribution: North Pacific and Antarctic waters, at depths of 3,423 to 5,740 m (Mooi & David 1996). Antrechinus mortenseni, Mooi & David, 1996 Test: Thin, in outline regularly oval, upper side low convex; maximum length 40 mm. Apical system: With four gonopores in males, and with an invagination in females for brooding the young. Ambulacra: Nonpetaloid with only one pore on each plate. Oral side: Peristome sunken, periproct on the posterior side with subanal fasciole beneath. Spines: Primary spines thin and sparsely distributed, miliary spines club-shaped. Color: Dark brown with whitish primary spines. Distribution: This species is restricted to the Weddell Sea at depths of 200 to 1,450 m. Antrechinus nordenskjoldi (Mortensen, 1905) This species is very similar to A. mortenseni, but differs by its pointed posterior end, which is ringed by the subanal fasciole and by its smaller size. The periproct lies above the fasciole on the upper side of the test. The development

140 

 3.3 Superorder Atelostomata von Zittel, 1879

B

A

C

C

G

F

E

H

D

B

I

J



of the young in the brood pouches of the female is the same. Genus Cystechinus A. Agassiz, 1879 The test is thin-plated and very large, reaching more than 100 mm in length. The periproct is positioned on the oral side, and no fasciole exists. The anterior genital plates, G2 and G3, are fused and perforated by one or two gonopores. Three species are recorded from the tropical west coast of America: Cystechinus giganteus A. Agassiz, 1898. Four gonopores. West coast of America. Cystechinus loveni A. Agassiz, 1898. Three gonopores. West coast of America. Cystechinus wyvillii A. Agassiz, 1879. Four gonopores. West coast of South America, subantarctic and circumantarctic waters. Cystechinus giganteus A. Agassiz, 1898 Test: Large, ovoid, lowly vaulted with rounded margins; length may reach more than 100 mm. Apical system: Four gonopores, the two anterior ones in the fused genital plate 2 and 3. Ambulacra: Apetaloid, with single pores, phyllodes double-pored. Oral side: Peristome subcentral, subcircular, at the end of a slightly sunken area in ambulacra III; periproct inframarginal. Tubercles: Primaries on aboral side distinct and set with rather long, slender, and curved spines; oral spines straight. Occurrence: Only two specimens are known from off Mazatlan, Mexico. Bathymetric range 1,820 m (Fig. 3.113). Cystechinus loveni A. Agassiz, 1898 Test: Large and high, distinctly conical with rounded margins; maximum length more than 100 mm.

3.3.1 Order Holasteroida Durham & Melville, 1957 

 141

Apical system: Three gonopores. Oral side: Peristome anterior and slightly transversely elongated; periproct inframarginal. Distribution: This species is recorded from the Northern Pacific, in the Bering Sea, the Okhotsk Sea, and off the Kurile Islands. Along the American coast, it is found off Southern California, Baja California and off Acapulco, Mexico. Bathymetric range 2,600 to 4,800 m. Cystechinus wyvillii A. Agassiz, 1879 Test: Large and high (ratio of height to length very variable, up to 65%), more or less conical; outline ovoid; length may reach 130 mm. Apical system: Slightly posterior, four gonopores. Oral side: Flat; peristome slightly anterior, transverse oval or circular; inflated area from peristome to periproct; periproct itself large, close to posterior ambitus, longitudinally elongate. Distribution: Off the west coast of South America and south of Juan Fernandez Island, subantarctic and circumantarctic; bathymetric range 2,012 to 3,935 m. Genus Pilematechinus A. Agassiz, 1904 The genus is characterized by a large and very high, conical test, usually exceeding 100 mm in length. The plating is extremely thin. Labrum and sternum are very small. The three species are difficult to distinguish by morphological characteristics, with only minor differences in shapes of some pedicellariae and of the sphaeridia (Fig. 3.114). Three species are included in this genus: Pilematechinus belyaevi Mironov, 1975. South of Cuba. Pilematechinus rathbuni (A. Agassiz, 1898). West coast of South America. Pilematechinus vesica (A. Agassiz, 1879). West coast of South America and Antarctica.

◂ Fig. 3.111: (A–E) Solenocystis imitans. Test length 52.1 mm, Northern Atlantic. (A) Aboral side: a sharp keel runs from the anterior to the posterior slope and the sunken periproct. (B) Oral side: the median keel is densely set with large tubercles. (C) Side view: posteriorly the test extends into a very conspicuous anal rostrum, the end encircled by a subanal fasciole [drawn after photos in Mironov (2008); tuberculation partly omitted]. (D) Plastron plate pattern: the large labrum (1) is separated from the sternum (2) by two pairs of plates (ambulacral plates yellow, interambulacral plates green). (E) Compact apical system: the fused anterior genital plate carries two pores, the oculars OII and OIV are also perforated by gonopores, followed by nonperforated posterior genital plates, which are in contact with OI and OV. The interpretation of Mironov’s “semifused” apical plates is not quite clear, and the anterior ocular plate OIII is only indicated by two small pores. Modified after Mironov (2008). Scale bar is 5 mm; genital plates in red, ocular plates blue. (F–J) Spatagocystis challengeri. (F) Test length 48 mm. Plate pattern of oral side. Modified after David (1984). (G) Test length is unknown. From Agassiz (1904). Oral side. (H–J) Test with spines, from Agassiz (1881). (H) Aboral side. (I) Side view. (J) Oral side. In younger specimens, the anal heel is distinctly more pronounced.

142 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

B

D

C ocellar plate O III G2 G3

G3

O II G1 OI

O IV G4

O IV G4 OV

G2 O II G1 OI

OV

E

F

G

Fig. 3.112: (A–E) Antrechinus mortenseni. (A) Diagrammatic longitudinal section showing fertilization, growth, and “birth” of juveniles: first, the sperm of the male fertilizes the egg (1.8 mm long), which has left the gonads to be deposited in one brood pouch. Then, the egg grows to a short-spined juvenile (3 mm) in the expanding brood pouch. In the third stage, the original juvenile has reached the long-spined stage (4.5 mm) as new eggs are fertilized in the other pouches. Finally, the apical plates hinge downwards to enlarge the aperture when the juvenile leaves the pouch. It is “born” and the young sea urchin starts his independent life, whereas the other eggs continue to develop. Modified after Mooi & David (1993). (B, C) Apical system of female and male. In the female (B), the terminal plates (TI–TV) are enlarged. The genital plates (G1–G4) are much reduced because there are no gonopores. The gonoducts coming from the gonads lead directly into the internal brood pouches. In the center of the plate system, the apical aperture opens, wherein the juveniles emerge after having finished their development. In the male (C), the apical system is elongated as usual in holasteroids. The ambulacral plates carry only a single pore. Modified after Mooi & David (1996). (D, E) A. mortenseni. (D) Aboral side of female; test length 22 mm, Weddell Sea, Antarctica. ZMH. The apical system is modified into four internal brood pouches. (E) Aboral side of male: test length 33 mm; Weddell Sea, Antarctica. ZMH. At the four gonopores, the papillae protrude. (F, G) Antrechinus nordenskjoldi. Length of test 16 mm; Weddell Sea, Antarctica. ZMH. (F) Aboral side: the apical aperture of the female is covered by densely set, club-shaped miliary spines. The posterior rostrum is ringed by the subanal fasciole. (G) Oral side: the peristome is sunken. The posterior tuft of spines supports the clearing of feces.

3.3.1 Order Holasteroida Durham & Melville, 1957 



A

B

C

D

B

 143

E

F

G

H

J I

K

N L

M

Fig. 3.113: (A,B) Apical system of Cystechinus: (A) Cystechinus wyvilli. (B) Cystechinus loveni. The anterior genital plates G2/G3 are fused and perforated by one or two pores (the genital plates are shaded in a red color, the oculars in blue, the interambulacral plates are gray). Scale bar is 5 mm. Modified after Mooi (1996). (C–E) Oral side of Cystechinus: (C) Cystechinus giganteus. (D) Cystechinus loveni. (E) Cystechinus wyvillii. Plate (1) is the labrum, plate (2) the sternum. Scale bar is 30 mm. Modified after Mooi (1996). (F,G) Cystechinus giganteus. Test length 88 mm, from Agassiz (1904). (F) Aboral side. (G) Side view. (H–K) Cystechinus loveni. Test length 88 mm, South of Acapulco, Mexico. From Agassiz (1904). (H) Aboral side. (I) Oral side. (J) Side view. (K) Posterior side. (L–N) Cystechinus wyvilli. From Agassiz (1881). (L) Aboral side. (M) Oral side. (N) Side view: the lines show the range of variation in the height.

144 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

C

F

B

E

D

G

I

H

Fig. 3.114: (A, B) Pilematechinus: (A) Oral side of Pilematechinus rathbuni. (B) Oral side of Pilematechinus vesica. The tiny confluent labral and sternal plates are marked in green, ambulacra in yellow. Scale bars are 20 mm. Modified after Mooi (1996). (C–E) P. rathbuni. Test length 92 mm, off Columbia. From Agassiz (1904). (C) Oral side: there is a distinctly depressed peristomial area. (D) Posterior side: the periproct is large, positioned above the ambitus. (E) Side view. (F) Apical system. All four genital plates are perforated by a gonopore. The genital plates are shaded in red, the ocular plates in blue, interambulacral plates in gray. Scale bar is 3 mm. Modified after Mooi (1996). (G–I) P. vesica. From Agassiz (1888). (G) Aboral side. (H) Oral side. (I) Side view. The plates are extremely thin, folding in air by its own weight.



Pilematechinus belyaevi Mironov, 1975 Known only from the description of the types. Occurrence: In the Cayman Trench, south of Cuba; depth range 5,800 to 6,740 m. Pilematechinus rathbuni (A. Agassiz, 1898) Test: Highly conical with rather sharp edges, in outline regularly oval; plates very thin; length more than 100 mm. Apical system: Subcentral, with four gonopores; madreporic pores spread over adjacent plates. Ambulacra: Apetaloid, single-pored except adjacent to the peristome. Oral side: Area surrounding the peristome distinctly sunken; labral and sternal plates very small. Periproct: Large, longitudinally oval, well above the ambitus. Tubercles: Primaries densely covering the oral side, aborally more scarcely distributed. Primary spines: Rather short and slender. Color: Brilliant dark violet to light claret. Occurrence: Reported from the Northern Pacific and from the Gulf of Panama to Ecuador; bathymetric range: 2,060 to 3,335 m. Pilematechinus vesica (A. Agassiz, 1879) Test: Extremely thin, almost translucent, tends to collapse in air by its own weight; in outline oval to ovoid. Apical system: Three gonopores, lacking in G2. Ambulacra: Apetaloid, plates with single pores except the one adjacent to the peristome. Oral side: Flat, with a circular, sunken peristomial area, labrum and sternum strongly reduced. Periproct: Just supramarginal. Tubercles: Primaries scattered over the surface, associated spines curved. Color: Dark brown. Distribution: This species is reported from off the west coast of South America and southward to Antarctica; 3,060 to 4,030 m. Genus Urechinus A. Agassiz, 1879 The genus is characterized by three gonopores on the aboral side, the pore is absent from G2, and on the oral side, the labral and sternal plates are not separated. The genus Urechinus consists of three species: Urechinus antipodeanus McKnight 1974. Off South Island of New Zealand. Urechinus naresianus A. Agassiz, 1879. Atlantic, subantarctic waters and North Pacific. Urechinus reticulatus H.L. Clark 1913. Off Baja California, Mexico. Urechinus antipodeanus is very closely related to U. naresianus, differentiated only by the number of teeth

3.3.2 Order Spatangoida L. Agassiz, 1840 

 145

on the valves of the globiferous pedicellariae. Mooi & David (1996) doubt the validity of the species because the number of teeth varies even in individuals. Occurrence: Off South Island of New Zealand; 1,500 m (Fig. 3.115). Urechinus naresianus A. Agassiz, 1879 Test: Lowly domed or subconical, outline ovoid; length may reach more than 50 mm. Apical system: Three gonopores, in G2 lacking; plates very rarely interrupted. Oral side: Slightly sunken toward peristome, interambulacrum 5 more or less inflated and keeled; slight hood above periproct; subanal fasciole in young specimens conspicuous, much less distinct in adults. Distribution: Mironov (1976) found the specimens from the Atlantic (including the Caribbean) and Antarctic undistinguishable from those in the Northern Pacific. They seem to be a single species; bathymetric range 770 to 4,400 m. Urechinus reticulatus H.L. Clark, 1913 Only the holotype is known from off Baja California, Mexico; depth 1,608 m.

3.3.2 Order Spatangoida L. Agassiz, 1840 The members of this order are characterized by having different forms of fascioles, which evolve in different ontogenetic stages. The marginal (or lateral) fasciole evolves very early in ontogeny, before the periproct shifts out of the apical system toward the posterior (see also Schizasterids). Therefore, it is always positioned below or behind the periproct (adoral). The peripetalous fasciole evolves later when the periproct has already shifted backwards (orange bands). It always passes above or in front of the periproct (adapical). As an adaptation to their endofaunal lifestyle, the taxa convergently evolved fascioles, which maintain a water current over the surface of the test. As in the likewise epifaunal Clypeasteroids, the Atelostomata have to keep their test clean and free from uplying particles. Very small spines set with mucus-producing fine hairs are arranged in bands leading in certain patterns over the test. The beating of these cilia creates a current that flushes over the whole surface (Fig. 3.116). Functions of the fascioles The fascioles are dense bands of very small spines that are set with two series of hairs (ciliae) opposite each other. The spinelets end distally in a mucus gland. They are

146 

A

 3.3 Superorder Atelostomata von Zittel, 1879

B

B

C

D

E

G

F

3.3.2 Order Spatangoida L. Agassiz, 1840 



 147

B

A

C apical system (as)

periproct

peripetalous fasciole (pf)

E young

adult

Fig. 3.116: (A–C) Fascioles: (A) The fresh water current maintained by the internal (upper side) and subanal/anal fasciole (posterior side) comes down the funnel running over the surface toward the rear of the animal (Echinocardium). After Nichols (1960). (B) Abatus cavernosus. Peripetalous fasciole, scale 10 mm. (C) Hemiasterids and the palaeostomatids show a fasciole, which lies above the periproct. Modified after Markov & Soloviev (2001).

developed mainly in burrowing sea urchins and serve two important functions: The mucus is spread over the spine canopy of the aboral surface to prevent fine sediment particles from falling onto the epidermis of the animal. The cilia beat in coordination creating a fresh water current running down from the sediment surface (through a funnel or through interstices in the substrate), pumping it over the test to provide the podia in the petals with oxygen and washing the wastewater to the rear of the echinoid into the sediment (see also Echinocardium).

3.3.2.1 Family Hemiasteridae H.L. Clark, 1917

The family is characterized by a conspicuous peripetalous fasciole running around the distal ends of the petals. Ontogenetically, its evolution dates from the time when the periproct had already shifted toward the back.

That means the peripetalous fasciole has evolved twice: as a complete ring around the petals in the Hemiasterids and the Palaeostomatids, and in the Palaeopneustina, as a bow from one side of the marginal fasciole to the other, which sometimes extends to the front of the test above of and separated from the marginal fasciole. The apical system is ethmophract. Numerous fossil genera are included in this family, only two of them survive today. Holanthus A. Agassiz, 1848. Three species. Kupeia McKnight, 1974. Monospecific. Literature: Lovén 1874; Koehler 1914; Grieg 1921; Mortensen 1950c; McKnight 1974. Genus Holanthus Lambert & Thiéry, 1924 The genus evolved in the Upper Cretaceous. In comparison to Hemiaster, Holanthus has broad petals and pinched ambulacral plates when they cross the fasciole.

◂ Fig. 3.115: (A) Urechinus naresianus. (B) Urechinus antipodeanus. The apical system is usually compact. The genital plates are shaded in red, the ocular plates in blue, interambulacral plates in gray. Scale bar is 2 mm. Modified after Mooi (1996). (C) Oral plate pattern of U. naresianus. The labral plate 1 is in contact with the sternal plate 2. Labral plate dark green, sternal lighter green, episternals still lighter green, ambulacra yellow. Scale bar is 20 mm. Modified after Mooi (1996). (D–G) U. naresianus. (D) Test length 33 mm, Davis Strait, Northwest Atlantic. ZMUC. Aboral side. (E) Oral side. (F) Side view. (G) Specimen with spines; test length 35 mm. North Atlantic, MfN Berlin.

148 

 3.3 Superorder Atelostomata von Zittel, 1879

Only three species are recorded today: Holanthus expergitus (Lovén, 1871). North Atlantic to the Caribbean. Holanthus gibbosus (A. Agassiz, 1879). Japan and Malayan Archipelago. Holanthus hickmani (Koehler, 1914). Gulf of Oman, Arabian Sea. Holanthus expergitus (Lovén, 1871) Test: In outline oval, in profile inflated, highest point near the steep posterior side, sloping straightly toward the anterior rounded margin; length up to 37 mm. Apical system: Central, ethmophract with four gonopores. Anterior ambulacrum: Anterior one narrow, adapically sunken, with large pore pairs set with large tube feet developed for funnel-building. Paired ambulacra: Petaloid, distally closed, posterior petals shorter than anterior petals; ambulacral plates strongly compressed beneath the end of the petals, where they are crossed by the peripetalous fasciole. Oral side: Flat to slightly convex, peristome anterior, sunken, kidney-shaped; labral plate bottle-shaped, sternal plates very long. Periproct: High on the truncated posterior side; subanal pore pairs. Peripetalous fasciole: Distinct. Distribution: Northern Atlantic from south of Iceland to the West Indies and the Cape Verdes at depths of 950 to 3,200 m (Fig. 3.117). Holanthus gibbosus (A. Agassiz, 1879) Very much like Holanthus expergitus. H.L. Clark (1917) thought the two species conspecific. Mortensen (1950) also had to admit that he could not distinguish them. He suggested that H. gibbosus should be taken as a geographical variety of H. expergitus. Distribution: The species is recorded from southern Japan and the Malayan Archipelago; bathymetric range approximately 1,500 m. Holanthus hickmani (Koehler, 1914) Main characteristics of the genus: length of holotype 35 mm. Test in outline almost circular, with typically very broad and long paired petals, the pore zones being wide, and the interporiferous zones much narrower. Distribution: This species is only known from the Gulf of Oman, at a depth of 1,260 to 1,520 m. Genus Kupeia McKnight, 1974 The genus is monospecific. Only fragments were found and described.

Kupeia toi McKnight, 1974 Test: Thin-shelled, seems to be subglobular with truncated posterior side. Apical system: Ethmophract, three gonopores associated with genital papillae; 10 madreporic pores. Ambulacra: Anterior one flush with the test, slightly depressing toward the fasciole; double pores very small, close-set, uniserial, associated with specialized penicillate podia; anterior pair short, bending anteriorly, anterior series rudimentary, posterior series larger, the inner pore being round, outer one transversely elongated; posterior petals distinctly longer than anterior ones, almost parallel with transversely oval pores, closed distally; ambulacral pores beyond fasciole single and tiny with small podia. Oral side: Labrum longer than wide, extending to second adjacent ambulacral plate, sternal plates amphisternous; peristome slightly anterior; kidney-shaped. Periproct: On vertically truncated posterior side. Peripetalous fasciole: Narrow, oval and just encircling the distal ends of the petals, no lateral indentations. Tuberculation: Aborally densely scattered in a groundmass of fine granules, no enlarged primaries. Color: Brownish with purple fasciole. Occurrence: West coast of the North Island of New Zealand, off Kaipara Harbour; no depth stated (Fig. 3.118).

3.3.2.2 Family Palaeostomidae Lovén, 1868 Palaeostoma Lovén in Agassiz, 1872. Indo West Pacific. Monospecific. Sarsiaster Mortensen, 1950. Mid-Atlantic. Monospecific. Genus Palaeostoma Lovén in A. Agassiz, 1872 The genus is monospecific. Palaeostoma mirabilis (Gray, 1851) Test: Small, ovoid in outline, highly vaulted with a slightly truncated posterior end; length up to 18 mm. Apical system: Subcentral, ethmolytic with two gonopores in the posterior genital plates, each surrounded by a rim, anterior gonopores absent, genital plates fused. Anterior ambulacrum: Not petaloid, paired pores one above the other, set with differentiated tube feet for building funnels. Paired ambulacra: Broad, closed distally, scarcely sunken; posterior petals about half the size of anterior ones; pores slit-like. Oral side: Convex, peristome pentagonal, covered by five triangular plates. Periproct: High on the posterior side, large radiating plates; subanal pore pairs. Fasciole: Peripetalous fasciole conspicuous.

3.3.2 Order Spatangoida L. Agassiz, 1840 

B

B

D

 149

E

A C

E

F

G

I

H

Fig. 3.117: (A–E) Holanthus expergitus. Young specimen of 14 mm length, off Portugal. After Lovén (1874). (A) Oral side. (B) Aboral side. (C) Side view. (D) Posterior side. (E) The apical system is ethmophract: the posterior genital plates and the ocellar plates join each other. After Mortensen (1950). (F–I) Holanthus gibbosus. Test length 30 mm; west central Pacific. After A. Agassiz (1881). (F) Aboral side. (G) Oral side. (H) Posterior side. (I) Profile.

Tuberculation: Aborally small, spines fine and slender. Distribution: This species is widely spread in the IndoWest Pacific from the Red Sea to the Kei Islands, Indonesia, and northward to the Riu-Kiu Islands, Japan. Bathymetric range 20 to 110 m. Genus Sarsiaster Mortensen, 1950 The genus is monospecific. Sarsiaster griegii Mortensen, 1950 Test: Thin and fragile; almost as long as broad with very shallow anterior notch; aboral side arched; maximum length reaches 53 mm.

Apical system: Central, ethmolytic with two gonopores. Anterior ambulacrum: Depressed adapically; pore pairs small, suckered tube feet having a rosette of short laminae. Paired ambulacra: Petals narrow, sunken, distally closed; anterior pair double the length of posterior pair. Oral side: Slightly convex, peristome D-shaped, labrum with prominent lip, extending to third adjacent ambulacral plates; sternal plates symmetrical. Periproct: Subcircular, on posterior, truncated side. Peripetalous fasciole: Well-developed, slightly indented laterally. Tuberculation: Dense and uniform. Occurrence: Mid-Atlantic; 3,120 m (Fig. 3.119).

150 

A

 3.3 Superorder Atelostomata von Zittel, 1879

B

C

E

D

F

E

F

Fig. 3.118: (A, B) Holanthus hickmani. Test length 35 mm, Gulf of Oman; from Koehler (1912). (A) Aboral side: the petals are extremely broad and long. (B) Profile. (C) Kupeia toi. Aboral side, oral side is unknown. Off the North Island, scale bar is 5 mm. Sketch after McKnight (1974). (D–F) Palaeostoma mirabilis. Test with spines, length 8 mm, Java, Indonesia. ZMUC. (D) Aboral side. (E) Side view. (G) Aboral side with spines, test length 9 mm.

Suborder Brissidina Stockley et al., 2005 There is no explicit description of this clade. It was established as Superfamily Brissidea by Stockley, but following Kroh & Smith (2010), it was classified as a suborder that included the Superfamily Spatangidea. This suborder contains eight families: Brissidae Gray, 1855 Palaeotropidae Lambert, 1896 The following six families are compiled in the superfamily Spatangoidea Kroh & Smith, 2010:

Eupatagidae Lambert, 1905 Eurypatagidae Kroh, 2007 Loveniidae Lambert, 1905 Maretiidae Lambert, 1905 Spatangidae Gray, 1825 Macropneustidae Lambert, 1895 3.3.2.3 Family Brissidae Gray, 1855 The members of this family live buried in the sediment. Except for Brissopsis, none have developed specialized tube feet serving for funnel-building in the anterior

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

D

B

B

 151

C

E

F Fig. 3.119: (A–C) Palaeostoma mirabilis. Young specimen with a length of 8 mm. From Lovén (1883). (A) Aboral side. (B) Oral side. (C) Side view. (D–F) Sarsiaster griegii. Test length 53 mm, Mid-Atlantic. After Grieg (1921). (D) Aboral side. (E) Oral side. The apical system is ethmolytic with two gonopores. Scale bar is 2 mm. After Mortensen (1950).

E

ambulacrum. Consequently, they have to stay near the surface with their apical spines reaching the surface, or they burrow in coarse substrate where they can get sufficient oxygen for respiration. The subanal fasciole below the periproct encloses pores associated with specialized tube feet with finger-like digits, which are used for building sanitary funnels. The bilobed fasciole surrounds two tufts of specialized spines, indicated by enlarged tubercles. They support the construction of two parallel funnels. The shield-shaped fasciole bounds a single tuft of spines, and a single posterior tube is built. The family is characterized as follows: Apical system: Ethmolytic. Paired petals: Sunken, narrow and with parallel sides. Anterior ambulacrum: Simple podia, not differentiated for funnel-building. Labrum (labral plate): Generally very short and wide. Fascioles: Peripetalous and subanal ones are present; in some species, an anal fasciole occurs additionally.

A

anterior

anterior ambulacrum

B

anterior

phyllodes peristome

anterior petals

labrum

apical system

sternal plates

peripetalous fasciole

episternal plates

posterior petals posterior

The composition of the enlarged sternal and episternal plates is called the plastron, the different patterns are important for specific determination. The plastron is covered to various degrees by tubercles set with spines specialized for locomotion in the sediment. They are often spatulate. The adjacent ambulacra AI and AV are more or less naked (Fig. 3.120). The Brissidae were a rather widely defined taxon. Mortensen (1950) already felt that some genera might not belong into this family. Smith (2005) restricted the family to 13 genera and shifted some other genera into the family Maretiidae. Anabrissus Mortensen, 1950. Brazil; monospecific. Anametalia Mortensen, 1950. Indo-Pacific; three species. Brissalius Coppard, 2008. Philippines; monospecific. Brissopsis L. Agassiz in L. Agassiz & Desor, 1947. Worldwide; 15 species. Brissus Gray, 1825. Indo-Pacific, Atlantic, and Mediterranean; six extant species.

posterior

subanal fasciole

Fig. 3.120: Characterization of a brissid spatangoid (Metalia spatagus). (A) The anterior ambulacrum is nonpetaloid. The pore pairs, in a uniserial or biserial row, are associated with specialized podia. They serve for food gathering, funnel building, or as sensory podia. The petals are sunken. The peripetalous fasciole surrounds the distal ends of the petals. (B) Oral side. The plastron is built by the paired sternal plates and, in some groups only, the paired episternal plates. The subanal fasciole lies at the posterior end below the periproct, being shield-shaped. In some genera, it may be bilobed.

152 

 3.3 Superorder Atelostomata von Zittel, 1879

Tab. 3.4: A comparison of the common Brissidae and their features. Brissus

Meoma

Metalia

No anterior notch, anterior side convex

Very faint to distinct anterior notch

Anterior notch

Bilobed subanal fasciole complete

Bilobed subanal fasciole, upper portion lacking, but complete in M. frangibilis and M. cadenati

Shield-shaped subanal fasciole with anal branches

Cionobrissus A. Agassiz, 1879. Indo-Pacific; monospecific. Idiobryssus H.L. Clark, 1939. East Pacific; only juveniles of a single species are known. Meoma, Gray, 1851. Caribbean and West Africa, Pacific Central America; four species. Metalia Gray, 1855. Nine species distributed in the IndoPacific. Neopneustes Duncan, 1889. Caribbean; one species, poorly known. Plagiobrissus Pomel, 1883. Tropical Atlantic, Mediterranean, and Pacific Central America. Rhynobrissus A. Agassiz, 1872. East Coast of USA and IndoPacific; four species. Taimanawa Henderson & Fell, 1969. Japan and Indonesia; two species. The most common genera, Brissus, Meoma, and Metalia look rather similar at first sight. They differ in the following features (Tab. 3.4): Literature: Mortensen 1907; Chesher 1968, 1969, 1970; A.M. Clark & Rowe 1971; Baker & Rowe 1990; Rowe & Gates 1995; Hendler et al. 1995; Miskelly 2002; Lessios 2005; Stockley et al. 2005; van Noordenburg 2008; Kroh & Smith 2010. Genus Anabrissus Mortensen, 1950 The genus is monospecific. Anabrissus damesi (A. Agassiz, 1881) Test: Oval, without any frontal depression; lowly vaulted, posteriorly highest, posterior side truncated. Apical system: With three gonopores, pore lacking in genital plate 3. Anterior ambulacrum: Not petaloid, flush, pore pairs rudimentary. Paired ambulacra: Paired petals short expanding halfway to ambitus, narrow and hardly sunken, interporiferous zone very narrow; anterior pair at an angle of almost 180°. Oral side: Flat; peristome anterior, broad; labrum short and wide; peripetalous fasciole rudimentary, only sporadically visible; subanal fasciole well-developed, bilobed. Periproct: On the truncated posterior side. Occurrence: Only two specimens were found off the coast of Brazil at a depth of approximately 640 m (Fig. 3.121).

Genus Anametalia Mortensen, 1950 The genus is characterized by a more or less conspicuous posterior prominence, the “anal snout”. Three species are recorded: Anametalia grandis Mortensen, 1950. South China Sea. Anametalia regularis (H.L. Clark, 1925). Indonesia. Anametalia sternaloides (Bolau, 1874). South China Sea to Australia. Anametalia grandis Mortensen, 1950 Test: Large, box-shaped with a nearly flat upper side, anterior side vertical, posterior side also vertical with a small, inconspicuous anal snout at the lower margin; outline ovoid with a very shallow anterior furrow; test narrowing toward the posterior end; recorded maximum length 118 mm. Apical system: Anterior, ethmolytic, four gonopores. Anterior ambulacrum: Adapically with close-set, simple pore pairs; lower down they are more distantly set in a shallow, broad depression. Paired ambulacra: Deeply sunken and of about the same size; the anterior pair at an angle of almost 180°, the posterior pair distally diverging. Oral side: Convex, anteriorly slightly sunken toward the peristome; labrum distinct forming a strong lip; peripetalous fasciole bending very slightly inward behind the anterior petals. Subanal fasciole: Present, probably also an anal branch. Periproct: Very high on the posterior side. Tuberculation: A few larger tubercles border the adapical part of the anterior ambulacrum; otherwise, the tubercles are dense and uniform. Occurrence: The sole recorded specimen was found off Nha Trang, Vietnam; at a depth of approximately 50 m. Anametalia regularis (H. L. Clark, 1925) Test: Moderately high, upper side flat and slightly sloping toward the anterior, frontal margin rounded, posterior side steeply sloping toward the anal snout; outline ovoid and with very shallow anterior sulcus; length more than 50 mm. Apical system: Anterior, ethmolytic, four gonopores. Ambulacra: Anterior petal in a shallow, broad depression; paired petals straight, only slightly sunken.

3.3.2 Order Spatangoida L. Agassiz, 1840 



C

A

 153

E

D

B

F

G

H

Fig. 3.121: (A–E) Anabrissus damesi. Test length approximately 20 mm; off Brazil. Rough sketches after the photos in Mortensen (1951). (A) Oral side. (B) Side view. (C) Aboral side: the rather coarse tuberculation is regularly scattered over the aboral surface. (D) The labrum is short and broad. (E) The apical system is ethmolytic. It is highly characteristic that the anterior gonopore perforates genital plate 2, not genital plate 3. Drawings by Ailsa Clark based on the holotype in London, length 22 mm. After Mortensen (1951). (F–H) Anametalia grandis. Test length 118 mm, width 100 mm, height 70 mm; Indochina. ZMUC Holotype ECH 194. (F) Aboral side: the paired petals are deeply indented. (G) Oral side. (H) Side view.

Oral side: Slightly convex, peristome sunken; periproct high on the posterior side; peripetalous fasciole distinct, subanal fasciole encircling the anal snout. Tuberculation: Some larger tubercles border the anterior ambulacrum, a few are scattered along the paired petals; otherwise, dense and uniform.

Distribution: The species is reported from a few specimens taken in the Java and Arafura seas, Indonesia, at a depth of 50 m (Fig. 3.122). Remarks: The three species of Anametalia differ in the following features: A. grandis has a vertical anterior side, a small anal snout, and deeply sunken paired petals. In

154 

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 3.3 Superorder Atelostomata von Zittel, 1879

B

C

D

E

Fig. 3.122: (A, B) Anametalia regularis. Test length 53 mm, Java Sea, Indonesia. ZMUC. (A) Aboral side. (B) Side view: the posterior side is obliquely sloping, the anterior side is rounded. (C–E) Anametalia sternaloides. Test length 49 mm; Thailand, ZMH. (C) Aboral side: the peripetalous fasciole passes around the petals shallowly indented in the interambulacra. (D) Oral side: the plastron is relatively narrow. (E) Side view: at the posterior, a prominent snout projects and is surrounded by the shield-shaped subanal fasciole.



A. regularis, the anterior side is rounded, the paired petals are less depressed, and the anal snout is not conspicuous. A. sternaloides shows a distinctly projecting anal snout and the test is anteriorly much lower than posteriorly. Anametalia sternaloides (Bolau, 1874) Test: Oval with shallow anterior notch; subanal region projecting like a snout; large forms. Apical system: Anterior, four gonopores. Anterior ambulacrum: Narrow and slightly depressed, groove continuing to peristome. Petals: Sunken, posterior pair longer than anterior pair. Oral side: Small peristome, partially covered by the labrum; plastron rather small, adjacent ambulacra broad; peripetalous fasciole scarcely indenting in interambulacra; subanal fasciole shield-shaped surrounding the snout; enclosing two or three tube feet. Periproct: On the obliquely truncated posterior side above the subanal snout. Tuberculation: Short rows of primary tubercles set with larger spines within peripetalous fasciole; coarse spines bounding anterior groove. Color: Young specimens are whitish, adults brownish. Distribution: The species is found from Hong Kong to northern Australia. Its bathymetrical range is 20 to 50 m. Genus Brissalius Coppard, 2008 At first sight, Brissalius is similar to Brissopsis in having naked depressions in its lateral ambulacra and confluent paired petals, but it differs in having a shield-shaped subanal fasciole. This fasciole and the tube feet on each side are associated with a single tuft of spines maintaining the funnel for wastewater. Brissopsis, in contrast, has a bilobed subanal fasciole with two tufts of spines (Fig. 3.123). Brissalius vannoordenburgi Coppard, 2008 Test: Thin and fragile, shallow anterior notch, slight naked depressions along the lateral ambulacral midlines; aboral surface on the whole depressed; sloping toward the anterior to the rounded margin, posterior side obliquely truncated; length up to 38 mm. Apical system: Central, sunken, ethmolytic with four gonopores. Anterior ambulacrum: Narrow and sunken, with funnelbuilding podia. Paired ambulacra: Anterior and posterior pairs conspicuously depressed, short and of equal length, parallelsided and inner pore zones adapically confluent, distally flexed outward with only the most distal pore pairs of the inner series developed. Oral side: Slightly convex; keel from the sternum toward the posterior, continuing into the shield-shaped subanal

3.3.2 Order Spatangoida L. Agassiz, 1840 

 155

fasciole; peristome anterior encircled by penicillate podia, labral plate protruding; peripetalous fasciole parallel to the petals; subanal fasciole shield-shaped, four pore pairs per column, indistinct anal branches. Periproct: On the posterior truncated end, partly visible from above. Tuberculation: On the aboral side, dense and uniform, naked lines along the plate sutures; tubercles on the oral side larger and less densely distributed; broad naked ambulacral areas. Spines: Short and thin, curved and distally widened, on the sternal plate spoon-shaped. Color: Test and spines are cream-white, sometimes orange-brown. Distribution: The species is found off Siquijor, Philippines, at a depth of 200 m. Remarks: With the confluent petals Brissalius looks like a typical Brissopsis, but the shield-shaped subanal fasciole is a characteristic that it shares with Metalia. Genus Brissopsis L. Agassiz in Agassiz & Desor, 1847 This genus has developed large, penicillate tube feet in the anterior ambulacrum specialized for funnel-building, in contrast to Brissus, which has only simple pores and podia. Test: Of medium size, thin, more or less elongated, shallow to moderately indented anterior notch; often faint depressions and/or naked bands along the midlines of the columns, especially in the lateral ambulacra. Apical system: Subcentral, ethmolytic, the madreporite extends far beyond the posterior ocular plates; two to four gonopores. All ambulacra depressed on the aboral side, on the oral side broad and naked. Anterior ambulacrum: With differentiated pore pairs associated with funnel-building tube feet. Petals: Posterior paired petals more or less confluent adapically with inner rows of pore pairs reduced or absent. Fascioles: Peripetalous and bilobed subanal fasciole present; three to four small pore pairs to each side within subanal fasciole; anal fasciole present or not. Periproct: Drop-like and high on the posterior side. Tuberculation: Uniform without distinctly larger primary spines. Remarks: The variation in the species is high, often making it difficult to distinguish. Another problem is that morphological features may change considerably with age. Biology: Chesher (1968) has recorded the ecology of Brissopsis alta: the animals live burrowed in fine, sandy, and muddy bottoms. By using the penicillate tube feet in the upper anterior ambulacrum, the echinoid maintains a respiratory funnel to the surface. The cilia, very fine spinelets in the peripetalous fasciole, create a fresh water current, which flows over the podia and the test surface. The cilia

156 

 3.3 Superorder Atelostomata von Zittel, 1879

B

A

C

E

D

F

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

 157

B

Fig. 3.124: Characteristics of the genus Brissopsis.

in the bilobed subanal fasciole sweep the wastewater into two sanitary funnels behind the animal (Fig. 3.124). Brissopsis differs from Moira and Echinocardium in that the funnel to the surface obviously serves only for pumping fresh water for respiration, not for gathering food (Fig. 3.125). The genus includes 17 species, most of them reported from deeper water: Brissopsis alta Mortensen, 1907. Caribbean, Gulf of Mexico. Brissopsis atlantica Mortensen, 1907. From Florida to Colombia. Brissopsis bengalensis Koehler, 1914. Indian Ocean to Japan. Brissopsis columbaris A. Agassiz, 1898. California to Ecuador. Brissopsis elongata Mortensen, 1907. West Indies, Gulf of Guinea. Brissopsis evanescens Mortensen, 1950. Off southwest Africa. Brissopsis jarlii Mortensen, 1951. Gulf of Guinea. West Africa. Brissopsis luzonica (Gray, 1851). Indo-West Pacific. Brissopsis lyrifera (Forbes, 1841). East Atlantic including the Mediterranean. Brissopsis lyrifera capensis (Mortensen, 1907). South Africa from Agulhas Bank to Gulf of Guinea. Brissopsis mediterranea Mortensen, 1913. Mediterranean to Gulf of Guinea. Brissopsis micropetala Mortensen, 1948. Philippines. Brissopsis obliqua Mortensen, 1948. Philippines. Brissopsis oldhami Alcock, 1893. Bay of Bengal, Malayan region to New Zealand. Brissopsis pacifica A. Agassiz, 1898. California to the Galapagos Islands. Brissopsis parallela Koehler, 1914. Bay of Bengal. Brissopsis similis Mortensen, 1948. Philippines.

Brissopsis zealandiae Mortensen, 1921. New Zealand. Most probably synonymous with B. oldhami. Brissopsis alta Mortensen, 1907 Main characteristics of the genus: Test: Broad and solid; subglobular, length may reach 65 mm; posterior side vertically truncated. Paired petals: Short, almost straight and divergent. Fascioles: Peripetalous fasciole with sharp indentation, distant from petals; subanal fasciole with two to three pore pairs, anal branches absent. Distribution: Endemic to the south of USA, from the Atlantic coast off Florida into the Gulf of Mexico to the mouth of the Mississippi; bathymetric range 90 to 310 m. Brissopsis atlantica Mortensen, 1907 Main characteristics of the genus: Test: Elongated, low, highest point posterior, length may reach more than 70 mm; posterior side more or less obliquely sloping. Paired petals: Short and flexed toward the sides, posterior pair confluent proximally. Periproct: Visible from above. Fascioles: Peripetalous fasciole shallowly indented laterally; subanal fasciole with five entering ambulacral plates and four pore pairs on each side; distinct anal branches. Distribution: Known from Florida and the Bahamas through the Gulf of Mexico to Colombia, at depths of 100 to 3,200 m. Remarks: Mortensen (1951) could not decide if B. atlantica and B. mediterranea are separate species or only subspecies. Chesher (1968) clarified the differences: B. atlantica has developed four ambulacral plates entering the subanal fasciole with three tube feet on each side. This species is

◂ Fig. 3.123: Brissalius vannoordenburgi. (A–D) Specimen with test length 29 mm, Palawan, Philippines. (A) Aboral side: the paired petals are short and confluent, diverging only in the most distal part. (B) Oral side. (C) Oblique view from the posterior: the subanal fasciole is shield-shaped. (D) Side view. (E, F) Specimen with spines, length 28 mm; Palawan, Philippines. (E) Aboral side. (F) Oral side: the distal ends of the plastronal spines are spoon-shaped.

158 

 3.3 Superorder Atelostomata von Zittel, 1879

B

A

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G



distributed in the Caribbean from Florida through the Gulf of Mexico to Colombia and the Bahamas. B. mediterranea developed five ambulacral plates entering the subanal fasciole with four tube feet on each side, and is distributed in the Mediterranean, West Africa to the Gulf of Guinea, over the Atlantic in parts of the Bahamas and the East Coast of the USA. Brissopsis bengalensis Koehler, 1914 Main characteristics of the genus: Test: Oval, relatively low, maximum length 85 mm. Ambulacra: Anterior paired petals diverging distally, posterior pair slightly sinuous. Fascioles: Peripetalous fasciole with a conspicuous indentation laterally; subanal fasciole present. Distribution: Known from Ceylon to the Philippines and Japan (Sagami Bay), at depths from 130 to 2,980 m. Brissopsis columbaris A. Agassiz, 1898 Main characteristics of the genus: Test: Of moderate size, posterior side sloping obliquely, periproct visible from above; length may reach approximately 50 mm. Petals: Broad, anterior pair divergent, anterior column bowed; posterior pair less diverging. Fascioles: Lateral indentations in peripetalous fasciole very slight; subanal fasciole broad, anal branches absent. Distribution: Baja California, Mexico, to Ecuador, at depths of 730 to 2,280 m (Fig. 3.126). Brissopsis elongata Mortensen, 1907 Main characteristics of the genus: Test: Elongated, rather low, tapering toward the truncated posterior face, maximum length not reaching 100 mm. Ambulacra: Paired petals of about same length, anterior pair widely diverging, posterior pair confluent adapically and flexed outward distally; anterior ambulacra beyond petals tuberculated. Oral side: Convex, posterior face truncated, sloping almost vertical. Fascioles: Peripetalous fasciole with wide, shallow lateral indentation; subanal fasciole with four pore pairs on each side, no anal branches. Distribution: Known in the Caribbean from the Florida Keys to Belize and Venezuela, bathymetric range 13 to 72 m.

3.3.2 Order Spatangoida L. Agassiz, 1840 

 159

Brissopsis evanescens Mortensen, 1950 Test: Compact, upper and underside convex, length may reach 50 mm. Paired petals: Very large and broad; pore pairs hardly reduced adapically. Fascioles: Peripetalous fasciole with shallow lateral indentation, subanal fasciole reduced or even absent in adults. Distribution: This species was found in enormous numbers on the west coast of South Africa off Walfish Bay, Namibia; 135 to 320 m (Mortensen 1950). Brissopsis jarlii Mortensen, 1951 Very much resembling Brissopsis elongata. Test: Broad and rather low, aboral side very slightly arched, oral side convex; length reaching 70 mm. The petaloid area is relatively small; posterior pair of petals confluent adapically, strongly diverging distally. Fascioles: Similar to B. elongata. Distribution: Endemic to the Gulf of Guinea, West Africa, at a depth of 37 to 59 m. Brissopsis luzonica (Gray, 1851) Test: Thin and fragile, elongated with more or less conspicuous indentations at the margin in the anterolateral ambulacra; anterior notch moderately deep; on the whole rather low, upper side more or less flattened, highest point at the posterior end; maximum length more than 80 mm. Apical system: Subcentral or slightly anterior. Anterior ambulacrum: Sunken and set with differentiated pore pairs. Paired petals: Sunken; anterior pair strongly diverging distally, posterior pair almost parallel, only very slightly flexed outward at the ends; inner rows of pore pairs greatly reduced or absent near the apical system. Oral side: Slightly convex, sternal plates entirely tuberculated, adjacent naked ambulacra broad; peristome kidney-shaped; peripetalous fasciole follows the ends of the petals, conspicuously indenting in the paired interambulacra; subanal fasciole bilobed with three to four pore pairs on each side, anal branches narrow, but distinct. Periproct: High on the steeply rounded posterior end. Tuberculation: Rather uniform, larger on the oral side; no conspicuous primary spines at the upper side. Color: The spines are light brownish, the peripetalous fasciole distinctly darker. The bare test is whitish.

◂ Fig. 3.125: (A–C) Brissopsis alta. Test length 50 mm, Gulf of Mexico. ZMUC ECH 58. (A) Aboral side. (B) Oral side: the naked ambulacral areas are broad. (C) Side view. (D, E) Brissopsis atlantica. Test length 44 mm, Gulf of Mexico, USA. ZMUC ECH 68. (D) Aboral side. (E) Side view. (F, G) Brissopsis: part of subanal fasciole. (F) B. atlantica: Five ambulacral plates enter the subanal fasciole on each side, and four pore pairs with specialized tube feet are developed. (G) Brissopsis mediterranea: four ambulacral plates enter the subanal fasciole on each side, and three pore pairs set with specialized tube feet are developed (marked in red). Scale bar is 6 mm. After Mortensen (1951).

160 

 3.3 Superorder Atelostomata von Zittel, 1879

C

A

B D

E

F

H G

3.3.2 Order Spatangoida L. Agassiz, 1840 



Distribution: This species is widely known in the IndoWest Pacific from the Red Sea to Mozambique, from Japan to northeast Australia, and from Tahiti and Hawaii to the North Island of New Zealand; bathymetrical range 10 to 1,500 m.

A

D

 161

Remarks: This specimen is distinguished from the typical B. luzonica by the anterior position of the apical system and by the consequently shorter proximal part in the anterior pair of, before it spreads outward distally in a much wider angle. It is also unusually elongated (Fig. 3.127).

B

C

E

F

G

Fig. 3.127: (A–D) Brissopsis luzonica. Outline of test, length 37 mm. (B) Young specimen; test length 25 mm; Indian Ocean. Aboral side: the narrow petaloid area of this rather young specimen is still small. (C) Petaloid area: the peripetalous fasciole follows the petals closely. The inner series of pore pairs are reduced adapically. Scale bar is 10 mm; after Mortensen (1951). (D) Profile: length of test 55 mm. (E–G) Brissopsis cf. luzonica. Test length 77 mm, off northeast Taiwan. (E) Aboral side: the apical system is positioned anteriorly, the anterior paired petals broadly flexed outward. (F) Oral side: the naked areas are broad and straight. (G) Side view. ◂ Fig. 3.126: (A, B) Brissopsis bengalensis. Test length 64 mm, Ceylon. (A) Aboral side. (B) Side view. Sketches after photos of Koehler (1914). (C, D) Brissopsis columbaris. Test length 31 mm, Baja California, Mexico. ZMUC cotype ECH 114. (C) Aboral side. (D) Profile: the posterior side slopes obliquely toward the rounded rear. The periproct is supramarginal. (E, F) Brissopsis elongata. Test length 45 mm; Venezuela. ZMUC ECH 399. (E) Upper side: the anterior pair of petals is straight, the posterior pair flexed outward. (F) Side view. The oral side is convex. (G, H) Brissopsis evanescens. Test length 46 mm; off Walfish Bay, Namibia. ZMUC ECH 133. (G) Aboral side. The petals are broad and long. (H) Side view. Aboral and oral sides are convex.

162 

 3.3 Superorder Atelostomata von Zittel, 1879

Brissopsis lyrifera (Forbes, 1841) Main characteristics of the genus: Test: Thin with anterior notch, more or less shallowly indented at the anterolateral ambulacra; in profile the shape may vary: upper side convex or flattened with the highest point posterior; the rear subvertical or obliquely truncated with a short rostrum; length up to 70 mm. Paired petals: Strongly divergent with parallel sides, the posterior pair at a less wide angle; inner series of pore pairs only proximally rudimentary. Fascioles: Peripetalous one strongly indented in the paired interambulacra, subanal fasciole. Distribution: This species is known in the eastern Atlantic from the Lofoten Islands, Norway to Iceland, along the European coasts into the Mediterranean and southward to the Canaries and West Africa. It is found at depths from 5 to 1,400 m (Fig. 3.128). Brissopsis lyrifera capensis (Mortensen, 1907) Characteristics on the whole similar to Brissopsis lyrifera, but with higher test of more than 60% of total length, broad, upper side sloping to the anterior margin, posterior side vertically truncated. Fascioles: Peripetalous fasciole distant from petals; subanal fasciole present, no anal branches. Distribution: This subspecies is known from the Agulhas Bank, South Africa to the Gulf of Guinea, at depths of approximately 275 to 485 m. Brissopsis mediterranea Mortensen, 1907 Test: Posteriorly higher than anteriorly, posterior side more or less vertically truncated; length may reach 66 mm. Paired petals: Anterior pair not very divergent; in younger specimens, posterior pair parallel and confluent in most part of its length, only distally flexed outward, in larger specimens more divergent. Fascioles: Peripetalous one fairly parallel to petals, subanal fasciole with three pore pairs; anal branches well-developed, halfway up to peripetalous fasciole. Distribution: Mediterranean, West African coast to the Gulf of Guinea; East Coast of USA from Florida to Nantucket and parts of the Bahamas; 46 to 3,200 m. Remarks: For the distinction between B. mediterranea and B. atlantica, see Fig. 3.129. Brissopsis micropetala Mortensen, 1948 Test: Broad, anteriorly low, rising posteriorly toward the steeply truncated side; length up to 78 mm. Paired petals: Short, anterior pair diverging, distal part flexed, posterior pair confluent in most of the length, straight, distally very slightly bent outward.

Oral side: Flat; naked ambulacra very broad; peripetalous fasciole rather narrow, slightly indented laterally; subanal fasciole with three podia in each column; anal branches lacking. Distribution: Philippines, at depths of 185 to 1,040 m (Fig. 3.130). Brissopsis obliqua Mortensen, 1948 Test: Very narrow and elongated, greatest width in the middle, tapering gently to both ends; in profile low, posterior side sloping very gradually toward the posterior end, which projects rather conspicuously; largest length recorded 40 mm. Apical system: Madreporite rather short, not extending beyond the posterior ocular plates. Anterior ambulacrum: With very small adapical pores, set with conspicuously long, reddish tube feet. Paired petals: Only slightly sunken, posterior pair with parallel pore zones proximally, but barely confluent, the inner series being a little smaller. Oral side: Flat, naked ambulacral areas broad; peristome rather distant from anterior margin; periproct visible from above, distinctly pointed aborally; peripetalous fasciole with very slight indentation laterally; subanal fasciole almost circular with four tube feet to each side; anal branches present. Spines: Forming a distinct pair of tufts within the anal fasciole, approximately 10 mm long. Color: Test and spines whitish, fascioles may be darker. Occurrence: South of Panay, Philippines; 250 m. Brissopsis oldhami Alcock, 1893 Test: Very fragile; anteriorly lower than posteriorly; oral side convex; maximum length 83 mm. Paired petals: Short and almost straight, anterior pair strongly divergent, posterior pair less so, confluent adapically; ambulacral plates outside the paired petals well tuberculated. Fascioles: Peripetalous fasciole closely following the outline of the petals; subanal fasciole with three podia, no anal branches. Color: Yellowish-pink, fasciole darker, juveniles dull pink, fascioles purple. Distribution: Bay of Bengal, Malayan region, Philippines to New Zealand; 91 to 2,012 m. Remarks: Having at hand diverse ontogenetic stages of this species from the Cook Strait, Fell (1958) and McKnight (1969) stated that Brissopsis zealandiae Mortensen, 1921, which based on relatively young specimens, is synonymous with B. oldhami. Brissopsis pacifica A. Agassiz, 1898 Test: Low, not raised posteriorly, oral side slightly convex, posterior end vertical; length up to 75 mm.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

D

 163

C

B

E

F G

Fig. 3.128: (A–E) Brissopsis lyrifera. Test length 42 mm; Kattegat, Sweden. (A) Aboral side: the petals are straight and divergent. (B) Oral side: the posterior ambulacra are broad and naked, set only with fine miliary spines. (C, D) Specimen with spines. Test length 44 mm; North Sea. (C) Aboral side: the peripetalous fasciole is distinct. (D) Oral side: the spines used for burrowing are longer than those of the upper side. (E) Enlargement of apical field. Test length 54 mm, Senegal, West Africa. Aboral side: near the apical system, the pore pairs in the petals become rudimentary. The madreporite extends greatly beyond the ocular plates. (F, G) Brissopsis lyrifera capensis. Specimen with spines, test length 55 mm, Agulhas Bank, South Africa. ZMUC. (F) Aboral side. (G) Side view.

164 

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 3.3 Superorder Atelostomata von Zittel, 1879

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Fig. 3.129: Brissopsis mediterranea. (A, B) Test length 54 mm, Canaries. ZMUC. (A) Aboral side: the posterior pair of petals is adapically parallel, distally flexed outward. (B) Side view. (C–E) Specimen without spines, test length 90 mm, Bay of Guinea, West Africa. ZMUC. (C) Aboral side: in each petal, one of the columns of pore pairs is rudimentary near the apical system. (F) Oral side: the naked areas in the posterior ambulacra are straight. (E) Side view.

3.3.2 Order Spatangoida L. Agassiz, 1840 



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Fig. 3.130: (A–D) Brissopsis micropetala. Test length 78 mm, Philippines. ZMUC ECH 226. (A) Aboral side: the posterior pair of petals is almost parallel. Specimen with spines, test length 57 mm, Philippines. ZMUC ECH 517. (B) Aboral side: the petaloid area is small. (C) Oral side: the naked areas are very broad. (D) Side view: the posterior side is truncated. (E) Brissopsis obliqua. Aboral side within the peripetalous fasciole. Scale bar is 8 mm, after Mortensen (1951). (F–I) Brissopsis oldhami. (F, I) Test length 69 mm; Philippines. ZMUC. (F) Aboral side. (I) Profile. (G, H) Specimen without spines (as Brissopsis zealandiae Mortensen, 1921, ZMUC ECH 341), test length 28 mm, Bare Island, New Zealand. (G) Aboral side: the posterior pair of petals is less confluent than in the older specimen above. (H) Side view.

Apical system: The number of gonopores may vary from two to four. Paired petals: Very long, anterior pair divergent with rudimentary pore pairs in the anterior columns; posterior pair only adapically confluent, distally widely diverging. Fascioles: Peripetalous fasciole broad, shallowly indented laterally; subanal fasciole more than twice as broad as long; anal branches absent.

Color: Brown with dark fascioles. Distribution: Reported from off southern California, USA, to Panama and the Galapagos Islands at 36 to 1,430 m (Fig. 3.131). Brissopsis parallela Koehler, 1914 Main characteristics of the genus: Test: More or less broad and low; posterior side sloping to a rounded margin; length up to 50 mm.

166 

 3.3 Superorder Atelostomata von Zittel, 1879

B

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I

Fig. 3.131: Brissopsis pacifica. (A–C) Test length 40 mm, La Jolla, CA, USA. ZMUC. (A) Aboral side. The specimen has only three welldeveloped gonopores, the fourth one is much reduced. (B) Oral side. (C) Side view. (D) Test length 34 mm; La Jolla, CA, USA. Aboral side: there is much variation in the shape. This specimen has a more anteriorly positioned apical system and more laterally spread anterior petals. (E) Specimen without spines, test length 40 mm; La Jolla, CA, USA (aboral side).



Paired petals: Anterior pair slightly diverging, distally bending abruptly toward the sides; posterior pair parallel almost in its whole length; inner pore series reduced. Peripetalous fasciole: Shallowly indented laterally; subanal fasciole present. Occurrence: Known from the Bay of Bengal off the Andaman Islands; 340 to 1,290 m (Fig. 3.132). Brissopsis similis Mortensen, 1948 Test: Inflated with rounded margin, posteriorly much higher than anteriorly; in outline narrow and elongated, lateral naked depressions in ambulacra very conspicuous; anterior notch shallow, posterior end sloping obliquely; length up to 66 mm. Apical system: Subcentral to slightly posterior. Anterior ambulacrum: Pores large, associated with large and dark podia. Paired petals: Short, anterior pair flexed outward only distally, reduced pore pairs adapically, posterior pair parallel and confluent in most of their length, flexed outward only distally. Oral side: Convex; periproct visible from above; naked ambulacral areas broad. Fascioles: Peripetalous fasciole parallel to outline of petals, subanal fasciole with four pore pairs (one may be rudimentary), anal branches distinct. Distribution: Philippines; 500 to 1,420 m. Remarks: This species is very similar to B. luzonica. It differs by its much higher posterior part. Furthermore, the anterior ambulacrum outside the petals is totally naked, in B. luzonica, each plate carries a tubercle. Brissopsis zealandiae Mortensen, 1921 Most probably synonymous with Brissopsis oldhami Alcock, 1893 (see under that species). Genus Brissus Gray, 1825 The genus is characterized by its well-developed, deeply sunken paired petals. Test: Large, regularly oval, no anterior notch; upper side vaulted, posterior end more or less truncated. Apical system: Anterior, ethmolytic, four gonopores. Anterior ambulacrum: Flush, small pore pairs with simple tube feet. Paired petals: Sunken, narrow and parallel sided. Oral side: With broad plastron, side bowed; labral plate short and wide, peristome kidney-shaped. Periproct: On the posterior side. Peripetalous fasciole: Well-developed, indented in all five interambulacra.

3.3.2 Order Spatangoida L. Agassiz, 1840 

 167

Subanal fasciole: Bilobed, usually enclosing four specialized podia on each side. Tuberculation: Anteriorly larger than posteriorly. The genus Brissus includes four recent species according to Smith (2005): They strongly resemble each other. They can be distinguished mostly by the shape of the posterior side and by the pedicellariae, although the variability is considerable. Larger tests are relatively broader and the aboral posterior interambulacrum is more keeled. Biology: The species prefers coarse sediments, where they live buried down to a depth of approximately 20 cm, and usually in large numbers. The species are ecologically important because they “clean” the sea floor by swallowing large quantities of substrate, ingesting only the organic content and discharging the sand particles. They are often preyed on by snails. Literature: Mortensen 1913; Hendler et al. 1995; Rowe & Gates 1995; Miskelly 2002. Brissus (Allobrissus) agassizii Döderlein, 1885. Japan to East Australia (includes B. gigas Fell). Brissus latecarinatus (Leske, 1778). Indo-Pacific. Brissus obesus Verrill, 1867. Tropical west coast of America from the Gulf of California to Panama. Brissus unicolor (Leske, 1778). East and west Atlantic, Mediterranean. Brissus (Allobrissus) agassizii Döderlein, 1885 General characteristics of the genus. Test: Strong; outline elliptical without anterior notch, the aboral side is scarcely raised and scarcely keeled in the posterior interambulacrum; length may reach more than 150 mm. Paired petals: Anterior pair long, reaching to the sides, conspicuously sinuous; posterior pair even longer, slightly diverging and distally flexed. Posterior side: More or less vertically truncated, periproct slightly depressed. Tubercles: Coarse, set with robust spines around the anterior half of the aboral part of the test. Distribution: This species is widely known from Japan throughout the Malayan Archipelago to southern Australia from the intertidal zone to a depth of 120 m. Remarks: The subgenus Allobrissus differs from Brissus only in the shape of its pedicellariae. Brissus (A.) agassizii and Brissus latecarinatus have widely overlapping distributions, and they are very similar (Fig. 3.133). Brissus latecarinatus (Leske, 1778) Test: Strong, aborally rather high with a distinct keel in the posterior interambulacrum (not yet developed in small specimens); posterior end overhanging the periproct,

168 

A

 3.3 Superorder Atelostomata von Zittel, 1879

C

B

D E

F

Fig. 3.132: (A,B) Brissopsis parallela. Test length 37 mm. (A) Side view: the test is rather low. (B) Petaloid area: the posterior paired petals are parallel, hardly diverging. Sketches after the photos in Koehler (1912). (C–F) Brissopsis similis. Test length 63 mm; Philippines. ZMUC ECH 514. (C) Aboral side: the petals are distally flexed outward. (D) Posterior side with subanal fasciole. (F) Side view. The posterior side is much higher than the anterior side. (E) Oral side: the naked areas are very broad.

3.3.2 Order Spatangoida L. Agassiz, 1840 



B

A

C

 169

D

E

F H G

Fig. 3.133: (A–C) Brissus (A.) agassizii; test length 94 mm; Fukui Prefecture, Japan. (A) Aboral side: the tubercles of the anterior part of the test are distinctly larger than those on the posterior side. (B) Oral side: the periproct is not visible in oral view. (C) Side view. (D) Specimen with spines, test length 89 mm; Fukui Prefecture, Japan. Aboral side: the spines around the anterior part of the test are coarser than those around the posterior. (E–H) Brissus latecarinatus. Test length 116 mm, Ile de la Réunion. (E) Aboral side: the tubercles of the anterior part of the test are only slightly enlarged compared with those of the posterior part. (F) Oral side: the periproct is entirely visible in oral view. (G) Side view. (H) Specimen with spines, test length 49 mm; Dumaguete, Negros, Philippines. Oral side: the spatulate spines of the plastron are directed to the anterior and specialized for digging.

170 

 3.3 Superorder Atelostomata von Zittel, 1879

which is consequently visible in oral view; length up to 140 mm. Paired petals: Anterior ones directed laterally at an angle of more than 180°, not reaching the sides, may be slightly flexed anteriorly; posterior pair slightly divergent and hardly flexed distally. Spines: Short and densely packed, increasing in size around the anterior half of the aboral side. Color: The species is brown in life, adults are darker than smaller specimens. The bare test is grayish white. Distribution: B. latecarinatus is known from the Red Sea and Mauritius through the Malayan Archipelago to Hawaii and the Marquesas Islands and in Australia from the northeast round the south to the Ningaloo Reef on the west coast. It lives from the intertidal to a depth of 45 m. Remarks: This species differs from Brissus (A.) agassizii by the following characteristics: the test is higher, the posterior interambulacrum more keeled and there is a smaller angle between the posterior petals; periproct entirely visible from the oral side; plastron broader, more convex; tubercles smaller and more closely set associated with smaller spines around the anterior part of the aboral side. Brissus obesus Verrill, 1867 Main characteristics of the genus: length may reach more than 100 mm. Aboral side: Vaulted, sloping to a low posterior end. Distribution: This species is recorded in the East Pacific from the Gulf of California to the Gulf of Panama in shallow water (Fig. 3.134). Brissus unicolor (Leske, 1778) Main characteristics of the genus: Test: Solid and relatively low; outline oval without anterior notch, length up to 130 mm; highest point behind the apical system, posterior end rounded. Anterior pair of petals: Diverging at an angle of 180°, distally slightly flexed. Spines: Short and densely packed, more solid and coarser around anterior part of test, as indicated by larger tubercles. Habitat: This species lives under rocks, or buried in sandy sediments. Distribution: Brissus unicolor is known on either side of the Atlantic, from Florida and Bermuda through the Caribbean to Brazil and in the Mediterranean, the Cape Verde Islands and the Azores. It lives from the intertidal zone to a depth of 240 m. Genus Cionobrissus A. Agassiz, 1879 The genus is monospecific.

Cionobrissus revinctus A. Agassiz, 1879 Test: Oval with very conspicuous anterior notch, posterior side pointed by the prominent subanal heel; test vaulted, highest posteriorly; moderately sized. Apical system: Ethmolytic, but the madreporite extends hardly beyond the posterior ocular plates; four gonopores. Anterior ambulacrum: Anterior ambulacrum adapically flush, deeply depressed at the ambitus, continues as a deep furrow toward the peristome. Paired petals: Slightly bowed, interporiferous zone narrow. Oral side: Convex, peristome small and facing forward; labrum elongated, sternal plates long; peripetalous fasciole not bending inward behind the anterior petals; subanal fasciole encircling the posterior heel. Periproct: Round, high on the truncated posterior side above the anal snout. Tuberculation: A few distinctly larger tubercles within the peripetalous fasciole, aborally coarse tubercles along the anterior sulcus, smaller tubercles scarce. Occurrence: This species is only recorded from the Arafura Sea at a depth of 1,460 m. Remarks: Both Cionobrissus and Anametalia have developed similar anal snouts. Cionobrissus differs in having a deep frontal notch continuing toward the peristome. With its elongated labral plate, Cionobrissus stands alone in the family Brissidae (Fig. 3.135). Genus Idiobryssus H.L. Clark, 1939 The genus is monospecific. Idiobryssus coelus H.L. Clark, 1939 Only juveniles of less than 12 mm are known. Test: Oval, no frontal depression; aboral side slightly concave; small subanal heel. Apical system: Anterior, no gonopores developed; plating unknown. Anterior ambulacrum: Narrow and flush; simple pores. Paired petals: Almost flush, narrow and straight, pore zones parallel, interporiferous zone narrow; anterior pair longer than posterior pair. Oral side: Convex; peristome D-shaped, not overarched by labrum; plastron plating unknown, but labral plate appears to be short and wide; sternal plates fully tuberculated; peripetalous fasciole not indented in posterior interambulacra, subanal fasciole present. Periproct: On the posterior aboral side. Occurrence: Galapagos Islands, East Pacific; depth 70 to 120 m. Remarks: The specimens show the typical features of a juvenile: The gonopores are not yet developed, and the periproct is still positioned on the aboral side. Adult specimens are unknown.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

B

 171

C

F

D

E

Fig. 3.134: (A, D) Brissus obesus. Test length 100 mm, Taboguilla, Panama. ZMUC. (A) Aboral side. (D) Side view: the posterior end is low. (B, C, E, F) Brissus unicolor. Test length 89 mm; Crete, Greece. (B) Aboral side: the anterior petals lie at an angle of 180° to each other. The peripetalous fasciole surrounds the distal ends of the petals indenting at each interambulacrum. (C) Oral side: the periproct is not visible from below. (E) Side view. (F) Young specimen, test length 37 mm; Siquijor, Philippines. Small individuals are relatively more elongated than fully grown specimens.

Genus Meoma Gray, 1851 The genus is characterized as follows: Test: More or less solid, ovoid with more or less shallow anterior notch; upper side almost hemispherical, oral side flattened; very large forms. Apical system: Small, four gonopores. Anterior ambulacrum: Narrow, shallow furrow continuing to the peristome; pore pairs small and undifferentiated. Paired petals: Long and narrow, anterior and posterior pair of about equal length, conspicuously sunken. Oral side: Fully tuberculated plastron; peristome kidneyshaped, widely covered by prominent labrum; periproct on short, steeply undercut posterior side. Fascioles: Peripetalous fasciole indented behind the anterior petals, subanal fasciole bilobed, well-developed in juveniles; in adults, the portion next to the peristome may have disappeared (not in Meoma cadenati and Meoma frangibilis). Tuberculation: Within peripetalous fasciole slightly enlarged, set with larger spines.

Color: In life, the spines are light to dark brown; the bare test is white. Remarks: The genus consists of four species or subspecies: two common species live on each side of Central America. They are almost equal in shape having evolved from a single, large population, which divided after the closing of the Panamic seaway during the Pliocene. According to Chesher (1970), the Pacific species M. grandis should be a subspecies of the Caribbean species M. ventricosa. Meoma cadenati Madsen, 1957. West Africa. Meoma frangibilis Chesher, 1970. Caribbean in deeper water. Meoma ventricosa ventricosa Lamarck, 1816. Caribbean from Florida to Venezuela. Meoma ventricosa grandis Gray, 1851. East Pacific from California to the Galapagos Islands. Affinities: Both subspecies of M. ventricosa differ rather little in the distance from the subanal fasciole to the periproct and sometimes in the color: M. v. grandis is dark brown to black with coarse tuberculation, whereas M. v. ventricosa is

172 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

D

C

G

E F

H

I

Fig. 3.135: (A–G) Cionobrissus revinctus. (A, B) Test length 48 mm, Arafura Sea. ZMUC. (A) Aboral side: the peripetalous fasciole is not indented between the lateral petals. (B) Side view: the usually prominent anal snout is broken. (C) Plastron. (E) Apical system. Scale bar is 4 mm, after Mortensen (1950). (E–G) Test length approximately 45 mm, Arafura Sea, Indonesia. After A. Agassiz (1881). (E) Aboral side with spines. (F) Side view of denuded test. (G) Oral side with spines. (H, I) Idiobryssus coelus. Juvenile specimen with a length of 12 mm, Galapagos Islands. (H) Aboral side: the periproct is still positioned on the aboral side. (I) Profile: the upper side is concave, the oral side convex. After H.L. Clark (1939).

mostly more or less dark reddish-brown with finer tubercles. They both have shallow anterior notches, the subanal fasciole is usually incomplete and there are 8 to 10 oral pores in the posterior phyllodes. In contrast M. cadenati and M. frangibilis have distinctly deeper frontal notches, the subanal fasciole is complete, and the posterior phyllodes are set with five to six pores. Meoma cadenati Madsen, 1957 Test: Robust, aboral side moderately inflated, sloping steeply toward the anterior margin, less steeply toward the undercut posterior side; deep frontal notch, length almost 140 mm. Apical system: Distinctly less than one-third of test length away from the anterior.

Ambulacra: Anterior and posterior paired petals of about the same size, anterior pair slightly bowed. Oral side: Flat, peristome at the end of a shallow depression, narrow compared with M. ventricosa; broadly oval plastron; peripetalous fasciole present, bilobed subanal fasciole complete. Distribution: The species is only recorded from West Africa in deeper water (Fig. 3.136). Meoma frangibilis Chesher, 1970 This species is similar to M. cadenati in most characteristics. It differs from that species in the thin, very fragile test and in the shorter anterior petals; length up to 140 mm (Lessios 2005).

3.3.2 Order Spatangoida L. Agassiz, 1840 



B

A

D

B

 173

C

E F

G

Fig. 3.136: (A–C) Meoma cadenati. Test length of specimen 132 mm, Sierra Leone. Sketches after photos in Chesher (1970). (A) Aboral side. (B) Oral side. (C) Side view. (D–F) Meoma frangibilis. Length of specimen 130 mm, Bay of Panama. Sketches after Chesher (1970). (D) Aboral side: the anterior pair of petals is distinctly shorter than the posterior pair. (E) Oral side: the bilobed subanal fasciole is complete. (F) Side view. (G, H) Schematic cross-section of Meoma ventricosa in its burrow. (G) Small specimens live entirely buried in coarse sediment with scattered blocks of coral. They still have a complete subanal fasciole and create two short sanitary funnels. (H) Larger specimens burrow in fine sand only as deep as the apical spines, which still reach the surface. In coarse substrates, they may dig deeper; in silt or mud, the sea urchins are only half buried. The arrows indicate the direction of locomotion. Behind the animals, the sediment is disturbed. Modified after Chesher (1969).

174 

 3.3 Superorder Atelostomata von Zittel, 1879

Occurrence: Bay of Panama, Pacific; at a depth of 55 to 98 m. Meoma ventricosa grandis Gray, 1851 Main characteristics of the genus: Distribution: The species is reported on the Pacific side of tropical America from the Gulf of California, Mexico, to Columbia and the Galapagos Islands from the littoral zone to a depth of 100 m. Remarks: The two species evolved from a single large population that existed before the Central American land bridge was raised separating the Atlantic and Pacific components by the closing of the Panamic seaway during the Pliocene (Chesher 1970) (Fig. 3.137). Meoma ventricosa ventricosa Lamarck, 1816 The differences between the two common species are very slight, except the geographical range. They may only be distinguished by the position of the subanal fasciole: In M. ventricosa, the distance of the upper portion of the fasciole to the periproct is equal to or greater than the vertical diameter of the periproct, whereas in M. v. grandis, this distance is shorter than the vertical diameter of the periproct (right). In direct comparison, the primary tubercles, mainly within the peripetalous fasciole, may be coarser in M. grandis than in M. ventricosa, and the Pacific species is dark brown to black, the Caribbean species more red-brown. Main characteristics of the genus: Distribution: The species is known in the Caribbean from Florida to Venezuela. No other West Indian spatangoid is found in as diverse habitats as M. ventricosa. It was collected from the intertidal zone to a depth of 200 m; in fine, silty sand to coral rubble and in biotopes like sea grass beds, coral reef tops and the deeper sea floor (Chesher 1969). Biology: M. ventricosa (and most probably also M. v. grandis) lives buried in the sediment. In fine substrates, the adult sea urchins stay near the surface, their apical spines showing above the bottom. In coarse sand, they may burrow deeper; in silt and mud, they plough half buried. They do not develop specialized tube feet in the anterior ambulacrum to build funnels for fresh water currents. In large specimens, the posterior portion of the subanal fasciole is lost and the waste is washed into the loose sand behind the animal. Usually, specimens of more than 100 mm in length come to the surface at night to feed, staying buried during daylight. With their enlarged penicillate tube feet around the peristome (forming the phyllodes), the animal grasps organic particles from the bottom of its burrow and leads them to the mouth. In contrast, young specimens burrow approximately 5 to 8 cm

below the surface of coarse sand scattered with large boulders of coral, where sufficient oxygen for respiration is available. Their subanal fasciole is still complete, and they build two short sanitary funnels by means of specialized tube feet and spines (Chesher 1969). Genus Metalia Gray, 1855 The species of Metalia resemble each other. Tab. 3.5 shows the combinations of features that are typical for the IndoWest Pacific species. Metalia is a tropical genus; eight species are described from the Indo-West Pacific, a single one from the East Pacific: Metalia angustus De Ridder, 1984. Southeast Australia. Metalia dicrana H.L. Clark, 1917. Réunion to Fiji Islands and Northern Australia. Metalia kermadecensis Baker & Rowe, 1990. Kermadec Islands. Metalia latissima H.L. Clark, 1925a. India to Japan and East Australia. Metalia nobilis Verrill, 1867. West coast of America. Metalia persica (Mortensen, 1940a). Persian Gulf. Metalia robillardi de Loriol, 1876. Madagascar to Mauritius and east coast of Australia. Metalia spatagus (Linné, 1758). Red Sea to Japan and Hawaii, East Australia. Metalia sternalis (Lamarck, 1816). Red Sea to Japan, Hawaii and Polynesia. Metalia townsendi Bell, 1904. Persian Gulf and Gulf of Oman. The genus is characterized by its shield-shaped subanal fasciole with anal branches on each side of the periproct. Test: Oval with more or less shallow anterior notch; very large forms. Apical system: Conspicuously anterior, four gonopores. Anterior ambulacrum: Narrow and more or less sunken, pore pairs simple and without specialized tube feet. Paired petals: Sunken, strongly diverging, pore zones more or less parallel. Oral side: Short, narrow labrum, more or less covering the peristome; large periproct at the posterior end; peripetalous fasciole with more or less shallow indentations in the interambulacra. Spines: Within peripetalous fasciole more or less enlarged, especially along the ambulacra; the lateral spines longer than the spatulate ones on the plastron, as indicated by distinctly larger tubercles. Remarks: The genus Metalia differs from Plagiobrissus in its lack of primary tubercles set with large spines within the peripetalous fasciole.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

D

 175

C

B

E

F

G

Fig. 3.137: (A, D) Meoma ventricosa grandis. Test length 99 mm; Gulf of California, Mexico. ZMH. (A) Aboral side: the primary tubercles are slightly coarser than in M. v. ventricosa. (D) Side view: the anterior pair of petals reaches almost to the margin. (B, C) Specimen with spines, test length 101 mm; west coast of Mexico. MfN Berlin. (B) Aboral side. (C) Oral side: the spines are rather coarse and of uniform size. (E–G) M. ventricosa ventricosa. Test length 139 mm; Florida, USA. (E) Aboral side: the anterior ambulacrum is shallowly depressed, the paired petals more deeply so. (F) Oral side: the periproct is entirely visible from the underside. (G) Side view: the upper side is hemispherically rounded. The periproct lies low on the short truncated posterior side.

176 

 3.3 Superorder Atelostomata von Zittel, 1879

Tab. 3.5: A comparison of the Metalia species and their characteristics. M. angustus

M. dicrana

M. robillardi

M. spatagus

M. sternalis

Anterior slope of test gently curved, deep notch

Anterior side steep, Anterior slope of test broad, shallow notch curved, deep notch, AIII depressed

Anterior side steep, notch deep

Anterior slope curved, virtually no notch

Anterior slope more or less steeply curved, broad notch

Five (to six) pores within subanal fasciole

Four pores within subanal fasciole

Five to eleven pores within subanal fasciole

Subanal fasciole Two points at anterior Subanal fasciole tapering to one sharp part of subanal tapering to one point fasciole point

Subanal fasciole tapering to one point

Subanal fasciole tapering to one sharp point

Subanal fasciole tapering to one point

Often heel below the periproct

More or less pointed vertex at the apical system

Subanal fasciole with Three (to four) pores four pores on either within subanal side fasciole

M. latissima

Six pores within subanal fasciole

Test broad

Metalia angustus De Ridder, 1984 Test: Rather solid, outline oval with deep anterior notch; in profile low with obliquely truncated posterior side and sharp margin; length up to 153 mm. Anterior ambulacrum: Adapically almost flush, increasing in depth toward the ambitus and continuing to the peristome. Petals: Sunken, parallel-sided, almost cross-like, the angle of the anterior pair a little larger than that of the posterior pair. Oral side: Pointed labrum covering the peristome; periproct at posterior end, uppermost portion strongly pointed and extending above the margin; often with subanal heel. Peripetalous fasciole: Shallowly indenting at the three posterior interambulacra. Subanal fasciole: Shield-shaped, strongly pointed at lower end, four pores at either side; anal branches present. Tubercles: Larger primaries border the anterior ambulacrum and are scattered over the three posterior interambulacra, mostly arranged in short, oblique bands. Spines: Short on the aboral side; on the oral side, the lateral spines are longer than the spatulate spines of the plastron. Color: In life, the sea urchin is dark red-brown with light brown to reddish spines, the bare test is mostly whitish, sometimes the test shows the individual plate boundaries. Distribution: This species is known from Bowen, Queensland to Sydney, New South Wales, along the east coast of

Posterior petals fused, inner pore pairs rudimentary or absent adapically

Australia, and from New Caledonia from the littoral zone to a depth of 25 m (Fig. 3.138). Metalia dicrana H.L. Clark, 1917 Test: Oval with shallow, broad anterior notch; in profile high and box-shaped, highest point slightly behind the apical disc or more posterior, anterior side vertical; length up to 124 mm. Anterior ambulacrum: Distinctly sunken at the margin continuing to the peristome; small undifferentiated pore pairs; no specialized tube feet. Petals: Sunken, anterior pair shorter than posterior pair at an angle of more than 180°. Oral side: With kidney-shaped mouth; labrum short and hardly covering the peristome, faintly pointed; periproct on truncated posterior end, elongated and pointed at upper and lower edge. Peripetalous fasciole: Moderately indented at the interambulacra. Subanal fasciole: Shield-shaped, anterior most portion with two points, enclosing three (rarely four) pores at either side; anal branches present. Tubercles: Larger primaries at either side of the anterior ambulacrum. Spines: Short and densely packed aborally, longer in front of the peristome on the oral surface. Color: In life, the sea urchin is brown with lighter brown spines, the bare test is whitish.

▸ Fig. 3.138: (A–C) Metalia angustus. Test length 124 mm; Sydney Harbour, New South Wales, Australia. (A) Aboral side: the peripetalous fasciole is only shallowly indented in the interambulacra. (B) Oral side: the distinct anterior notch continues to the peristome. (C) Side view: the anterior margin is sharp. At the posterior truncated side, the lower part of the periproct is often distinctly depressed, and the upper portion of the subanal area is inflated. (D–G) Metalia dicrana. Test length 62 mm; Ile de la Réunion. (D) Aboral side: the apical system lies near the anterior edge. (E) Oral side: the subanal fasciole includes three pores at either side. The anterior-most portion shows two points. (F) Side view: the test is highly inflated. (G) Anterior side.

3.3.2 Order Spatangoida L. Agassiz, 1840 



B

A

C

D

F

G

E

 177

178 

 3.3 Superorder Atelostomata von Zittel, 1879

Distribution: This species is widely found throughout the Indo-West Pacific from Ile de la Réunion and Mauritius to Samoa and Fiji Islands, and from the Philippines to Northern Australia. It prefers very shallow water between the littoral zone and 5 m. Metalia kermadecensis Baker & Rowe, 1990 The description of the holotype was based on eight fragments, being reconstructed to most of the aboral side. Test: Length approximately 110 to 115 mm; anterior margin with notch; slightly arched, highest point at the apical system; sloping to the anterior more steeply than to the posterior. Apical system: Subcentral, no plate details, broken. Anterior ambulacrum: More or less straight, with 28 pore pairs, depressed below the ambitus. Paired petals: Well-developed, depressed; anterior pair diverging at an angle of 120°; posterior pair not confluent, distally slightly bent outward, diverging at an angle of 60°. Oral side with peristome and periproct unknown.

A

B

Distribution: Raoul Island, Kermadec; 90 to 120 m. Metalia latissima H.L. Clark, 1925a Test: Large and broad, deep anterior notch; anterior side curved, upper side flattened; length up to 180 mm. Anterior ambulacrum: Conspicuously depressed. Paired petals: Slightly flexed and sunken. Oral side: Peristome anterior, periproct at the posterior steep and slightly undercut end. Peripetalous fasciole: Only shallowly indented in the lateral interambulacra. Subanal fasciole: Surrounding five to six pores on either side, pointed at the lower end. Distribution: Known in the Indo-West Pacific from India to Japan and along the east coast of Australia from the littoral zone to a depth of 74 m (Fig. 3.139). Metalia nobilis Verrill, 1867 Main characteristics of the genus: Length may reach more than 100 mm. In M. nobilis, the anterior sulcus is lacking

C

D

E

F

G

Fig. 3.139: (A–C) Metalia latissima. Test length 90 mm, Tuticorin, India. After photos in Clark (1925). (A) Aboral side: the anterior notch is deeply indented. (B) Oral side. (C) Side view. In profile, the test is compact. (D–G) Metalia nobilis. Test length 50 mm, Tabonilla, Panama. ZMUC. (D) Aboral side: the posterior pair of petals is longer than the anterior pair. (E) Oral side. (F) Side view: the upper surface is almost horizontal. (G) Specimen with spines, test length 102 mm, Taboga, Panama. ZMUC. Aboral side.

3.3.2 Order Spatangoida L. Agassiz, 1840 



similar to M. spatagus but the species differs in its longer posterior petals. In M. spatagus, the anterior and the posterior pairs of petals are of about the same length. Distribution: M. nobilis is recorded from the Gulf of California to the Bay of Panama, Pacific. It lives in the littoral and down to a depth of 10 m. Metalia persica (Mortensen, 1940) Mortensen (1940) described some brissid echinoids from the Persian Gulf as Brissopsis persica, including a specimen, which was more inflated, as a variety B. persica var. elevata. However, the specimens had divergent posterior petals, enlarged tubercles along the anterior ambulacrum, and a shield-shaped subanal fasciole. Anal branches extending from the subanal fasciole are lacking. The anterior ambulacrum is set with simple tube feet not able to build a funnel.

 179

Specialized funnel-building podia and a bilobed subanal fasciole used for managing two waste funnels are typical for the genus Brissopsis (sensu Smith et al. 2005). Based on these characteristics, the specimens have been placed in the genus Metalia by Coppard (2008). Test: Compact and rather broad, outline subcircular with distinct frontal groove; upper side rather flattened. Apical system: Anterior, four gonopores. Anterior ambulacrum: With simple podia, bordered by larger tubercles. Paired petals: Diverging, posterior pair slightly longer than anterior pair. Oral side: Convex; subanal fasciole shield-shaped, no anal branches. Distribution: Known from the Gulf of Oman and the Persian Gulf in shallow water (Fig. 3.140).

A

B

C

D

Fig. 3.140: (A–C) Metalia persica. Test length 38 mm; Persian Gulf. ZMUC ECH 261 as Brissopsis persica. (A) Aboral side. (B) Oral side. (C) Side view: the upper surface slopes toward the anterior, the highest point is posterior. (D) Metalia persica variety elevata. Test length 67 mm, Persian Gulf. ZMUC ECH 275 as B. persica elevata. Side view: the upper side is elevated, the vertex lies shortly behind the apical system.

180 

 3.3 Superorder Atelostomata von Zittel, 1879

Metalia robillardi de Loriol, 1876 Test: In outline subcircular, anterior part distinctly raised, highest point just behind the apical system, sloping gently to the short-cut posterior end. In Sydney Harbour, this species reaches a length of 124 mm. Apical system: Anterior, but not so far as in M. dicrana. Anterior ambulacrum: Broad with simple pores, bordered by larger tubercles. Paired petals: The anterior pair is directed downward at less than 180°, not backward at an angle of more than 180°. The posterior petals are more flexed than those in M. dicrana. The subanal fasciole includes five (sometimes six) pores on either side, whereas M. dicrana has three (to four) pores. Distribution: M. robillardi is found in the Indian Ocean at Mauritius, Ile de la Réunion and Madagascar, and at the east coast of Australia in the vicinity of Sydney; bathymetrical range: shallow water from 2 to 14 m. Remarks: M. robillardi differs from M. dicrana by its less anterior apical system, by the smaller angle of its anterior pair of petals, and by its lower posterior side (Fig. 3.141). Metalia spatagus (Linné, 1758) Test: Oval with very faint anterior notch; in profile rather low, highest point approximately at the apical disc, anterior side gently sloping, posterior side steep; length up to 110 mm. Anterior ambulacrum: Flush with the test; small undifferentiated pore pairs, no specialized tube feet. Paired petals: Straight and sunken, anterior pair shorter than posterior pair and at an angle of a little less than 180°. Oral side: With kidney-shaped mouth; labrum very short; plastron narrow, adjacent ambulacra parallel, periproct at truncated posterior end, elongated and pointed at upper and lower edge; peripetalous fasciole slightly indented on the interambulacra; subanal fasciole shield-shaped, lower end pointed, enclosing in adults four pores on either side; anal branches present. Tubercles: Larger primaries bordering the anterior ambulacrum and along the paired petals in short, oblique rows set with longer spines. Color: In life, the sea urchin is brown to yellowish-brown; the bare test is white, sometimes grayish with white sutures. Distribution: This species is found throughout the IndoWest Pacific from East Africa and the Red Sea to Japan and Hawaii, and in Australia from the northwest coast to Queensland and southward to Sydney. It lives from the ebb zone to a depth of 130 m. Metalia sternalis (Lamarck, 1816) Main characteristics of the genus: Test: Large and more or less box-shaped, length may reach up to 185 mm.

Petals: Posterior pair more or less fused in the adapical part, a unique characteristic in the genus Metalia. Subanal fasciole: May include up to 11 pores on either side. Distribution: Found in the Indo-West Pacific from the Red Sea to Japan, Hawaii and Polynesia, but not in New Zealand. It lives in the littoral zone and to a depth of 90 m (Fig. 3.142). Metalia townsendi Bell, 1904 This species differs from the other species of Metalia in its subcentral apical system. Other characteristics are similar for the genus. Distribution: Metalia townsendi is only known from the Gulf of Oman and the Persian Gulf, living in shallow water (Fig. 3.143). Remarks: It is very likely that there occur more species of Metalia. Van Noordenburg (2008) shows a specimen from the Philippines with a high test rising to an even higher posterior side with a vertical rear. The diverging posterior petals are almost as short as the anterior ones, which nearly spread at an angle of 180°. In addition, Miskelly (2005) also showed rather flat specimens without anterior notches and more or less subcentral apical systems from the Philippines and East Australia. Genus Neopneustes Duncan, 1889 The genus is monospecific. Neopneustes micrasteroides A. Agassiz, 1876 Test: Small, egg-shaped in outline without frontal notch; lowly domed with truncated posterior side ending in a small subanal heel; length of the type specimen 32 mm. Apical system: Ethmolytic, anterior, four gonopores. Ambulacra: All five ambulacra similar: flush with the test and nonpetaloid, plates as tall as wide; anterior ambulacrum with single and rudimentary pores, paired ambulacra with double pores, which do not alter beyond the crossing of the peripetalous fasciole. Oral side: Rather flattened; peristome far from anterior margin, kidney-shaped, labral plate extending slightly over the mouth; plastron plating unknown, labrum seems to be short and wide, sternal and episternal plates long and with parallel sides; fully tuberculated. Periproct: On truncated posterior side. Peripetalous fasciole: Faint, not indented in the interambulacra. Subanal fasciole: Shield-shaped, positioned on the rostrum. Tubercles: On the aboral side uniform. Distribution: The species is known in the Caribbean from Cuba to South Carolina; bathymetric range 320 to 420 m.

3.3.2 Order Spatangoida L. Agassiz, 1840 

A

 181

B

D

C

F E

Fig. 3.141: (A,B) Metalia robillardi. (A) Aboral side and (B) profile: the anterior notch is deeper than in Metalia dicrana. The highest point is at the distinctly raised apical disc. Sketches after photos in Miskelly (2002). (C–F) Metalia spatagus. Test length 74 mm; Negros, Philippines. (C) Aboral side: there is effectively no notch at the anterior side. (D) Oral side: three pores are visible on each side of the subanal fasciole, the fourth will develop later in growth. The fasciole tapers to a sharp point at its lower end. (E) Side view: the test is rather low with steep posterior and gently sloping anterior sides. (F) Subanal fasciole. The subanal fasciole is shield-shaped and includes three pores on either side. They are set with tube feet specialized for maintaining the sanitary funnel, into which the waste material is washed. Two anal branches are developed, which surround the periproct (upper edge).

182 

A

 3.3 Superorder Atelostomata von Zittel, 1879

B

C

D

E

Fig. 3.142: Metalia sternalis. (A, B) Test length 53 mm; Negros, Philippines. (A) Aboral side: the inner adapical pore pairs of the posterior petals are rudimentary. (B) Side view: anterior and posterior ends are steeply rounded. (C–E) Specimen with spines; test length 55 mm; Negros, Philippines. (C) Aboral side: below the spines, the fused posterior petals are recognizable. (D) Oral side: within the subanal fasciole a tuft of longer spines is developed. (E) Side view: the spines are densely arranged; at the posterior end is a subanal tuft.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

 183

B

C

D

E F

Fig. 3.143: (A–C) Metalia townsendi. Test length 56 mm; Gulf of Oman. Photos courtesy of H. van Noordenburg. (A) Aboral side. (B) Oral side. (C) Side view. (D–F) Neopneustes micrasteroides. Test length 23 mm. From Agassiz (1883). (D) Aboral side: the pore pairs do not alter at the peripetalous fasciole, but continue in the same shape. (E) Oral side. (F) Posterior side: the subanal fasciole encircles the protruding subanal heel.

Genus Plagiobrissus Pomel, 1883 The genus is characterized by an elongated, low test with a peripetalous and a shield-shaped subanal fasciole with anal branches. All interambulacra within the  ­ peripetalous fasciole are set with more or less

numerous, large primary tubercles carrying long, needlelike spines. Five species belong to the genus: Plagiobrissus (Plagiobrissus) africanus Verrill, 1871. West Africa.

184 

 3.3 Superorder Atelostomata von Zittel, 1879

Plagiobrissus (Plagiobrissus) grandis (Gmelin, 1788). Caribbean to Brazil. The subgenus Rhabdobrissus Cotteau, 1889 was established for those species, which differ from P. Plagiobrissus in having virtually no anterior notch: Plagiobrissus (Rhabdobrissus) costae (Gasco, 1876). Mediterranean and east Atlantic. Plagiobrissus (Rhabdobrissus) jullieni Cotteau, 1889. West Africa. Plagiobrissus (Rhabdobrissus) pacificus H.L. Clark, 1940. Gulf of California to Ecuador. Plagiobrissus (Plagiobrissus) africanus Verrill, 1871 Test: Low, upper side flattened, elongated oval in outline with shallow anterior depression; large form. Apical system: Ethmolytic with four gonopores. Anterior ambulacrum: Slightly depressed, pore pairs small and simple. Paired petals: Narrow and depressed, posterior pair slightly flexed outward at the end. Oral side: Flattened, peristome D- to kidney-shaped; periproct high on posterior side, not visible from below; labral plate short and wide, sternal plates long and narrow with parallel sides; plastron fully tuberculated. Fascioles: Peripetalous fasciole well-developed, subanal fasciole shield-shaped with four podia on either side, anal branches present. Tubercles: Many primaries set with long spines within peripetalous fasciole, forming horizontal series on interambulacral plates. Color: Reddish to yellowish brown. Distribution: This species is known from the West African coast, living in shallow water. Remarks: Plagiobrissus (P.) africanus differs from Plagiobrissus (Rhabdobrissus) jullieni, also native to West Africa, in having a shallow anterior notch, which is lacking in P. (R.) jullieni (Fig. 3.144). Plagiobrissus (Plagiobrissus) grandis (Gmelin, 1788) Test: Thin, outline ovate with more or less shallow anterior notch, depressed in profile; very large forms with a maximum length of 220 mm. Apical system: Anterior with four gonopores, madreporite expanding to the posterior. Anterior ambulacrum: Narrow and faintly sunken; pore pairs small and undifferentiated. Paired petals: Narrow and distinctly depressed, the anterior pair in an obtuse angle, the posterior longer and flexed outward distally. Oral side: Flattened, sternal and episternal plates symmetrical, fully tuberculated; adjacent ambulacra narrow

and subparallel; peristome kidney-shaped; periproct on short, slightly undercut, truncate posterior end. Fascioles: Peripetalous fasciole not indented behind anterior petals; subanal fasciole shield-shaped and with anal branches; primary tubercles within peripetalous fasciole, arranged in horizontal series. Spines: Primaries long and needle-like, directed to the posterior; the spines of the plastron conspicuously short, those in interambulacra 1 and 4 much longer. Color: The spines are tan-colored, the bare test is white. Distribution: Plagiobrissus grandis is widely recorded from Florida and the Bahamas to Sao Paulo, Brazil, through the Caribbean and along the central and South American coasts in the littoral zone and to a depth of 200 m, most common in less than 50 m. Biology: The species lives buried approximately 50 mm below the sea floor in sandy areas with little or no sea grass or algae. In the sediment, the primary spines are usually laid flat, but when disturbed, the animal emerges and moves rapidly over the substrate with a speed of up to 100 mm per second raising its long spines. Literature: Hendler et al. 1995. Genus Plagiobrissus (Rhabdobrissus) Cotteau, 1889 The three species in the subgenus differ generally in the arrangement of the primary spines within the peripetalous fasciole, but within each species, there occurs a certain degree of variation. Often, it is difficult to separate the species, especially younger specimens (Tab. 3.6). Plagiobrissus (Rhabdobrissus) costae (Gasco, 1876) Test: Thin, elongated, no anterior notch; depressed in profile, length may reach 100 mm. Apical system: Anterior, four gonopores; madreporite expanding to the posterior. Anterior ambulacrum: Narrow and flush, pore pairs small and undifferentiated. Paired petals: Narrow and parallel-sided, anterior petals with anterior column of pores rudimentary near the apical system; posterior ones slightly longer and flexed. Oral side: Peristome large and semilunate, periproct on truncated posterior end, pointed. Fascioles: Peripetalous fasciole surrounding the petals, not indented in interambulacra, sometimes with small crack behind the anterior paired petals; subanal fasciole shield-shaped. Tubercles: Primaries restricted to area within the peripetalous fasciole, mainly developed in the posterior columns of the lateral interambulacra (IA 1 and 4). Distribution: This species is found throughout the whole Mediterranean and along the West African coast to the Gulf

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

 185

C

B

D

E

F

G

Fig. 3.144: (A–C) Plagiobrissus (Plagiobrissus) africanus. Test length 55 mm; Dakar, Senegal. ZMUC. (A) Aboral side: within the peripetalous fasciole, many large primary tubercles are developed. The long primary spines are broken. (B) Oral side: the periproct is not visible from below. (C) Side view. (D–G) Plagiobrissus (Plagiobrissus) grandis; test length approximately 170 mm; Caribbean. (D) Aboral side: the long spines within the peripetalous fasciole are flattened against the test. After Mortensen (1950). (E, F) Specimen without spines, test length 172 mm, width 122 mm, height 45 mm; Martinique, Caribbean (HVT). (E) Aboral side: the petals are narrow and sunken. (F) Oral side: the plastron is narrow, and the periproct is visible from below. (G) Plagiobrissus (P.) grandis. Test length 133 mm, height 49 mm (profile). Tab. 3.6: A comparison of the three species in the subgenus. P. (Rhabdobrissus) costae

P. (Rhabdobrissus) jullieni

P. (Rhabdobrissus) pacificus

Primary tubercles in IA 1 and IA 4, present only in posterior column

Primary tubercles in IA 1 and IA 4, and in IA 2 and IA 3

Very few primary tubercles in IA 1 and IA 4, present in both columns

of Guinea, also at the Canaries, the Cape Verde Islands, and St. Helena. It lives at depths of 25 to 200 m (Fig. 3.145). Plagiobrissus (Rhabdobrissus) jullieni Cotteau, 1889 This species is very similar to Plagiobrissus (Rhabdobrissus) costae. It differs in the arrangement of the primary

spines: They are not concentrated in the posterior columns of the lateral interambulacra, but scattered across all interambulacral areas within the peripetalous fasciole. Distribution: Recorded in the Gulf of Guinea, West Africa, from the littoral zone to a depth of approximately 300 m.

186 

 3.3 Superorder Atelostomata von Zittel, 1879

B

A

C

D

E

F

G

Fig. 3.145: (A–C) Plagiobrissus (Rhabdobrissus) costae. Test length 105 mm, height 46 mm; Hvar, Croatia, Mediterranean. Photo courtesy of C. Neumann, MfN Berlin. (A) Aboral side: the densely packed primary tubercles in the posterior columns of the posterior paired interambulacra are well-developed. (B) Oral side: the plastron is narrow, the adjacent ambulacra subparallel. (C) Side view. (D–F) Plagiobrissus (Rhabdobrissus) jullieni. (D) Test length 32 mm; Liberia, West Africa. ZMH. Aboral side: young specimen. The large tubercles carrying long spines are distributed across all interambulacra within the peripetalous fasciole. (E, F) Specimen with spines, test length 34 mm, Cape Palmas, Liberia, West Africa. (ZMH). Young specimen. (E) Aboral side: the spination is uniform, except for the long primary spines within the peripetalous fasciole. (F) Oral side: the plastron is narrow, covered by short, flexed spines. (G) Plagiobrissus pacificus. Sketch after the photos in H.L. Clark (1948). Aboral side.

3.3.2 Order Spatangoida L. Agassiz, 1840 



Plagiobrissus pacificus H.L. Clark, 1948 Very similar to Plagiobrissus costae and P. jullieni. The species differs mainly by the distribution of the primary tubercles in the aboral interambulacra IA 1 and IA 4. There are only very few large tubercles scattered over both columns. Test: Length reaches 56 mm. Distribution: The species is found at the tropical west coast of America from the Gulf of California to Ecuador including Cocos Island; bathymetrical range 10 to 135 m. Genus Rhynobrissus A. Agassiz, 1872 The genus consists of small to moderate sized, highly vaulted species with a well-developed peripetalous fasciole, a shieldshaped subanal fasciole and anal branches. There are no pores with specialized tube feet within the subanal fasciole. Four species are with certainty included in the genus Rhynobrissus: Rhynobrissus cuneus Cooke, 1957: North Carolina, USA. Rhynobrissus hemiasteroides A. Agassiz, 1872. Hawaii and Tahiti to Australia. Rhynobrissus pyramidalis A. Agassiz, 1872. India to China Sea, North Australia. Rhynobrissus tumulus McNamara, 1982. Northwest Australia. Two additional species are named in the World Echinoidea Database. However, McNamara (1982a), having studied the genus in detail, following Mortensen (1951), thought it very likely that Rhynobrissus placopetalus A. Agassiz & H.L. Clark, 1907 is the juvenile form of R. pyramidalis. In that case, the geographical range of R. pyramidalis extends to the Hawaiian Islands. Rhynobrissus macropetalus H.L. Clark, 1938 was already damaged prior to publication. McNamara (1982) supposed that, among others, the large size and the broad anterior petals diverging strongly anteriorly suggest a closer relationship to Metalia (Tab. 3.7). Rhynobrissus cuneus Cooke, 1957 Test: Highest point posterior to apical system, sloping strongly to the rather sharp anterior margin, posterior side vertically truncate or even overhanging; outline ovoid, anterior side convex, posterior side bluntly pointed; length up to 37 mm.

 187

Apical system: Four gonopores, the anterior round, the posterior larger and diverging. Anterior ambulacrum: Narrow, flush with the test, pore pairs rudimentary. Paired petals: Rather long, of about the same size; depressed; anterior pair almost horizontally diverging, posterior pair much less diverging. Oral side: Flattened, peristome kidney-shaped and partly covered by projecting labral plate. Periproct: Near the top of the vertically truncated posterior side. Peripetalous fasciole: Well-developed, surrounding the petals parallel to the margin. Subanal fasciole: Shieldshaped, in contact with anal branches. No larger tubercles within the peripetalous fasciole. Occurrence: Only very few specimens were found in South Carolina, USA, in shallow water (Fig. 3.146). Rhynobrissus hemiasteroides A. Agassiz, 1879 Test: Very thin, oval in outline without anterior notch; highly inflated with steeply sloping anterior and posterior sides, highest point in front of the apical disc at the top of the somewhat conical area within the peripetalous fasciole; maximum length 50 mm. Apical system: Small with four gonopores. Anterior ambulacrum: Flush, pore pairs simple and very small. Paired petals: Short and deeply sunken, high keel between posterior pair, inner columns of pore pairs reduced. Oral side: Peristome kidney-shaped, labrum short and wide, plastron parallel-sided, sharply pointed at the rear. Periproct: High on top of the truncate posterior side, pointed at the upper and lower ends. Peripetalous fasciole: Broad, following the distal ends of the petals closely. Subanal fasciole: Shield-shaped, broad; lower end pointed; enclosing two pores at each side, which lie immediately within or on the fasciolar band; anal branches distinctly separated from subanal fasciole. Tubercles: Uniform, some larger ones at the vertex. Spines: On the aboral side, mainly short and uniform, some longer spines anterior of the apical system; longer

Tab. 3.7: The four species included in the genus Rhynobrissus. R. cuneus

R. hemiasteroides

R. pyramidalis

R. tumulus

Highest point posterior to apical system, test strongly wedge-shaped

Highest point: a “hump” anterior of apical system

Highest point posterior of apical system, test wedge-shaped, upper side rather flattened

Highest point posterior of apical system; upper side convex

Subanal fasciole in contact with anal branches

Subanal fasciole not in contact with anal branches

Subanal fasciole not in contact with anal branches

Subanal fasciole just in contact with anal branches

188 

A

 3.3 Superorder Atelostomata von Zittel, 1879

B

C

E D

F

I

H

G

J



on the oral side in the lateral interambulacra, short and spatulate over the plastron. Color: The spines are light brownish, the peripetalous fasciole being darker. The bare test is whitish. Distribution: R. hemiasteroides is known from Hawaii and Tahiti. In Australia, it is found along the east coast from Queensland to Sydney, New South Wales, and along the west coast. It lives in the littoral zone and down to a depth of 75 m. Biology: This species lives buried in sandy sediments to a depth of 10 mm. When uncovered, the animal immediately begins to bury itself again, and disappears from view within only 20 to 30 s. The longer spines at the apex serve for defense (McNamara 1983). Rhynobrissus pyramidalis A. Agassiz, 1872 Test: Highest point posterior to apical system, sloping to the rounded anterior margin, posterior side truncate; outline oval, tapering toward the posterior; anterior side convex; length up to 50 mm. Apical system: With four gonopores. Anterior ambulacrum: Narrow, flush with the test, pore pairs very small. Paired petals: Depressed, anterior pair spread at an angle of 140°; pore pairs of inner series reduced. Oral side: Flattened, scarcely sunken toward the kidneyshaped peristome, labral plate very short, plastron distinctly keeled toward the lower point of the subanal fasciole, fully tuberculated; plates of paired posterior interambulacra not contacting the peristome. Periproct: Near the top of the vertically truncated posterior side. Fascioles: Peripetalous fasciole well-developed; subanal fasciole shield-shaped, anal branches broad, almost encircling the periproct, not connected to the s ubanal fasciole. No larger tubercles within the peripetalous fasciole. Distribution: The species is recorded in the Indo-West Pacific from India to Singapore and Thailand and to southern China and Taiwan, in shallow water to depths of approximately 200 m. Remarks: Mortensen (1951) and McNamara (1982a) think that R. placopetalus A. Agassiz & H.L. Clark, 1907 may most probably be a young R. pyramidalis (Fig. 3.147).

3.3.2 Order Spatangoida L. Agassiz, 1840 

 189

Rhynobrissus sp. From the East China Sea, another kind of Rhynobrissus is reported. It differs in the following features: Test: It is lower in the whole and more rounded, not pointed toward the posterior; aboral side slopes rather straight toward the anterior rounded margin, not arched; highest point is very close to the posterior side, just before it falls down almost vertically. Petals: Anterior pair of petals extends toward the sides horizontally; in R. pyramidalis, it spreads at an angle of approximately 140°. Subanal fasciole: Not in contact with anal branches; a single pore pair to each side. This specimen might be a variety of R. pyramidalis or perhaps a subspecies. Further studies are needed (Fig. 3.148). Rhynobrissus tumulus McNamara, 1982 Test: Ovoid, highly vaulted, highest point posterior of subcentral apical system; aboral side gently convex, posterior side steep but rounded; length up to 60 mm. Anterior ambulacrum: Flush with very small pore pairs. Paired petals: Slightly depressed, anterior pair diverging horizontally, shorter than posterior pair. Oral side: Gently convex, peristome broadly semilunar, labral plate very narrow, plastron conspicuously keeled. Periproct: High at the posterior side, small and longitudinally oval. Peripetalous fasciole: Encircling the distal ends of the petals parallel to the margin. Subanal fasciole: Well-developed, just in contact with the almost closed anal fasciole. Tuberculation: Aborally fine and uniform, except some slightly larger tubercles just in front of the apical system, probably bearing a tuft of longer spines. Spines: On the plastron, spatulate serve for burrowing. Distribution: This species is endemic to the southern part of Australia, recorded from Fremantle on the west coast to Port Jackson in the east, living in shallow water down to depths of 50 m Genus Taimanawa Henderson & Fell, 1969 The genus is characterized by large to very large tests with depressed aboral interambulacra, on which a few very

◂ Fig. 3.146: (A–D) Rhynobrissus cuneus. Holotype, test length 37 mm. Sketches after photos in Cooke (1959). (A) Aboral side. (B) Oral side. (C) Posterior side. (D) Side view. (E) Rhynobrissus hemiasteroides. Plate pattern of oral side. Scale bar is 5 mm, after Mortensen (1951). The interambulacra 1 and 4 (shaded) are excluded from the peristome. The labral plate (green) is very short. (F–J) R. hemiasteroides. (F–H) Length of test 41 mm; Queensland, Australia. (F) Aboral side: the area within the peripetalous fasciole is conically raised to the apex, which is set with larger tubercles. (G) Oral side: the plastron is long and narrow; the tubercles being very regularly arranged. (H) Side view. (I, J) Specimen with spines, test length 39 mm; Sydney Harbour, New South Wales, Australia. (I) Aboral side: the petaloid area surrounded by the peripetalous fasciole is relatively small. (J) Oral side: the plastron is parallel-sided and set with short, spatulate spines.

190 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

C

D

E

F

G

Fig. 3.147: Rhynobrissus pyramidalis. (A–C) Test length with spines 45 mm; off northwest Taiwan. (A) Aboral side. (B) Oral side: the lateral spines are long and slender, those on the plastron have a paddle-like distal end. (C) Side view: the fine, silky spines cover the test as densely as a fur. Below them, the water currents flow over the epidermis and, maintained by the fascioles, provides the podia with oxygen when the animal burrows through the substrate. (D–G) Specimen without spines, test length 27 mm; northwest of Taiwan. (D) Aboral side: the peripetalous fasciole encircles the petals parallel to the margin. (E) Oral side: the posterior end is pointed. (F) Side view: the upper side is arched from the high posterior side toward the rounded anterior margin. (G) Posterior side: the subanal and anal fascioles are broad. There are no pores developed nor associated tube feet.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

B

 191

C

D

E

F

G

Fig. 3.148: (A–D) Rhynobrissus sp. Test length 41 mm, Northeast Taiwan. (A) Aboral side. (B) Oral side. (C) Side view. (D) Posterior side: both anal and subanal fasciole are broad. They are not in contact. One pore to each side is developed within the subanal fasciole. (E–G) Rhynobrissus tumulus. Test length 41 mm, Broome, Western Australia. Photos courtesy of H. van Noordenburg. (E) Aboral side. The anterior pair of petals extends horizontally to the sides. (F) Oral side. (G) Side view. The upper side is inflated and regularly arched.

conspicuous primary tubercles occur being restricted to the posterior column of the interambulacra within the peripetalous fasciole. A subanal fasciole is present, sometimes also traces of an internal fasciole.

Besides a few fossil species from the Tertiary of New Zealand, two extant species from the West Pacific are described. There will probably be additional species. Taimanawa mortenseni Henderson & Fell, 1969. Indonesia.

192 

 3.3 Superorder Atelostomata von Zittel, 1879

Taimanawa relictus Shigei, 1975c. Japan and East China Sea. Taimanawa sp. Philippines. Taimanawa mortenseni Henderson & Fell, 1969 Described from fragments, the largest has a width of 105 mm. Test: Seems to be large and approximately as wide as long with a deep anterior notch; moderately inflated with rounded margin. Anterior ambulacrum: Adapically shallowly depressed, increasing in depth toward the frontal margin; minute pore pairs. Petals: Anterior petals slightly depressed, spread at an angle of approximately 120°, weakly flexuous; adapical pore pairs rudimentary; posterior petals and apical system unknown. Oral side: Mostly unknown. Fascioles: Peripetalous fasciole well-developed, not indented on the interambulacra, disappearing on anterior ambulacrum; thin internal fasciole from anterior of the apical system downward bordering half the anterior ambulacrum; subanal fasciole well-developed, enclosing seven pore pairs on each side. Tubercles: Very conspicuous, large primary tubercles cover the posterior column of the anterior interambulacrum within the peripetalous fasciole, the areoles surrounded by a distinct ring of scrobicular tubercles. Occurrence: Kei Islands, Indonesia, at a depth of 260 m (Fig. 3.149). Taimanawa relictus Shigei 1975c Test: Large, nearly as broad as long, upper side more or less lowly arched, anterior ambulacrum depressed continuing to a deep notch at the more or less sharp margin; posterior side truncated, overhanging the periproct. Apical system: Small, slightly anterior; four gonopores of equal size. Anterior ambulacrum: Increasingly sunken to a very conspicuous anterior notch; small pore pairs in a regular series, small tubercles bordering the sides. Paired ambulacra: Petaloid, more or less sunken forming the posterior limit of the depressed adjoined interambulacra; distally open; anterior pair slightly curved and longer than the posterior pair; pore pairs adapically rudimentary, especially in the anterior series. Interambulacra: IA 2 and IA 3 distinctly sunken, anterior pair deeper than the posterior pair; set with three to four large tubercles surrounded by upset areoles restricted to the posterior column. In the holotype, there is additionally a single tubercle on the posterior interambulacrum IA

1 and IA 4 in the posterior columns; in the specimen here, those tubercles are lacking. Oral side: Flat, distinct keel from the peristome to the posterior end. Peristome: Semilunar, overarched by a distinct labral lip; sternum slightly bowed, naked areas of adjacent ambulacra rather broad. Periproct: Ovoid, pointed above, clearly visible from the oral side. Fascioles: Peripetalous fasciole not indented between the petals; narrow inner fasciole(?), subanal fasciole bilobed, broad, encircling four to five pore pairs on each side. Primary tubercles: Large, crenulated, and perforated; length of primary spines not known. Color: The test and the spines are yellowish-white, the skin of the base of the spines reddish-brown. Distribution: This species is recorded from Japan, Sagami Bay to the China Sea in deeper water. Remarks: The specimen here differs slightly from the holotype described by Shigei. Both specimens are about the same size with a length of 75 and 78 mm, respectively. i. The posterior aboral interambulacra are not set with a large tubercle, which is present in the holotype. ii. The inner fasciole, narrow, but distinct here, is not mentioned by Shigei in his detailed description. Henderson & Fell (1969) report on a narrow inner fasciole visible on some fragments of T. mortenseni. Taimanawa sp. Distribution: Philippines; depth recorded 250 m.

3.3.2.4 Family Palaeotropidae Lambert, 1896 The family is characterized by ambulacra that are apetaloid or even rudimentary. Test: Very delicate, ovoid or ovate in outline without an anterior notch. Apical system: Ethmolytic with two, three, or four gonopores; genital plates often fused. Labral plate: Formed like a mushroom and rather short, at most reaching to the rear of the second ambulacral plate. Subanal fasciole: Present, or at least traces of it, sometimes only in young specimens. being lost in adults. Tubercles: Aboral surface with scattered small primary tubercles among finer granules. The family consists of five genera: Kermabrissoides Baker, 1998. Kermadec Islands, New Zealand. Monospecific.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

C

 193

B

E

D

Fig. 3.149: (A) Taimanawa mortenseni. Width of grouped fragments 120 mm; Kei Islands, Indonesia. ZMUC ECH 344. Part of aboral side: the anterior ambulacrum is deeply depressed. (B–D) Taimanawa relictus. Test length 78 mm, East China Sea. Photos by Bas van der Steld. (B) Aboral side: there is a peripetalous fasciole around the petals and a narrow inner fasciole around the apical system, continuing on both sides of the anterior ambulacrum. (C) Oral side. (D) Side view: the anterior margin is rather sharp; posteriorly, the margin is overhanging the periproct. (D) Taimanawa sp. Test length 161 mm, Mactan, Philippines. Aboral side. By its large size, this specimen resembles more T. mortenseni. Photo courtesy of H. van Noordenburg.

Palaeobrissus A. Agassiz, 1883. Caribbean. Monospecific. Palaeotropus Lovén, 1872. Azores, Bay of Biscay, West Indies. Two species. Paleotrema Koehler, 1914. Malayan region. Two species.

Scrippsechinus Allison, Durham & Mintz, 1967. East Pacific. Monospecific. Literature: Agassiz 1883; Allison et al. 1967; van Noordenburg 2008.

194 

 3.3 Superorder Atelostomata von Zittel, 1879

The four, mostly monospecific, genera are very similar in general shape, but well defined by plate pattern and development of pores in the ambulacra (Tab. 3.8): Genus Kermabrissoides Baker, 1998 The original name was Acanthotrema but had to be changed because it was already taken. Kermabrissoides siculum (Baker & Rowe, 1990) Test: Oval, low, very fragile; upper side slightly arched, posterior side vertically truncated; anterior side rounded, no notch; length may reach 25 mm. Apical system: Four genital pores on top of small cones. Ambulacra: All flush, with double rows of plates, pore pairs mince, usually obliquely aligned, sometimes single pores. Oral side: Depressed around the D-shaped peristome; anterior peristomial plates each with a rugose, spine-like extension that protrudes from the opening; labrum long and dagger-shaped, reaching beyond the middle of the third adjacent ambulacral plate; short, strongly tuberculated plastron. Periproct: At the vertical posterior side, transversely oval, plates may have thorny extensions. Fasciole: Subanal fasciole shield-shaped, with two tube feet to either side. Tuberculation: On the aboral side usually small, but up to six larger tubercles on the frontal and lateral interambulacra; longest primary spine 8 mm, curved at the base. Occurrence: Kermadec Islands, New Zealand; depth 256 to 347 m (Fig. 3.150). Genus Palaeobrissus A. Agassiz, 1883 Palaeobrissus hilgardi A. Agassiz, 1883 Test: In outline regularly oval, no frontal notch; in profile low with thickly rounded margin, vertex central or posteriorly higher than anteriorly; maximum length 47 mm. Apical system: Anterior; two gonopores in the posterior genital plates surrounded by a rim; in larger specimens, two additional smaller anterior gonopores; genital plates fused, but according to Smith (2005) basically ethmolytic.

Ambulacra: All flush with hexagonal plates; remains biserial adapically; pores in anterior ambulacrum single, paired ambulacra apetaloid with double pores, reaching in large specimens almost to the ambitus. Oral side: Flattened; peristome not sunken and transversely oval; periproct at the subvertical posterior side, round or slightly oval. Labral plate: Mushroom-shaped, slightly projecting, a few larger tubercles; sternal plates triangular, short, fully tuberculated. Subanal fasciole: Rounded, with only one tube foot to each side. Tuberculation: Aborally heterogeneous. Color: The color is stated by Agassiz (1883) as dark greenviolet or greenish red, the smaller specimens being more pink. Occurrence: West Indies; 150 to 240 m. Genus Palaeotropus Lovén, 1871 The genus included two species. Palaeotropus josephinae (Lovén, 1871) Test: In outline regularly oval, no frontal notch; aboral side gently arched, anteriorly with round margin, posteriorly vertically truncated; length usually 18 to 20 mm, exceptionally 25 mm. Apical system: Anterior; two gonopores in the posterior genital plates; genital plates fused, a few hydropores restricted to the anterior part of disc. Ambulacra: All flush; plates higher than wide; adapically in a single row. Paired ambulacra: Apetaloid with microscopic pores. Oral side: Labral plate mushroom-shaped, straight cut at the peristomial edge; sternal plates bowed and almost fully tuberculated; distinct keel toward the posterior; peristome D-shaped, not sunken. Periproct: At the subvertical posterior side, pointed at the upper end. Subanal fasciole: Broad and subcircular. Primary tubercles: Rather small, scattered in finer granules, a row of larger tubercles bordering the frontal ambulacrum. Distribution: The species is known with certainty only from the Azores and the Bay of Biscay and the Gulf of

Tab. 3.8: A comparison of four genera from the Palaeotropidae family Palaeobrissus

Palaeotropus

Paleotrema

Scrippsechinus

Ambulacral pores double

Ambulacral pores single

Ambulacral pores single

Ambulacral pores double

Double row of ambulacral plates adapically

Uniserial ambulacral plates adapically

Double row of ambulacral plates adapically

Uniserial ambulacral plates adapically

3.3.2 Order Spatangoida L. Agassiz, 1840 



B

A

C

 195

D

E

Fig. 3.150: Palaeobrissus hilgardi. (A–C) Test length 22 mm, Barbados, West Indies. ZMUC. (A) Aboral side: there are two gonopores surrounded by a rim, and double pores in the double rows of ambulacral plates. (B) Oral side: the slightly projecting labral plate and the symmetrical sternal plates are tuberculated. (C) Side view. (D) Adapical area: there is a double row of ambulacral plates adapically (interambulacra shaded). Scale bar is 5 mm. After Mortensen (1950). (E) Peristomial area: the labrum is formed like a mushroom and slightly arched at the peristomial edge. Scale bar is 5 mm. After Mortensen (1950).

Mexico, recorded from seamounts and knobs. Bathymetric range 350 to 1,600 m (Fig. 3.151).

Distribution: Atlantis seamount, northeast Atlantic; 1,580 to 1,900 m.

Palaeotropus uniporum Mironov, 2006 The species is based on a single incomplete specimen. The main characteristics are typical of the genus, but the single pores on the apical system (unipores) are in contrast with those P. josephinae. Oral side: Labrum reaching second pair of ambulacral plates. Spines: Miliary spines curved. Pedicellariae: Absence of rostrate pedicellariae.

Genus Paleotrema Koehler, 1914 The genus is characterized by ambulacra with plates developed in double rows up to the apical system. The plates are perforated only by a single pore. The genus consists of two species: Paleotrema loveni A. Agassiz, 1879. Malayan Archipelago and the Philippines. Paleotrema ovatum Koehler, 1914. Indian Ocean.

196 

A

 3.3 Superorder Atelostomata von Zittel, 1879

B

C

D

E

F Fig. 3.151: Palaeotropus josephinae. (A–D) Test length 12 mm, Azores. After Lovén (1884). (A) Aboral side. (B) Oral side. (C) Posterior side. (D) Profile. (E) Apical region: the ambulacral plates form a single row toward the apical disc. (Interambulacra shaded). Scale bar is 4 mm. After Mortensen (1950). (F) Peristomial area: the labrum is straight cut at the peristomial edge. Scale bar is 3 mm. After Mortensen (1950).

The species differ mainly by the shape of the test: in Paleotrema loveni, the posterior end is obliquely truncated, the periproct visible from above, whereas in Paleotrema ovatum, it is subvertical, the periproct not visible from above. Paleotrema loveni A. Agassiz, 1879 Test: Very delicate, in outline more or less regularly oval, anterior side convex or straight, no anterior notch; in profile low arched, anterior end with thickly rounded margin, posterior end higher than anteriorly and obliquely truncated; length may reach 26 mm. Apical system: Conspicuously anterior, slightly sunken; fused genital plates, three gonopores. All ambulacra flush and apetaloid; plates in double rows adapically. Ambulacra: Anterior one with small pores, adapically single, more anteriorly becoming double, placed one above the other; paired ones with minute pores; rudimentary and distinctly smaller than in anterior ambulacrum.

Oral side: Slightly concave, posterior plastron forming a distinct keel; naked area relatively broad; peristome D-shaped; periproct supramarginal on the posterior surface; labral plate elongate, upper and lower end widened, straight cut at the peristomial edge; sternal plates bowed and symmetrical; fully tuberculated. Fasciole: Subanal fasciole distinct, rounded; two pore pairs to each side. Tuberculation: Fine and uniform with a few small primary tubercles aborally, some larger tubercles border the anterior ambulacrum above the margin. Primary spines: May reach the length of half the test, spines on the sternal plates widened distally. Color: The test and spines are white to purplish or pink, the periproct is often conspicuously purple. Distribution: The species is recorded from the Philippines and the Malayan region; bathymetric range 175 to 685 m (Fig. 3.152).

▸ Fig. 3.152: Paleotrema loveni. (A–C) Test length 20 mm, Philippines. ZMUC ECH 198. (A) Aboral side: there are three gonopores, the frontal slope is set with large primary tubercles. (B) Oral side: the peristome is surrounded by short phyllodes, the sternum is fully tuberculated. (C) Test length 19 mm, Philippines. ZMUC. Side view. (D–G) Specimen with spines: test length 18 mm, Philippines. ZMUC. (D) Aboral side: the spination is rather sparse. (E) Oral side: the naked areas on the posterior ambulacra are broad. (F) Apical area: the ambulacral plates are double toward the apical disc, perforated by rudimentary single pores (interambulacral plates shaded). (G) Labrum, peristome, and adjacent ambulacral plates. Scales bar is 5 mm. After Mortensen (1950).

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

B

C

D

E

F

G

 197

198 

 3.3 Superorder Atelostomata von Zittel, 1879

Genus Scrippsechinus Allison, Durham & Mintz, 1967 The genus is monospecific. Scrippsechinus fisheri Allison, Durham & Mintz, 1967 Test: Thin and delicate; ovoid in outline without anterior notch; highly vaulted with rounded margin; length up to 81 mm. Apical disc: Subcentral; ethmolytic with four gonopores. All ambulacra flush, plating to varying degrees; uniserial near the apical system. Ambulacra: Anterior one with small and oblique pore pairs; paired ambulacra apetaloid with small pore pairs— slightly larger than those in ambulacrum III—weakly enlarged on adapical plates. Oral side: Flat, peristome large, kidney-shaped, and sunken, facing obliquely toward the anterior; labral plate short and broad, lip projecting over the peristome; sternal plates bowed and symmetrical. Periproct: Inframarginal, vertically elongated and pointed above and below. Subanal fasciole: None developed; however, Allison et al. (1967) reported on traces of it in the form of a faint ridge. Tubercles: Aborally small and uniform; orally curved and widened distally. Occurrence: The species is only recorded from seamounts between the Nasca Ridge, off Peru, and the Easter Islands in the east Pacific at a depth of approximately 200 to 400 m (Fig. 3.153).

Superfamily Spatangoidea 3.3.2.5 Family Eupatagidae Lambert, 1905 The apical system of this family is ethmolytic with three or four gonopores. It consists of only one genus. Genus Eupatagus L. Agassiz in L. Agassiz & Desor, 1847 Test: Usually low (except in E. rubellus and E. flindersi), upper side flattened; anterior notch faint or lacking. Anterior ambulacrum: Flush and narrow, pores small and simple. Paired petals: Flush, pore zones more or less lanceolate, narrowing adapically and distal. Oral side: Flat, keeled toward the posterior; peristome large and kidney-shaped; labrum elongated and wedgeshaped, nearly as long as sternal plates; sternum fully tuberculated. Periproct: On short truncated posterior side, large. Fascioles: Peripetalous and shield-shaped subanal fasciole well-developed.

Primary tubercles: Large in interambulacra within peripetalous fasciole, without sunken areoles; enlarged tubercles border anterior ambulacrum. Literature: Döderlein 1906; McNamara 1986; Baker & Rowe 1990; Rowe & Gates 1995; van Noordenburg 2008. Six species are included in the genus: Eupatagus flindersi Baker & Rowe, 1990; South Australia to Tasmania. Eupatagus lymani Lambert & Thiery, 1924; Hawaii and ­Philippines. Eupatagus micropetalus H.L. Clark, 1917; Japan, Philippines, and Malayan region. Eupatagus obscurus (A. Agassiz & H.L. Clark, 1907); Hawaii. Eupatagus rubellus Mortensen, 1948. Philippines, Malayan region to Northwest Australia. Eupatagus valenciennesi L. Agassiz in L. Agassiz & Desor, 1847. Southeast coast of Australia. Eupatagus flindersi Baker & Rowe, 1990 Test: Delicate, regularly oval, width not much smaller than length (in holotype 83%); aboral side high, not flattened, sloping steeper toward the anterior than toward the posterior end. Oral side: Labrum long, naked areas on oral posterior ambulacra relatively narrow. Fascioles: Peripetalous fasciole thin, shallow indentations in lateral interambulacra, disappearing anteriorly; subanal fasciole broader, encircling a distinctly protruding plastron with four pore pairs per column. Primary tubercles: Restricted to the anterior interambulacra. Distribution: Endemic to Southern Australia and Tasmania; depth 25 m (Fig. 3.154). Eupatagus lymani Lambert & Thierry, 1924 Test: Low, upper side flattened, with the posterior interambulacrum slightly elevated, anterior margin rounded; posterior side vertical; in outline ovoid, distinctly tapering to the posterior truncated end, no frontal sulcus; length may reach 72 mm. Paired petals: Anterior ones spread at almost 180°, posterior pair longer, pores small. Oral side: Flattened; naked areas on posterior ambulacra and around the peristome broad, posterior part of plastron raised and sharply keeled toward the posterior end; peristome D-shaped with a straight cut labrum. Periproct: Circular at the truncated posterior side. Fascioles: Peripetalous fasciole thin, not indented laterally, disappearing in front of the anterior petals; subanal fasciole heart-shaped with four pore pairs to each side.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

B

E

 199

D

C

F

G

Fig. 3.153: Scrippsechinus fisheri. (A–D) Test length 54 mm; Nasca Ridge, off the Chilean coast, East Pacific. (A) Aboral side (interambulacra shaded). (B) Oral side. (C) Side view. (D) Apical system. Modified after the drawings of Allison et al. (1967). (E–G) Test length 81 mm; Nasca Ridge, off Chile. ZMUC. (E) Aboral side. (F) Oral side: the lip strongly projects over the peristome. (G) Side view: the test is strongly inflated. In the center of some interambulacral plates, the nodes are positioned.

Tubercles: Some large primary tubercles scattered on the paired interambulacra within the peripetalous fasciole; in large specimens, there may be numerous primary tubercles; secondary and miliary tuberculation not uniform, leaving naked areas mainly along the plate sutures. Spines: Very slender and curved. Color: The spines and the test may be aborally creamyellow to reddish-pink with lighter plate sutures, the oral side is white.

Distribution: Known in the Philippines and the Hawaiian Islands at a depth of 95 to 425 m. Eupatagus micropetalus H.L. Clark, 1917 Test: Low, flattened above and below, anterior margin rounded, posterior side steep but rounded toward the oral side; outline regularly oval, anterior side slightly depressed, posterior end truncated; maximum length 70 mm.

200 

A

 3.3 Superorder Atelostomata von Zittel, 1879

B

C

D

E

F

G

H

Fig. 3.154: (A–C) Eupatagus flindersi: (A, B) Sketch after the photos in Baker & Rowe (1990). (C) Labral plate. Scale bar is 5 mm. (D–F) Eupatagus lymani. Test length 45 mm; Philippines. (D) Aboral side: the primary tubercles are developed in all paired interambulacra. (E) Oral side: the naked areas are broad. (F) Side view: the test is low with a truncated end. (G, H) Specimen with spines. Test length 49 mm, Philippines: (G) Aboral side: the pointed aboral primary spines reach more than half the test length. (H) Oral side: the long and slender lateral spines border a broad naked area. Photos courtesy of Henk van Noordenburg.

Paired petal: Rather short and narrow; in the anterior pair, the pore pairs of the anterior column are distinctly smaller than the posterior column; posterior pair of petals longer. Oral side: Keeled toward the posterior end; very broad naked area on posterior ambulacra and around the peristome; labrum relatively short, straight cut; peristome D-shaped. Periproct: Subcircular and positioned on the truncated posterior face.

Fascioles: Peripetalous fasciole thin, hardly indenting laterally; subanal fasciole heart-shaped with four pore pairs on each side. Primary tubercles: None or very few large primary tubercles on the anterior interambulacra within the fasciole, some more on the posterolateral interambulacra; very slender and curved, they may reach half the test length.

3.3.2 Order Spatangoida L. Agassiz, 1840 



Color: The test is grayish, the spines can be silvery to light purple with faint bands. Distribution: This species is recorded from the Malayan region over the Philippines to Japan; bathymetrical range 80 to 400 m (Fig. 3.155). Eupatagus obscurus (A. Agassiz & H.L. Clark, 1907) Test: Low; upper and lower side almost parallel; shallow, distinct frontal sulcus; several primary tubercles in posterior interambulacra within the peripetalous fasciole. This species differs from Eupatagus lymani, which also occurs around Hawaii, in having three pores within the

A

B

 201

sub­anal fasciole at each side, whereas E. lymani has four. The species is endemic to Hawaii; bathymetric range 365 to 600 m. Eupatagus rubellus Mortensen, 1948 Test: High and egg-shaped, in contrast to the other species of Eupatagus; oral side convex, posterior interambula­ crum keeled; length a little more than 36 mm. Primary tubercles: No primary tubercles in the posterior interambulacrum. Fascioles: Peripetalous fasciole closely following the distal parts of the paired petals; subanal fasciole shieldshaped.

C

D

E

F

G

H

Fig. 3.155: (A–C) Eupatagus micropetalus. Young specimen, test length 31 mm; Sagami Bay, Japan. ZMUC: (A) Aboral side: the petals are short and narrow. (B) Oral side: the naked areas on the posterior ambulacra and around the peristome are very broad. (C) Side view: the posterior side is steeply truncated, the periproct just visible from above. (D) E. micropetalus: the labrum is straight cut. Scale bar is 3 mm; after Mortensen (1951). (E–H) Eupatagus rubellus. Test length 36 mm; Shark Bay, Western Australia: (E) Aboral side. (F) Oral side. All aboral ambulacra are flush with the test. The plastron is short. (G) Side view. (H) Plate pattern of the oral side. The labrum is short and straight cut. Scale bar is 4 mm; after Mortensen (1951).

202 

 3.3 Superorder Atelostomata von Zittel, 1879

Color: The species has spines banded in red with yellowish, or red-brown with whitish; the test is also reddish or redbrownish with yellowish or whitish edges along the sutures. Distribution: The species is recorded from the Philippines and recently from Northwest Australia in depths of 10 to 25 m and probably deeper (Fig. 3.156). Eupatagus valenciennesi L. Agassiz in L. Agassiz & Desor, 1847 General characteristics like genus; test length not much more than 66 mm. Test: Low. Oral side: Flattened, posterior interambulacrum keeled; labrum long, toward the sternum wedge-shaped. Fascioles: Peripetalous fasciole without indentations; subanal fasciole transversely oval. Color: In life brownish pink, within the peripetalous fasciole darker; bare test light brown. Distribution: Bass Street and east coast of Australia; depth 10 to 65 m. Biology: The species is brood-protecting. The female retains her growing young below the appressed primary spines at the apical pole (McNamara 1994).

3.3.2.6 Family Eurypatagidae Kroh, 2007 This taxon is characterized by the following features: Apical system: Ethmolytic with four gonopores. Petals: Very long and subparallel, distally open. Oral side: Ambulacral plates AIV enlarged and indenting the posterior interambulacrum. Subanal fasciole: Shield-shaped, present at least in young specimens. Literature: Koehler 1912; Baker & Rowe 1990; Rowe & Gates 1995; van Noordenburg 2008. Five genera are established in this taxon. Elipneustes Koehler 1914. India and Sri Lanka. Two species. Eurypatagus Mortensen, 1948. Indian Ocean and Philippines. Three species. Linopneustes A. Agassiz, 1881. Indo-West Pacific and West Indies. Six species. Paramaretia Mortensen, 1950. Malayan Region. Monospecific. Platybrissus Grube, 1865. East Australia and New Zealand. Monospecific. Genus Elipneustes Koehler, 1914 The genus consists of two species: Elipneustes denudatus Koehler, 1914. Sri Lanka. Elipneustes rubens Koehler, 1914. South India and Sri Lanka.

Elipneustes denudatus Koehler, 1914 Test: Oval, tapering to the posterior; faint anterior notch; in profile lowly vaulted; short, obliquely truncated posterior side; length may reach 104 mm. Apical system: Anterior, ethmolytic, four gonopores. Anterior ambulacrum: Flush; simple pores. Paired ambulacra: Petaloid, petals very long extending to the ambitus, with parallel sides, broad interporiferous zones, distally open; pore pairs conjugate. Oral side: Flat, peristome D-shaped, slightly sunken; periproct on truncated posterior side, transversely elongate; labral plate wedge-shaped, long and narrow, sternal plates short and triangular, fully tuberculated. Fascioles: Peripetalous fasciole thin, not indented late­ rally subanal fasciole with three tube feet to each side. Larger primary tubercles aborally scattered in all interambulacra. Distribution: Off Sri Lanka, 285 m. Elipneustes rubens Koehler, 1914 The two species differ in general size: in E. rubens, the length reaches only 53 mm. In E. denudatus, the petals are longer than in E. rubens, and the labrum is shorter. The posterior side is more oblique in E. denudatus and not subvertical as in E. rubens, where the distinct subanal fasciole is positioned more on the oral surface. Distribution: Off South India and Sri Lanka, depth 125 to 270 m (Fig. 3.157). Genus Eurypatagus Mortensen, 1948 This genus differs from Elipneustes by the lack of a peripetalous fasciole. A subanal fasciole is only present in young specimens. Eurypatagus has long, narrow, distally open petals with almost parallel pore zones. Maretia differs in having broad, distally closed petals. Eupatagus differs by its petals being short and bowed. Test: In outline ovoid without frontal notch; in profile low with flattened aboral side, posterior side short and truncated. Apical system: Anterior, ethmolytic with four gonopores. Anterior ambulacrum: Flush, pore pairs very small, placed vertically; upper plates with simple pores. Paired ambulacra: Flush and petaloid; petals extending to the ambitus, anterior pair flexed anteriorly, posterior pair more straight, pore zones subparallel; pore pairs conjugate; no occluded plates. Oral side: Flat, peristome slightly sunken; phyllodes strongly developed; peristome kidney-shaped and large, relatively distant from anterior margin; labral plate long and wedge-shaped; may be separated from the sternum by adjacent ambulacral plates; sternal plates short and triangular, tuberculated only posteriorly.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

C

 203

B

E

D

F

Fig. 3.156: (A, B) Eupatagus valenciennesi. Test length 66 mm; Bass Strait, Australia. ZMUC: (A) Aboral side. The petals are long. (B) Oral side. The naked areas are narrow. (C) Apical system: the madreporite does not expand very far to the posterior. Scale bar is 1 mm. (D) Oral side with labrum and adjacent ambulacral plates. Scale bar is 5 mm. After Mortensen (1951). (E) Aboral side: the paired petals are flexed and taper toward the apical disc. Length 55 mm. (F) Side view.

204 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

C

D

E

H

F

I

G

J

K

Fig. 3.157: (A–D) Elipneustes denudatus. Test length 89 mm; Ceylon: (A) Aboral side. (B) Oral side. (C) Side view. (D) Anterior side. From Koehler (1914). (E–G) Eurypatagus, part of posterior left ambulacrum. Scale bars are 5 mm, after Mortensen (1950): (E) Eurypatagus grandiporus. (F) Eurypatagus ovalis. (G) Eurypatagus parvituberculatus. (H–K) Eurypatagus grandiporus: (H) Test length 66 mm, Gulf of Aden, Indian Ocean. ZMUC ECH 150, aboral side. (I–K) Specimen with spines, length 65 mm; Gulf of Aden. ZMUC: (I) Aboral side: the upper surface is reddish, the spines being paler. (J) Oral side: the peristome lies rather distant from the anterior margin. Only the very posterior end of the sternal plates has some scattered tubercles. (K) Side view.



Periproct: Large, on short truncated posterior face. Fasciole: Subanal fascioles in young specimens present, but lost in adults. Primary tubercles: Larger ones scattered over aboral interambulacra; perforated and crenulated. on oral side large aligned tubercles with sunken areoles, bordering late­rally the broad naked area. Primary spines: Long, slender, at the base curved with distant thorns distally. Color: All species are more or less reddish. Three species, which are rather similar, are included in this genus: Eurypatagus grandiporus Mortensen, 1948. Cape Gardafui, Gulf of Aden. Eurypatagus ovalis Mortensen, 1948. Philippines. Eurypatagus parvituberculatus (H.L. Clark, 1924). South Africa, Mauritius. The morphological differences are not conspicuous, but the geographic ranges leave a good distinction. Eurypatagus grandiporus Mortensen, 1948 Main characteristics like genus: maximum length 73 mm. Test: Low and slightly arched aborally, posterior interambulacrum hardly keeled. Paired petals: Pore pairs very conspicuous, anterior row of anterior petals may be rudimentary toward the apical system. Primary tubercles: Rather few primary tubercles scattered over the aboral interambulacra; secondary and miliary tubercles scarce. Occurrence: Gulf of Aden, Indian Ocean at a depth of 366 m. Eurypatagus ovalis Mortensen, 1948 Main characteristics like genus: test may reach 73 mm. Posterior interambulacrum only slightly raised, the aboral side is lowly arched. Smaller pore pairs in paired petals than in E. grandiporus; more numerous primary tubercles on the aboral side, tending to form horizontal series above the ambitus. Color: Spines and test more or less bright red on the aboral side, cream-yellow on the oral side; the primary spines may be banded in red. Distribution: Recorded from the Philippines at 80 to 190 m depth (Fig. 3.158). Eurypatagus parvituberculatus (H.L. Clark, 1924) Main characteristics of the genus: maximum length 75 mm. This species differs from the other species of Eurypatagus by the small pore pairs in the petals. The aboral side in the posterior interambulacrum is inflated, forming a rounded, but distinct keel, so in profile the test is flat on the

3.3.2 Order Spatangoida L. Agassiz, 1840 

 205

upper side. In other species, the upper side is lowly arched. In young specimens, a subanal fasciole may be developed. Distribution: This species is recorded from South Africa and Mauritius, Indian Ocean. Bathymetric range 150 to 325 m (Fig. 3.159). Genus Linopneustes A. Agassiz, 1881 Linopneustes strongly resembles Elipneustes in general shape, but it differs in its shorter sternal plates, its narrower and straighter petals, and its deeper anterior notch. Test: Broadly oval, with shallow, but distinct anterior notch; in profile more or less high, upper side evenly arched; test fragile, especially the oral side; large forms. Apical system: Central, ethmolytic with four gonopores, situated on inner part of genital plates. Anterior ambulacrum: Adapically flush, gradually sunken toward the anterior margin, pore pairs small and positioned one above the other. Petals: Very long, open distally, parallel-sided, without occluded plates at the end. Oral side: Flat or slightly concave, peristome D-shaped, slightly sunken; periproct just inframarginal; posterior ambulacra naked. Labral plate: Wedge-shaped, extending to third ambulacral plate. Sternal plates: Short and triangular; fully tuberculated; posterior ambulacra on the oral side naked. Peripetalous fasciole: Thin, leading around the margin, posteriorly above the submarginal periproct. Subanal fasciole: Present in young, lost in adults, with two pore pairs to each side. Larger primary tubercles: Aborally in all interambulacra, mostly in short, oblique rows. Six species are included in this genus: Linopneustes brachypetalus Mortensen, 1950. East Australia, New Caledonia. Linopneustes excentricus de Meijere, 1903. Malayan ­Archipelago. Linopneustes fragilis (de Meijere, 1903). Philippines and Japan. Most probably to be synonymized with L. murrayi. Linopneustes longispinus (A. Agassiz, 1878). West Indies. Linopneustes murrayi (A. Agassiz, 1879). Malayan region, Philippines, Japan. Linopneustes spectabilis (de Meijere, 1903). Indo-West Pacific. Linopneustes brachypetalus Mortensen, 1950 Test: Highly domed, vertex centrally, anterior slope flattened, posterior slope arched; margin rather sharp; in

206 

A

 3.3 Superorder Atelostomata von Zittel, 1879

B

C

D

E

K

Fig. 3.158: Eurypatagus ovalis. (A–C) Test length 67 mm; Surprise Island, Pacific: (A) Aboral side. (B) Oral side. (C) Side view. (D, E) Test length with spines about 100 mm; Bohol, Philippines: (D) Aboral side: the primary spines usually reach half the test length. The spines below the periproct form a conspicuous tuft; they are broken here. The spines are very brittle. (E) Oral side.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

 207

B

C

D

E

F

Fig. 3.159: Eurypatagus parvituberculatus. (A, B) Test length with spines 85 mm; off Durham, South Africa. ZMUC: (A) Aboral side: the primary spines reach half the test length. Young specimen, length with spines 55 mm; Port Louis, Mauritius. (B) Oral side: the phyllodes are long and very conspicuous, especially those in the lateral ambulacra. (C) Test length 31 mm, off southern Natal, South Africa. Posterior end of test: this young specimen has kept a thin subanal fasciole. In larger specimens, it is usually lost. (D–F) Test length 31 mm, off southern Natal, South Africa: (D) Aboral side: the pore pairs in the ambulacra are rather inconspicuous. (E) Oral side: the naked area is broad, the sternum being almost without tuberculation. (F) Side view: the test is flattened.

208 

 3.3 Superorder Atelostomata von Zittel, 1879

outline broad with wide and fairly deep frontal notch; maximum length recorded 127 mm. Apical system: Slightly anterior. Paired petals: Long, anterior pair flexed to front, posterior pair with straight pore zones, pore pairs small. Oral side: Concave; labrum long, naked and overarching the peristome, sternal plates fully tuberculated; periproct below the margin. Peripetalous fasciole: Runs just around the margin; subanal fasciole absent (in adults?). Primary tubercles: A few primary tubercles forming an irregular arch on each aboral interambulacral plate; irregular, not very dense coat of small secondary and minute miliary tubercles. Distribution: East Australia from Queensland to Victoria, New Caledonia, depth range 270 to 640 m (Fig. 3.160). Linopneustes excentricus de Meijere, 1903 This species is characterized by having a thin, fragile test. There are some larger tubercles in the upper part of the anterolateral interambulacra and along the anterior ambulacrum. The peripetalous fasciole runs rather high around the test (as in L. murrayi). The primary spines are much longer and less numerous than in L. murrayi. Distribution: West Pacific; depth range 700 to 1,800 m. Linopneustes fragilis (de Meijere, 1903) Test: Very high; apex at slightly anterior apical system; in outline ovoid with very shallow anterior notch; length may reach 160 mm. Apical system: Subcentral. Paired petals: Long and parallel-sided; interporiferous zone broad and densely tuberculated. Oral side: Flat; peristome conspicuously sunken; labrum long and fairly well tuberculated; sternum densely tuberculated; periproct marginal; naked zones in posterior ambulacra not very broad. Fascioles: Peripetalous fasciole absent, subanal fasciole probably present only in young specimens. Primary tubercles: Small, densely covering the aboral surface; spines small and uniform. Distribution: West Pacific from Japan over the Philippines to the Malayan Archipelago; 560 to 1,170 m. Remarks: It is likely that L. fragilis and L. murrayi are conspecific (Fig. 3.161).

Linopneustes longispinus (A. Agassiz, 1878) Test: Fairly high, almost regularly arched from the sharp anterior margin to the very short, obliquely truncated posterior end; in outline oval with a moderately deep frontal sulcus; maximum length 142 mm. Paired petals: Relatively short and almost closed distally. Oral side: Slightly concave; labrum and especially the sternum distinctly keeled; peristome sunken in a groove, which leads toward the anterior sulcus. Periproct: At the short posterior end, visible from below. Peripetalous fasciole: Thin, running just along the margin, rising above the periproct; subanal fasciole only in young specimens. Primary tubercles: In one or two bows or irregularly distributed on all aboral interambulacra. Primary spines: Smooth, not thorny as in other species, terminating in a very sharp point; scarce miliary tubercles set with short, fine spinelets. Color: Upper side light brown with white primary spines, oral side darker brown. Distribution: West Indies from Cuba and the Gulf of Mexico to Barbados; at depths of 70 to 570 m. Linopneustes murrayi (A. Agassiz, 1879) Test: Hemispherical or subconical; in outline elliptical with a conspicuous frontal notch; maximum length 130 mm. Apical system: Subcentral. Anterior ambulacrum: Slightly depressed. Paired petals: Long, but not reaching the margin; parallelsided; pore pairs not conjugate; interporiferous zone well tuberculated. Oral side: Flat, sunken in front of the peristome; periproct round on the short, truncated posterior end. Peripetalous fasciole: Relatively high on the aboral surface, obliterated anteriorly; no subanal fasciole (in adults?). Aboral tuberculation: Heterogeneous, but not very dense, often the plate sutures may be naked. Primary spines: Ten to 15 mm long, sharply pointed, miliaries very short and fine. Color: The test may be brown to reddish-brown, the spines are cream-white, well cleaned. Distribution: West Pacific from the Malayan region over the Philippines to Japan at depths of 620 to 1,615 m.

▸ Fig. 3.160: (A–C) Linopneustes brachypetalus. Test length 127 mm, off New South Wales, Australia. ZMUC ECH 92: (A) Aboral side: there are only scattered primary tubercles, not arranged in rows. (B) Oral side: the fully tuberculated sternal plates are strongly bowed at the sides. (C) Side view. (D–F) Linopneustes fragilis. Test length 115 mm, Philippines. ZMUC: (D) Aboral side: there are no larger primary spines (or tubercles) developed. (E) Oral side. (F) Side view: the test is highly inflated to conical.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

B

C

D

E

F

 209

210 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

C

D

Fig. 3.161: Linopneustes longispinus. Test length 131 mm, Frederiksted, St. Croix, Lesser Antilles. ZMUC. (A) Side view. (B) Aboral side. (C) Oral side. (D) Labrum and sternum are tuberculated. Scale bar is 10 mm. After Mortensen (1950).

Remarks: Shigei (1986) synonymized L. murrayi with L. fragilis. Mortensen (1950) listed the following differences: L. murrayi has shorter petals than L. fragilis, it has longer and less numerous spines, and it has a peripetalous fasciole, although partly obliterated, which is obviously(?) totally lacking in L. fragilis. These are gradual differences. Furthermore, Mortensen could not find any tangible difference between the pedicellariae of the two species. Intermediate forms, like the specimen from Suruga Bay, demonstrate that the two “species” cannot be clearly separated (Fig. 3.162). Linopneustes spectabilis (de Meijere, 1903) Test: Lowly arched, highest point just in front of the apical system; short posterior end obliquely truncated; in outline

ovoid with shallow frontal notch; maximum length 180 mm or more. Apical system: Anterior; four gonopores. Anterior ambulacrum: Narrow and slightly depressed; small pore pairs. Paired petals: Long with conspicuous pore pairs, anterior pair may be flexed toward the anterior. Oral side: Faintly concave, midline of sternum keeled toward the posterior; peristome relatively shallowly sunken; labrum tuberculated in the anterior part, otherwise naked, very long, sternal plates fully tuberculated. Periproct: Transversely oval, rather small and inframarginal. Fascioles: Peripetalous fasciole thin, but distinct, running just above the margin, with several anterior branches;

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

 211

B

C

D

E

Fig. 3.162: (A, B) Linopneustes murrayi. Test length 93 mm, Sagami Bay, Japan. ZMUC: (A) Aboral side: the parallel-sided petals do not reach the margin. (B) Side view: the spines form heterogeneous clusters or bows on each interambulacral plate. (C–E) Specimens with spines: test length 116 mm, Suruga Bay, Central Japan, Pacific: (C) Aboral side: the spines are short and rather dense. (D) Aboral side, denuded test: the petals are long. (E) Side view.

212 

 3.3 Superorder Atelostomata von Zittel, 1879

bilobed subanal fasciole thin, in very large specimens absent. Large primary tubercles: Variable: in short rows, or in irregular double bows (in this latter case, the tubercles are conspicuously more numerous); miliary tubercles small and sparse. Primary spines: Long, robust and very sharply pointed. Color: The test and the spines are cream-yellow to light brown. Distribution: The species occurs from the Indian Ocean over the Malayan Archipelago to the Philippines at depths of 150 to 1,325 m (Fig. 3.163). Genus Paramaretia Mortensen, 1950 The genus is monospecific. This genus differs by its narrow petals, which are parallel-sided and, to a varying degree, obliterated in their anterior columns. The ambulacra do not indent the posterior interambulacrum to the rear of the episternal plates, as it is in the case of Maretia. Paramaretia peloria (H.L. Clark, 1916) (synonymous with Paramaretia multituberculata Mortensen, 1950) Test: Oval or ovoid in outline, anterior notch inconspicuous; in profile lowly arched aborally, oral side may be more or less concave, margin sharp; maximum length 125 mm. Apical system: Slightly anterior, ethmolytic, four gonopores. Anterior ambulacrum: Flush and narrow, pore pairs small, vertically positioned. Paired petals: Flush and narrow, pore pairs in anterior petals to a varying degree rudimentary or totally obliterated; pore pairs in posterior pair more developed, parallelsided, and distally open. Oral side: Peristome kidney-shaped, overarched by a lip, phyllode well-developed; labrum long, extending to third or fourth ambulacral plate; sternal plates narrow, naked except posteriorly. Periproct: Inframarginal on short, obliquely truncated posterior face. Fasciole: Subanal fasciole more or less developed; three to five pore pairs to each side; internal fasciole developed also to a varying degree. Primary tubercles: Numerous with sunken areoles in all interambulacra, small secondary and miliary tubercles sparse; oral primary tubercles in distinct rows with sunken areoles and spiral parapets. Primary spines: Coarse, slightly curved basally, length about one-third of test length. Color: The test is uniformly brownish, the spines are lighter.

Distribution: This species is only known along the southeast coast of Australia and off the south island of New Zealand; bathymetric range 165 to 657 m. Remarks: Baker & Rowe (1990) synonymized Paramaretia peloria and P. multituberculata. Based on numerous specimens, between 33 and 125 mm in length, they observed that the differences in the number of tubercles and spines, in the shape of the posterior end, and in the development of the subanal fasciole and internal fasciole are within the range of variation and are independent of size (Fig. 3.164). Genus Platybrissus Grube, 1865 The genus is monospecific. Platybrissus roemeri Grube, 1865 Test: Flat, aboral and oral side almost parallel; length may reach more than 70 mm. Apical system: Four gonopores, slightly anterior to center. Anterior ambulacrum: Narrow, flush, pore pairs very small and rudimentary. Paired petals: Flush and long, extending almost to ambitus; posterior pair longer than anterior pair. Oral side: Flat, peristome D-shaped and scarcely sunken, labrum elongated and wedge-shaped, not projecting the peristome; plastron and adjacent ambulacra naked except for posterior-most part, slightly raised; phyllodes welldeveloped. Periproct: At truncated posterior end. Subanal fasciole: Developed in young specimens, mostly lost in adults. Tuberculation: Aborally heterogeneous, but no distinct primary tubercles, coarse. Color: The spines are faintly banded in darker and lighter brown, the bare test is yellowish white. Distribution: This species is known in the Malayan Archipelago from the sublittoral zone into deeper water (Fig. 3.165).

3.3.2.7 Family Loveniidae Lambert, 1905 The family is characterized by having an internal fasciole. In Breynia, a peripetalous fasciole is additionally present. Five extant genera belong to this family: Araeolampas Serafy, 1974. Two species. Worldwide. Breynia Desor, in L. Agassiz & Desor, 1847. Five species. Indo West-Pacific. Echinocardium Gray, 1825. Ten species. Worldwide. Lovenia Desor, 1847. Eight species. Indo-Pacific. Pseudolovenia A. Agassiz & H.L. Clark, 1907. One species. Central Pacific. Literature: Clark & Rowe 1971; McNamara 1982; Rowe & Gates 1995; van Noordenburg 2008.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

B

 213

C

D

E

F

G

H

I

Fig. 3.163: Linopneustes spectabilis. (A) Young specimen of 55 mm length, Philippines. Photo courtesy of H. van Noordenburg. (B–D) Adult specimen, test length 145 mm, Maldives, Indian Ocean. ZMUC: (B) Aboral side: the primary tubercles are less numerous than in the specimen from the Philippines. (C) Oral side. (D) Side view. (E–G) Specimen with spines, test length 148 mm, Philippines. (E) Aboral side. (F) Oral side. The spines are thin and numerous. (G) Side view. (H, I) Posterior end of L. spectabilis: the bilobed subanal fasciole is thin, but still visible (size of test 114 mm). Scale bar is 10 mm: (I) Anterior edge: the peripetalous fasciole forms at least two branches. Scale bar is 10 mm.

214 

A

 3.3 Superorder Atelostomata von Zittel, 1879

B

C

D

E

F

G

Fig. 3.164: (A, B) Paramaretia peloria. Test length 115 mm, off Victoria, Southeast Australia. ZMUC: (A) Aboral side. (B) Oral side. (C) Part of left anterior petal: the pore pairs of the anterior column are, to a varying degree, rudimentary or totally obliterated. Scale bar is 10 mm; after Mortensen (1951). (D, F) Specimen with spines, length 97 mm, off Victoria, Southeast Australia. ZMUC: (D) Aboral side: the primary spines reach a length of about one-third of the test length. (F) Side view. (E, G) Specimen without spines; test length 104 mm; off Victoria, Southeast Australia. ZMUC ECH 41: (E) Aboral side. (G) Side view.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

B

D

 215

C

E

F

Fig. 3.165: Platybrissus roemeri. (A, B, D) Test length 63 mm; Negros, Philippines: (A) Aboral side: the paired petals reach almost to the margin, the posterior ones are longer. (B) Oral side: the plastron and the adjacent ambulacra are naked except toward the posterior end. (D) Side view: the test is very flat; on the oral side, only the posterior end of the plastron is slightly raised. (C, E, F) Specimen with spines. Test length with spines 74 mm; Negros, Philippines: (C) Aboral side: the spines extend beyond the posterior end of the test. (E) Oral side: the broad plastron is naked except for the posterior-most end. (F) Side view: the spines are long and slender but, on the whole, the spination looks coarse.

Genus Araeolampas Serafy, 1974 All specimens collected and examined thus far have been fragmentary. Araeolampas is very close to Homolampas, a maretiid according to The Echinoid Directory (Smith 2005). Test: Very fragile, mostly large forms, up to 110 mm in length; distinct anterior groove. Apical system: Ethmolytic, four well-developed genital pores.

Ambulacra: Not sunken to form petalodia, pore pairs double or single, situated obliquely near the inner edge of plates. Oral side: Flat, peristome anterior, labrum strongly elongated toward the posterior, reaching to second or third pair of adjoining ambulacra; only posterior part of sternum set with primary tubercles. Fascioles: Peripetalous and subanal fasciole present.

216 

 3.3 Superorder Atelostomata von Zittel, 1879

Primary tubercles: Perforate, crenulated; present on interambulacra 1 to 5; areoles shallowly sunken with slight internal bulges. Primary spines: Long, curved, and serrated on one side of the shaft. The genus consists of five species: Araeolampas atlantica Serafy, 1974. Southeast of USA. Araeolampas fulva (A. Agassiz, 1879). Central and East Pacific. Araeolampas glauca (Wood-Mason & Alcock, 1891). Bay of Bengal. Araeolampas hastata (A. Agassiz, 1898). East Pacific. Araeolampas rostrata (de Meijere, 1903). Arafura Sea.

Remark: According to H.L. Clark (1917), the sharply cut posterior furrow is developed in the holotype, but absent in the cotype.

Araeolampas atlantica Serafy, 1974 Test: Large and flattened, length of holotype 92 mm, height approximately 35 mm, in outline heart-shaped with wide anterior sulcus. Apical system: Anterior with four gonopores. Ambulacral pores: Near the apical system single. Oral side: Unknown; posterior side vertically truncated, slightly indented above the periproct. Periproct: Large, not visible from above. Fasciole: Peripetalous and subanal fasciole present. Primary tubercles: Perforate and crenulated; present on all interambulacra but restricted to the area within the peripetalous fasciole, only in interambulacrum 5 do they occur just outside of the fasciole; areoles of primary tubercles sunken, posterior part more deeply than anterior part; internally slight bulges. Differentiation: The species is distinguished from A. fulva mainly by its tridentate pedicellariae: in A. atlantica, they are distinctly wider than in A. fulva, which occurs in the Pacific (Fig. 3.166).

Araeolampas hastata (A. Agassiz, 1898) Main characteristics of the genus: small form. Test: On the whole low, posterior end vertically truncated. Apical system: With three gonopores. The drawing of Westergren, showing four gonopores, is incorrect according to Mortensen (1950). Fasciole: Distinct peripetalous fasciole; obviously shieldshaped subanal fasciole. Primary tubercles: Large also in posterior interambula­ crum 5; secondary and miliary tuberculation very dense aborally, very few large primary tubercles; areoles of primary tubercles sunken; internally, they form small rudimentary processes, but no distinct ampullae as in H. fulva. Distribution: Cocos and Malpelo islands, East Pacific; depth 1,785 to 2,100 m (Fig. 3.167).

Araeolampas fulva (A. Agassiz, 1879) Main characteristics of the genus: length may reach more than 100 mm. Test: Elongated heart-shaped with broad and deep anterior sulcus; highest point in front of the apical disk, falling steeply toward the anterior margin and sloping slowly toward the shortly truncated posterior end. Apical system: Anterior, with four gonopores. Anterior ambulacrum: Deeply sunken. Peristome: Also deeply sunken. Fascioles: Peripetalous fasciole very thin, disappearing anteriorly; broad shield-shaped subanal fasciole. Primary tubercles: Enlarged in all aboral interambulacra; in the anterior interambulacra in somewhat vertical series; distinct internal ampullae. Distribution: Central and eastern Pacific; bathymetric range 3,665 to 4,500 m.

Araeolampas glauca (Wood-Mason & Alcock, 1891) Main characteristics of the genus; maximum length 93 mm. Large form similar to Araeolampas fulva, but differing in its shallower anterior groove and broader test, although the general shape may vary considerably according to Koehler (1926). Four gonopores, thin peripetalous fasciole, shape of subanal fasciole unknown. Occurrence: Bay of Bengal; 2,985 m

Araeolampas rostrata (de Meijere, 1903) Test: Aborally slightly domed, posteriorly sloping to the low posterior end, which is encircled by the subanal fasciole; in outline oval, posteriorly narrowing to a somewhat pointed end; length of the two known specimens 18 mm. Apical system: Anterior, with four gonopores and a single madrepore. Ambulacra: Nonpetaloid. Oral side: Flat, not sunken in front of the peristome, posterior part of plastron raised; labrum not protruding, sternum obviously tuberculated only in the posterior part. Periproct: Supramarginal, visible from above. Fascioles: Peripetalous fasciole thin, leading anteriorly just along the margin, rising in a bow on both sides toward the aboral side; subanal fasciole broader than peripetal fasciole, positioned almost vertically. Primary tubercles: Large along the anterior ambulacrum and a few within the peripetalous fasciole on the lateral interambulacra; secondary and miliary tubercles rather scarce. Primary spines: Strong and long, more than half the test length.

3.3.2 Order Spatangoida L. Agassiz, 1840 

A

C

 217

B

D

E F

G

H

G

Fig. 3.166: (A, B) Araeolampas atlantica: (A) Holotype, the oral side is lacking, off Virginia, USA; 2,575 m; Albatross Station 2105; length 92 mm. (B) USNM E37549, from off South Carolina at 32°09′43″ N, 76°42′57″ W to 32°15′40″ N, 76°39′53″ W, 2,100 m; length 71 mm. Photos courtesy of D.L. Pawson. (C–F) Araeolampas fulva. Test length 95 mm; east of Christmas Island, Central Pacific. From A. Agassiz (1881): (C) Aboral side. (D) Oral side. (E). Side view. (F) Subanal fasciole and periproct. (G, H) Araeolampas glauca. Length 82 mm. Bay of Bengal, India: (G) Aboral side. (H) Side view. The oral side is lost. Sketches after photos in Koehler (1926).

218 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

D

E

G

C F

H

I

J

Fig. 3.167: (A–E) Araeolampas hastata. Test length 32 mm; Cocos Island, East Pacific. From A. Agassiz (1904): (A) Aboral side. (B) Side view. (C) Periproct. (D) Side view of spine. (E) Serrated primary spine from the underside. (F) Milled ring and base of spine. (G–I) Araeolampas rostrata: (G) Test length 18 mm, Arafura Sea, Indonesia. NHM Amsterdam. Aboral side: the whole specimen was very fragile, being partly fragmentary. Other views were unavailable. (H–J) Drawings after de Meijere (1903): (G) Aboral side. (H) Oral side. (I) Side view.

Occurrence: Arafura Sea, Indonesia, at depths of 538 and 694 m. Remark: De Meijere (1903) has only made rough sketches of the two specimens in natural size. As can be seen, the outline drawings show the posterior rostrum rather exaggerated. Genus Breynia Desor, in L. Agassiz & Desor, 1847 Test: Oval with a shallow frontal notch, more or less tapering to the posterior rounded end. Apical system: Small, ethmolytic with four gonopores (except in Lovenia triforis). Anterior ambulacrum: Slightly depressed, pore pairs set with specialized tube feet. Paired petals: Flush, pore pairs reduced within internal fasciole, just outside pore zones widest, distally narrowing and closed. Oral side: Flat, inflated, and keeled toward the adoral point of the subanal fasciole; peristome kidney-shaped, wider than long; labral plate narrow and long, paired sternal plates narrow and triangular, more or less tuberculated. Periproct: Inframarginal on truncated posterior face.

Fascioles: Shield-shaped subanal fasciole and inner fasciole, peripetalous fasciole not indented laterally. Primary tubercles: Aborally conspicuous with deeply sunken areoles, absent in posterior interambulacrum; larger tubercles bordering anterior ambulacrum; oral primaries in distinct series. Primary spines: Bent at the base and lying over the test; secondary spines curved. The genus consists of five species, living in the IndoWest Pacific: Breynia australasiae (Leach, 1815). East coast of Australia. Breynia desorii Gray, 1851. South to northwest coast of Australia. Breynia elegans Mortensen, 1948. Indonesia to East China Sea. Breynia neanika McNamara, 1982. Northern Australia. Breynia vredenburgi Andersen, 1907. Andaman Island. Breynia australasiae (Leach, 1815) Test: Thick; ovate with slight anterior notch, upper side convex, oral side keeled toward posterior; length up to 120 mm.



Apical system: Subcentral, four gonopores, madreporite strongly expanding to the posterior. Anterior ambulacrum: Slightly depressed, pore pairs and tube feet specialized, tubercles along the frontal groove enlarged and arranged in rows. Paired petals: Petaloid and flush, wide at the internal fasciole, closed distally. Oral side: Peristome small and kidney-shaped, labrum long and narrow; sternum triangular and fully tuberculated. Periproct: Large, on short, undercut, truncated posterior end. Fascioles: Internal fasciole broad, crossing the anterior ambulacrum halfway down to the margin; peripetalous fasciole surrounding distal ends of petals and parallel to margin; subanal fasciole shield-shaped, number of enclosed pore pairs generally five or six. Primary tubercles: Distinct within the peripetalous fasciole, with deeply depressed areoles, up to 15 in each of the anterior pair, up to 25 in each of the posterior pair, and none in interambulacrum 5. Distribution: B. australasiae is restricted to the east coast of Australia from Cape York to Fraser Island and around Lord Howe Island, living from the intertidal to a depth of 10 m in sandy sediments (Fig. 3.168). The two common species in Australia differ in the following features (Tab. 3.9): There is, however, a certain degree of variation, and smaller specimens with less numerous tubercles are difficult to distinguish. Breynia desorii Gray, 1851 This species is much like B. australasiae, but differs in the following features: Maximum length 110 mm. Within the peripetalous fasciole, many more primary tubercles are developed: up to 42 in each of the anterior pairs, up to 45 in each of the posterior pairs. The peripetalous fasciole passes nearer the ambitus around the anterior side of test. The internal fasciole is narrower and reaches further down the anterior ambulacrum. The subanal fasciole generally has six to seven pore pairs, but varies between three and eight. Distribution: B. desorii is restricted to the southern and western coasts of Australia from Esperance to Darwin. It is found in sandy sediments from the intertidal zone to a depth of 140 m. Breynia elegans Mortensen, 1948 Main characteristics of the genus: large form; maximum length reaches 126 mm.

3.3.2 Order Spatangoida L. Agassiz, 1840 

 219

Test: Distinctly narrowing toward the posterior, forming a more or less distinct rostrum; in profile, low with an overhanging, obliquely truncated end. Labral plate: Narrow and very long; consequently, the sternal plates are rather short, tuberculated only in the posterior part (not as in B. australasiae and B. desorii). Fascioles: Internal fasciole rather short, apical system nearly in the middle of it; subanal fasciole with up to eight pore pairs per column. Primary spines: Aborally about half the test length, bent at the base; secondary spines short, fine, and curved. Color: Primary spines banded in cream-yellow with purple or red-brown. Distribution: This species is known in the entire IndoMalayan region, in the Philippines, and the China Sea; bathymetric range 60 to 200 m (Fig. 3.169). Breynia neanika McNamara, 1982 This species differs from B. elegans in its more oval test, the aboral side flattened. There are fewer primary tubercles aborally and the apical system is more centrally placed; the petals are broader and shorter, the internal fasciole longer, the apical system positioned posteriorly within the internal fasciole. The sternum is fully tuberculated. Length up to 92 mm. Distribution: Restricted to Northern Australia, off Cobourg Peninsula and at the east coast of Cape Yorke Peninsula, Queensland: depth 10 to 82 m. Breynia vredenburgi Anderson, 1907 Only known specimen is 30 mm long. Aboral side: Primary tubercles scarce and confined to the anterolateral interambulacra, whereas there are none at all in the posterior. Occurrence: Port Blair, Andaman Island, India. Depth not recorded, but probably in shallow water. Genus Echinocardium Gray, 1825 The genus is characterized by a short labral plate, wider than long and just contacting the second ambulacral plate. The shield-shaped subanal fasciole is positioned at the posterior side, not orally. The internal fasciole is very distinct. Lovenia and Breynia are usually shallowly buried in the substrate. When disturbed, they move swiftly over the ground, raising their long aboral spines in defense. However, Echinocardium is adapted to burrowing into the sediment and staying there. Depending on the substrate, Echinocardium cordatum, the most specialized species, lives as much as 20 cm beneath the surface, whereas Echinocardium flavescens burrows only just below the sea floor.

220 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

C

D

E F

Fig. 3.168: (A–D) Breynia australasiae. Test length 63 mm, height 33 mm; Bowen, Queensland, Australia. ZMH: (A) Aboral side: most of the long primary spines within the peripetalous fasciole are broken. (B) Oral side: the spines are longer than those on the aboral side. The sternum is elongated triangular and fully covered with rather small, curved spines. Test length 97 mm. (C) Aboral side: there are fewer numerous primary tubercles than in Breynia desorii scattered within the peripetalous fasciole. (D) Labrum and adjacent ambulacral and sternal plates: the areoles of the large tubercles on the sternum are eccentric. Scale bar is 3 mm. After Mortensen (1951). (E, F) B. desorii. Test length 107 and 48 mm; Dirk Hartog Island, Western Australia: (E) Aboral side: the smaller specimen has not yet developed the full number of primary tubercles. (F) Oral side: the plastron is elongated triangular and tuberculated. The peristome is relatively small.

Tab. 3.9: A comparison of the two common Breynia species found in Australia: B. australasiae

B. desorii

Up to 15 primary tubercles in each anterior interambulacrum

Up to 42 primary tubercles in each anterior interambulacrum

Up to 25 primary tubercles in each posterior interambulacrum

Up to 45 primary tubercles in each posterior interambulacrum

Five to six subanal pore pairs

Six to eight subanal pore pairs

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

D

B

 221

C

E

F

G

H

Fig. 3.169: (A, B) Breynia elegans. Test length 126 mm, Philippines. Photo courtesy of H. van Noordenburg: (A) Aboral side: the primary spines are banded. (B) Oral side: the sternum is naked adorally. (C) Labrum and adjacent ambulacral plates: the labral plate is just in contact with the two sternal plates. Scale bar is 8 mm; after Mortensen (1951). (D, E) Specimen without spines. Test length 115 mm; Mactan Island, Philippines: (D) Aboral side: in this large specimen, the narrow peripetalous fasciole is partly obscured. (E) Side view: the test is very low. On the oral side, there is a prominent keel toward the posterior. (F) Ethmolytic apical system: the madrepores extend far beyond the posterior ocular plates. Scale bar is 3 mm, after Mortensen (1951). (G, H) Breynia neanika. Holotype, test length 71 mm, Albany Passage, Queensland, Australia. Sketches after the photos in McNamara (1982): (G) Aboral side. (H) Side view.

222 

 3.3 Superorder Atelostomata von Zittel, 1879

Biology: E. cordatum is found deeply buried in sandy to even muddy sediments, and is adapted to this extreme habitat with specialized tube feet and spines. It builds vertical funnels to the sea floor for obtaining fresh water. This funnel is plastered with mucus, distributed by the long penicillate tube feet in the anterior ambulacrum and compacted into the walls by the spines of the apical tuft. The cilia, tiny hairs that cover the fasciolar spines, create a continuous water current that comes down the funnel and flows over the test, providing the respiratory podia with oxygen. The cilia of the subanal and anal fascioles drive the current to the posterior and, added with the waste, into the sanitary funnel. This horizontal tube is built by the specialized podia and spines within the subanal fasciole. In clean sand in the littoral zone, the animal moves through the sediment at a maximum speed of 6 to 8 cm/h. As it advances by means of the plastronal spines, the echinoid builds new funnels with the aboral podia—every 4 cm in sand and at greater distances in mud. E. cordatum has developed two feeding strategies: 1. The penicillate podia around the peristome selectively collects particles from the surrounding sediment manipulating them into the mouth. 2. The long tube feet in the anterior ambulacrum on the upper side, extending to a length of 20 cm, examines the sea floor around the funnel opening and brings down organic-rich grains. These particles drop into the lower anterior ambulacrum, where a mucous string moves downward to the peristome. The adjacent spines are arched over this groove to shield the string from the substrate in front of the sea urchin. This behavior is very similar to Moira atropos, which is also specialized for living in muddy sediments (Fig. 3.170). Nine Recent species of Echinocardium are listed, most of them in shallow water: Echinocardium capense Mortensen, 1907. South Africa, the Mediterranean, and the West Pacific Echinocardium cordatum (Pennant, 1777). East Atlantic and the Mediterranean; the virtually identical subspecies E. cordatum australe Gray 1851 in Japan, New Zealand, and southern Australia. For the occurrence of Echinocardium cordatum in Brazil, there are inconsistent records. Echinocardium fenauxi Pequignat, 1964. Western Mediterranean and Japan. Echinocardium flavescens (O.F. Müller, 1776). Northeast Atlantic and the Mediterranean. Echinocardium laevigaster A. Agassiz, 1869. Caribbean and the Northwest Atlantic. Echinocardium lymani Lambert, 1924. Central Japan.

Echinocardium mediterraneum (Forbes, 1844). Mediterranean and Gulf of Biscay. Echinocardium meteorense Mironov, 2006. On seamounts in Northeast Atlantic. Echinocardium mortenseni Thiéry, 1909. Western Mediterranean, Banjuls Sea. Echinocardium pennatifidum Norman, 1868. From Southern Norway to England. It is often not easy to distinguish the species because intermediates and hybrids occur. The genus is most diverse in the Mediterranean, where five species are known. These populations became isolated from populations in the Atlantic and Pacific after the closure of the Mediterranean Sea in the early Miocene. Intermediates between E. cordatum and E. fenauxi are commonly found especially in the western Mediterranean along the coast of Spain (Tab. 3.10). Literature: Mortensen 1913; Nichols 1960; Baker 1969; Higgins 1974, 1975; Shigei 1986; David & Laurin 1996; Mironov 2006. Echinocardium capense Mortensen, 1907 E. capense was described from South Africa. David & Laurin (1996) have recently synonymized the species with E. mortenseni Thiéry, 1909 from the Mediterranean Sea and E. lymani Lambert & Thiéry, 1924 from Japan. Mironov (2006) admits that the morphological features are equal, but he has detected the pedicellariae and he thinks the species different. Main characteristics like genus: it has the following differences from E. cordatum: Test: Oval; length reaches up to 50 mm. Apical system: Slightly sunken and saddle-like. Anterior ambulacrum: Flush on the upper side, frontal notch faint. Paired petals: Columns are parallel, not widened at the internal fasciole. Fascioles: Internal fasciole is short; anal fasciole is joined with the subanal fasciole. Distribution: South Africa, bathymetrical range extends from 12 m to a depth of 400 m. Echinocardium cordatum (Pennant, 1777) This species is recorded all over the world, except the Americas. The regions are discontinuous as it lives in temperate waters: from Northern Norway along the European coasts to the Mediterranean and Morocco, in South Africa, in Japan, New Zealand, and southern Australia. It is the species with the widest distribution of all echinoids. Test: Thin, moderately high, highest point in the posterior interambulacrum; deep anterior notch; length 70 mm, rarely up to 90 mm.

3.3.2 Order Spatangoida L. Agassiz, 1840 

A

B

 223

C

D

E outer part of ambulacrum

internal fasciole inner part of ambulacrum

Fig. 3.170: (A, B) Echinocardium cordatum burrowing in the sediment: in sandy sediment (A), the sea urchin builds a new funnel at a distance of approximately 4 cm from the abandoned one. In muddy sediment (B), the animal, which is buried less deeply, maintains the funnel to the surface and the sanitary funnel for a longer distance. It is likely that the building of a new tube needs more effort than in sandy substrates. Direction of locomotion is to the left. After Nichols (1960). (C) Peristome with surrounding phyllodes: the pores, associated with penicillate tube feet, are enlarged. Closest to the peristomial edge in each ambulacrum, one plate has two pores, each with one phyllodal tube foot. Scale bar is 5 mm (Echinocardium mediterraneum). (D) Loveniid: Aboral side near the apical system: there is s sharp break between the large pore pairs of the outer part of the petals and the rudimentary or microscopic pores within the internal fasciole (Lovenia elongata). (E) Echinocardium capense. Aboral plate pattern. Scale bar is 10 mm. After David & Laurin (1996).

Tab. 3.10: Characteristics of some of the Echinocardium species: E. capense

E. cordatum

E. fenauxi

E. flavescens

E. mediterraneum

E. pennatifidum

Anterior notch lacking

Anterior notch distinct

Anterior notch distinct

Anterior notch very faint or lacking

Anterior notch shallow

Anterior notch very faint

Ambulacrum III flush on upper side

Ambulacrum III sunken on upper side

Ambulacrum III sunken on upper side

Ambulacrum III flush on upper side

Ambulacrum III slightly raised on upper side

Ambulacrum III flush on upper side

Irregular double series of pore pairs in ambulacrum III

Regular, dense, single series of pore pairs in ambulacrum III

Single series of widely spaced pore pairs in ambulacrum III

Single series of widely spaced pore pairs in ambulacrum III

Single series of widely spaced pore pairs in ambulacrum III

224 

 3.3 Superorder Atelostomata von Zittel, 1879

Apical system: Posterior, four gonopores, broader than long. Anterior ambulacrum: Broad, distinctly sunken on the upper side; approximately 50 plates within the fasciole; two broad series of irregularly arranged, large, single pores, each associated with a specialized tube foot. Paired petals: Slightly sunken, wide at the internal fasciole, tapering to the distal end, the lateral columns forming a continuous arc. Oral side: Convex with a large peristome slightly anterior, labrum prominent; plastron slightly keeled toward a sharp posterior rear. Periproct: Very variable in shape, from elongated to transverse; opening on the more or less truncated posterior side. Fascioles: Internal fasciole long; subanal fasciole shieldshaped with up to four pore pairs, sharply pointed at its lower end; anal fasciole expanding along the sides of the periproct onto the upper side and separated from the subanal fasciole. Spination: Uniform over the upper side, except for larger spines bordering the anterior ambulacrum; longer on the oral side, and spatulate on the plastron. Color: In life, the sea urchin is light brown, the bare test is whitish. Distribution: This species is found worldwide, but the regions are discontinuous: from Northern Norway along the European coasts into the Mediterranean, and to Morocco; in South Africa, in Japan, New Zealand, and ­southern ­Australia. It lives from the intertidal zone to a depth of 500 m (Fig. 3.171).

Remarks: Many intermediates between E. cordatum and E. fenauxi are recorded (Fig. 3.172).

Echinocardium fenauxi Pequignat, 1963 This species greatly resembles E. cordatum. It differs by the following features: Test: Large and depressed, distinct frontal notch and sunken anterior ambulacrum; length of more than 80 mm. Apical system: Posterior. Anterior ambulacrum: Two distinct series of large pores, each set with one specialized podium. The periproct being wider than long. Distribution: This species is known from the Mediterranean Sea in southern France, the Gulf of Genova, and the Adriatic Sea in the sublittoral. This form may also be found in Japan.

Echinocardium laevigaster A. Agassiz, 1869 Nearly related to E. flavescens. Test: Broad anteriorly and pointed posteriorly; length 25 mm. Fasciole: Four pairs of ambulacral plates within the subanal fasciole; internal fasciole. Distribution: Known from the Caribbean and the Northwest Atlantic at depths of 130 to 250 m.

Echinocardium flavescens (O.F. Müller, 1776) This small species differs by the following features from other species: Test: Fragile, high, frontal notch faint or absent; maximum length 35 mm. Anterior ambulacrum: Flush on the upper side; two series of widely spaced pores, tube feet not specialized. Paired petals: Also almost flush. Oral side: Labrum elongated, reaches to second ambulacrum; periproct opens on the vertical truncated posterior end. Fascioles: Internal fasciole broad, subanal fasciole broadly joined with the anal branches. Primary tubercles: Large within the apical part of interambulacrum V, set with long spines. Distribution: This species is known from Northern Norway and Iceland to the Mediterranean in the littoral and to a depth of 325 m. Echinocardium keiense Mortensen, 1950 Test: Oval, slightly wide anteriorly, frontal end almost vertical; length 35 mm. Anterior ambulacrum: Not sunken, pores in regular single series. No primary spines aborally, except along frontal ambulacrum within the inner fasciole, where they form a conspicuous raised tuft. Occurrence: Kei Islands, Indonesia; 250 m deep.

Echinocardium lymani Lambert & Thiéry, 1924 Main characteristics like genus: length reaches 28 mm. Test: Round without a frontal notch; small form; high posteriorly; flush anteriorly. Anterior petal: Very narrow with fine pores.

▸ Fig. 3.171: (A–E) Echinocardium cordatum. Length of test 49 mm; Brittany, France: (A) Aboral side: in the sunken anterior ambulacrum, two irregular series of pores are developed. (B) Oral side: the plastron is fully tuberculated. (C) Side view: the highest point lies in the posterior interambulacrum. (D) Aboral plate pattern. The ambulacral plates outside the internal fasciole are large but decrease sharply in size within the fasciolar area. Interambulacra lightly shadowed, internal fasciole dark. Scale bar is 10 mm. After David & Laurin (1996). (E) Posterior side: the subanal fasciole is not connected with the anal branches and extends far onto the upper side. Scale bar is 10 mm. (F–H) Echinocardium fenauxi. Test length 82 mm; Adriatic Sea, Mediterranean: (F) Aboral side. (G) Side view: the posterior interambulacrum is distinctly raised. (H) Posterior side: the anal branches are broadly connected with the subanal fasciole. Scale bar is 10 mm.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

C

F

B

D

E

G

H

 225

226 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

D

F

C

G

E

Fig. 3.172: (A–E) Echinocardium flavescens. Test length 32 mm; North Sea: (A) Aboral side: the general spination is uniform, the longer primary spines are mostly broken. (B) Oral side: the sternum is set with flattened, spatulate spines. (C, D) Specimen without spines; length 33 mm; North Sea: (C) Aboral side: the larger primary tubercles are distributed over all aboral interambulacra. (D) Side view: Anterior and posterior sides are steep. (E) Posterior side; scale bar is 10 mm. (F, G) Echinocardium lymani (as E. dubium). From H.L. Clark (1917): (F) Aboral side. (G) Posterior side.

Paired petals: Narrow and long; subequal in length; the distal ends do not join. Oral side: Labrum tuberculated, sternal plates only in the posterior part tuberculated. Fascioles: Internal fasciole broad, subanal fasciole with two or three pores to each side. No primary tubercles. Distribution: Sagami Bay, Suruga Bay and off Honshu Island, Central Japan; depth 195 to 350 m. Echinocardium mediterraneum (Forbes, 1844) This species differs markedly from E. cordatum as follows: Test: Maximum length 50 mm. Apical system: Rather anterior. Anterior ambulacrum: Slightly raised on the aboral side and sunken along the vertical front, the pores are not crowded, and the associated tube feet are not specialized. Paired petals: Lateral columns of pore pairs do not form a continuous arc.

Fascioles: Internal fasciole sharply pointed posteriorly and short; subanal fasciole barely connected with the anal branches, enclosing two pore pairs. Periproct: Elongated, depressed at its lower end and visible from above; the posterior end is slightly oblique. Distribution: This species is known only in the Mediterranean Sea and northward to the Bay of Biscay in depths between 5 and 280 m (Fig. 3.173). Echinocardium meteorensis Mironov, 2006 Main characteristics like genus: Test: Round, up to 24 mm. Anterior ambulacrum: Not sunken, pores in regular series. Paired petals: Parallel rows. Oral side: Labrum reaching the middle of the second adjacent ambulacra.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

C

 227

B

D

E

F G

Fig. 3.173: (A–C) Echinocardium mediterraneum. Test length 33 mm; Alanya, Turkey: (A) Aboral side: at the front there is a faint notch, which ends abruptly at the edge to the upper side. The area within the internal fasciole, the enlarged interambulacral plates, and the narrow ambulacrum are slightly raised. (B) Oral side: the peristome is large and kidney-shaped. (C) Side view: the test is high, the posterior side obliquely truncated. (D, E) Specimen with spines. Test length 39 mm; Algarve, Portugal: (D) Aboral side: the spines are smoothly flattened against the test, except for the apical and subanal tufts. (E) Oral side: the lateral, curved spines are specialized for burrowing; the spines of the plastron are used for locomotion and are also curved. (F) Aboral plate pattern: the interambulacral plates within the broad internal fasciole are enlarged. The plates of the petals are strongly reduced in size toward the apical system with a sharp break at the fasciole. Scale bar is 10 mm. After David & Laurin (1996). (G) Posterior side: the subanal fasciole is just connected to the anal fasciole. Scale bar is 10 mm.

Fasciole: Internal fasciole very small and thin; one to two large pores to each side of subanal fasciole; anal branch well-developed. Primary tubercles: Very few. Distribution: Seamounts in the Northeastern Atlantic, off Portugal; at depths between 180 and 450 m.

Echinocardium mortenseni Thiérry, 1909 Nearly related to E. flavescens, it differs by the following features. Oral side: Labrum elongated to second adjacent ambulacra. Primary tubercles in the anterior interambulacra. Distribution: West Mediterranean; depth 14 to 40 m.

228 

 3.3 Superorder Atelostomata von Zittel, 1879

Echinocardium pennatifidum Norman, 1868 It differs from E. cordatum: Test: Length reaches 70 mm. Anterior ambulacrum: Frontal notch being very faint. Anterior side: Being gently curved. Fascioles: Anal branches not joined to the subanal fasciole. Primary tubercles: No larger spines at the aboral side in all interambulacra. Distribution: Echinocardium pennatifidum is known from a rather restricted region, from Bergen, South Norway, the Faeroes Islands, and around Great Britain to the Channel Islands. The bathymetrical range is from the littoral zone to a depth of 275 m. It is very similar to E. flavescens, with which hybrids are commonly developed (Fig. 3.174). Genus Lovenia Desor, 1847 This genus is characterized by a depressed, delicate test. On the aboral side, there are large primary tubercles with deeply depressed areoles, which form worm-shaped ampullae on the inner surface of the test. The large lateral tubercles on the oral side, set with locomotory spines, have asymmetrical, deeply sunken areoles, which are like spiral parapets on the inner side of the test. These additional structures are necessary to strengthen the thin skeleton where the muscles of the spines attach. Test: Elongated, posteriorly narrower than anteriorly, shallow frontal sulcus, in profile flattened above and below. Apical system: Ethmolytic, four or three gonopores. Anterior ambulacrum: More or less sunken, wide, pore pairs set with specialized podia. Paired petals: Flush, pore pairs more or less conjugate, reduced within internal fasciole, outside pore zones widest, narrowing and distally closed; outer columns often continuous and forming an arc laterally. Oral side: Flat, peristome kidney-shaped; a few enlarged podia in the phyllodes; broad naked zone consisting of the posterior ambulacra, the anterior part of the sternum, and a wide area around the peristome; labral plate long and narrow, sternal plates narrow and triangular, tuberculated only on the posterior part. Fascioles: Subanal fasciole bilobed, internal fasciole present. Periproct: Submarginal on the short truncated posterior side; in some species, deeply invaginated. Primary tubercles: Scattered aborally with deeply sunken areoles, internal ampullae; oral lateral primaries very conspicuous, arranged in distinct series. Primary spines: Very long, serrated on the lateral side. Biology: The species of Lovenia live shallowly buried in sandy to gravelly sediment in sheltered waters from the ebb zone to the sublittoral. When an animal feels disturbed, it comes out of the substrate and moves swiftly

over the surface on its long oral spines. The primary spines of the aboral side are raised in defense to deter predators such as snails. In the intertidal zone, Lovenia also emerges from the sediment, when the water drains from the sand at low tide. Sometimes it “runs” over the sloping beach to the water, but it is also able to stay exposed for several hours in shallow pools till the tide returns and washes over the sea urchin again. Then, it quickly buries itself once again in the sand. The long aboral spines are also used for righting, when the animal becomes overturned. It takes only 10 s to re-right into the normal orientation. Ten species are included in the genus: Lovenia camarota H.L. Clark, 1917. North Australia. Lovenia cordiformis A. Agassiz, 1872. East Pacific from California to Ecuador. Lovenia doederleini Mortensen, 1950. Malayan region. Lovenia elongata (Gray, 1845). East Africa to Japan, the Malayan Archipelago, and Australia. Lovenia gregalis Alcock, 1893. Malayan region, Philippines, and Japan. Lovenia grisea A. Agassiz & H.L. Clark, 1907. One specimen known from Hawaii Islands. Differs from L. hawaiiensis by the lack of the deep anal invagination. Lovenia hawaiiensis Mortensen, 1950. Hawaii Islands. Lovenia lata Shigei, 1981. Sagami Bay, Central Japan. Lovenia subcarinata Gray, 1851. East Africa to Java Sea and Hong Kong. Lovenia triforis Koehler, 1914. Bay of Bengal, Indonesia to Philippines and Japan. The three Lovenia species, L. elongata, L. cordiformis, and L. hawaiiensis, are closely related and very similar, especially in sharing the deep anal funnel. They differ slightly in the shape of the tridentate pedicellariae and mainly in the separated geographical range (Fig. 3.175). Lovenia camarota H.L. Clark, 1917 Main characteristics of the genus: nearly related to L. elongata. Synonymized by Clark & Rowe (1971). It has the following ­differences from L. elongata: Test: The posterior interambulacrum raised to a low keel; holotype 53 mm. Fascioles: Four to five pores to each side of the subanal fasciole. Distribution: North Australia, Holothuria Bank to Torres Strait; depth 18 to 60 m (Fig. 3.176). Lovenia cordiformis A. Agassiz, 1872 Main characteristics of the genus: length of test may reach 83 mm. This species differs from the very similar L. elongata by the usually higher test and by the deeper conjugate pore pairs in the paired petals. Eight to 11 pore pairs are

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

B

 229

C

D

E

F

Fig. 3.174: (A) Echinocardium meteorense. Labrum and adjacent plates. Scale bar is 3 mm. After Mironov (2006). (B) Echinocardium mortenseni. Aboral side. Scale bar is 10 mm. After David & Laurin (1996). (C, D) Echinocardium pennatifidum: aboral plate pattern: the plates in the paired petals show a less marked break in size compared to E. mediterraneum, the columns are more parallel. The band of the internal fasciole is narrow. Scale bar is 10 mm. After David & Laurin (1996). (D) Posterior side. The anal fasciole is not connected to the subanal fasciole. Scale bar is 10 mm. (E, F) Echinocardium pennatifidum. ZMUC: (E) Aboral side; test length 51 mm; North Sea; depth 150 m. ZMUC. (F) Side view of specimen with spines; test length 53 mm; Channel Islands. ZMUC. There is a distinct anal tuft of spines, but no apical tuft is developed.

developed in the subanal fasciole and the area within the peripetalous fasciole may be more or less raised. Both have a sunken invaginated periproct, and the anterior part of the internal fasciole is double, it does not cross the ambulacrum. However, the geographical range is quite different. Distribution: The species is known from Southern ­California, USA to Ecuador and the Galapagos Islands; living from the ebb zone to a depth of 100 m. Lovenia doederleini Mortensen, 1950 Main characteristics of the genus: maximum length 36 mm. Test: Elongated, posterior part slightly narrowing, truncated at the end. Paired petals: Of about the same length, anterior column of anterior pair short.

Oral side: Labral plate with moderate lip, reaching posteriorly to the second adjoining ambulacral plate. Fascioles: Internal fasciole often double anteriorly, but usually not distinct on the anterior ambulacrum; posterior band of subanal fasciole widened. Periproct: At the truncated posterior end, not depressed. Primary tubercles: Few adorally; primary spines banded. Distribution: Known throughout the entire Malayan region; living in shallow water to a depth of 35 mm (Fig. 3.177). Lovenia elongata (Gray, 1845) Test: Delicate, elongated with distinct anterior notch, depressed in profile, flattened above and below; length may reach 85 mm.

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 3.3 Superorder Atelostomata von Zittel, 1879

A

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Fig. 3.175: (A) Lovenia elongata. Lembeh Strait, Sulawesi, Indonesia. Photo by Roland Kraft. (B–E) Lovenia. Anterior branch of the internal fasciole: (B) L. elongata (likewise in Lovenia cordiformis and L. hawaiiensis): the anterior branch is double, but does not cross the ambulacrum. In Lovenia doederleini, the double branches cross the ambulacrum, but are often indistinct. Scale bar is 4 mm. (C) Lovenia subcarinata: the single anterior branch crosses the ambulacrum. Scale bar is 4 mm. After Mortensen (1951). (D, E) L. elongata: internal structures of the aboral side: (D) View to the inner surface of the test. (E) Outer surface of the test: the muscles of the long primary spines are fixed in deeply depressed areoles around the tubercles, which form worm-shaped ampullae on the inner surface. The test itself is too thin to support these spines when the animal raises them in defense.

Apical system: Subcentral with four gonopores. Anterior ambulacrum: Slightly sunken, pore pairs differentiated; tube feet specialized. Paired petals: Flush, wide at the internal fasciole, tapering to the distal ends.

Oral side: Peristome kidney-shaped; labrum narrow and elongate, sternal plates tuberculated only over the posterior part. Fascioles: Internal fasciole present; subanal fasciole bilobed and leading into the anal funnel; enclosing 6 to 10 distinct pore pairs.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

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Fig. 3.176: (A–C) Lovenia camarota. Holotype, length 53 mm, from Clark (1917); (A) Aboral side. (B) Oral side. (C) Side view. The anterior is to the right. (D–G) Lovenia cordiformis. Test length 80 mm, Taboguilla, Pacific Panama. ZMUC: (D) Aboral side. (E) Oral side. The long spines on the lateral oral side serve for swift locomotion. (F) Side view: the long primary spines lie along the test when the animal is burrowed, but in case of danger, the sea urchin emerges out of the sediment and runs on its oral spines over the surface very quickly raising its sharply pointed spines against an eventual predator. (H) Young specimen with spines; test length 28 mm; Manta, Ecuador. ZMH. (G) Aboral side.

Periproct: On the upper side of a deep invagination on the posterior side. Primary tubercles: In the paired interambulacra on the aboral side and larger tubercles laterally on the oral side with deeply sunken areoles. Spination: Aboral primary spines longer than the test; long, lateral spines on the oral surface for locomotion over the sea floor. Color: The living animal is yellowish or reddish, the bare test is whitish. Distribution: The species is recorded from East Africa from the Red Sea to Mozambique, and from southern Japan to

the east coast of Australia, Sydney, New South Wales. It is not known from the South Pacific Islands. Its bathymetrical range reaches from the ebb zone down to a depth of 94 m. Literature: Rowe & Gates 1995; Miskelly 2002 (Fig. 3.178). Lovenia gregalis Alcock, 1893 Test: Elongate with a deep and narrow anterior sulcus and a relatively broad posterior end; in profile low, anterior side rounded, posterior side vertically truncated; aboral posterior interambulacrum raised forming a distinct keel; the length may reach up to 96 mm. Apical system: Sunken, positioned slightly anterior.

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 3.3 Superorder Atelostomata von Zittel, 1879

Anterior ambulacrum: Increasingly depressed toward the deep frontal notch. Paired petals: Anteriorly with closed pore zones, the posterior petals are open, pore pairs subconjugate; outer columns bending inward, thus not forming an arc laterally. Oral side: Flat, peristome sunken, labral plate straight, not forming a lip. Periproct: On vertical posterior side, pointed above and below; not invaginated. Fascioles: Internal fasciole disappearing anteriorly, bilobed subanal fasciole very conspicuous. Primary tubercles: Large on the lower part of the paired interambulacra. Color: The primary spines are not banded. Distribution: The species is found from the Bay of Bengal over the Philippines to Japan; bathymetric range 275 to 930 m (Fig. 3.179).

A

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Lovenia hawaiiensis Mortensen, 1950 Main characteristics of the genus: maximum length 39 mm. This species strongly resembles L. elongata in having a deep anal invagination. The pore pairs are likewise sunken, but not really conjugate, and the internal fasciole is also double, bending toward the frontal ambulacrum. Distribution: The species is endemic to the Hawaiian Islands. Lovenia lata Shigei, 1981 Test: In size, almost as wide as long; low; outline rounded; holotype in length 67.7 mm. Apical system: Slightly anterior. Anterior ambulacrum: Flush, indistinctly notched. Paired petals: Pore zones sunken and rudimentary within the internal fasciole; widely apart, the anterior

D

C

Fig. 3.177: Lovenia doederleini. (A–C) Test length 35 mm, Kei Islands, Indonesia. ZMUC ECH 125: (A) Aboral side. (B) Oral side. (C) Profile. (D) Specimen with spines. Test length 39 mm, Kei Islands, Indonesia. ZMUC. Aboral side: the primary spines are banded in beige and brown.

▸ Fig. 3.178: Lovenia elongata. (A, B) Test length 69 mm; Philippines: (A) Aboral side: the conspicuous primary tubercles are scattered over the paired interambulacra. (B) Oral side: the lateral tubercles are large, leaving a broad naked area in the anterior part of the plastron and the adjacent ambulacra and also in front of the peristome. (C) Side view. The test is flattened. The stippled line indicates the anal invagination, into which the periproct opens (arrow). (D, E) Specimen with spines; test length 46 mm; Gulf of Thailand: (D) Aboral side: the long primary spines are flattened to the test when the animal burrows in the sediment. (E) Oral side: the area around the peristome and the anterior part of the sternal plates is naked. The long lateral spines are used for locomotion. Two tufts of spines are set on larger tubercles at the sides of the bilobed subanal fasciole.

3.3.2 Order Spatangoida L. Agassiz, 1840 



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3.3.2 Order Spatangoida L. Agassiz, 1840 



much shorter; the posterior ones straight, narrowing rather abruptly; all pores strongly conjugate. Oral side: Labrum keeled, reaching the anterior angle of the third ambulacral plates, sternal plates not broad. Fascioles: Internal fasciole broad, subanal fasciole with four pores to each side. Primary tubercles: Five to six to each anterolateral interambulacrum, 14 to 15 to each posterolateral one; with worm-shaped ampullae on the aboral side. Distribution: Restricted to the Sagami Bay, Central Japan; depth 80 to 120 m. Lovenia subcarinata Gray, 1851 Test: Thin, elongated, anterior and posterior part tapering, truncated posterior end; length up to 74 mm. Apical system: Posterior, four gonopores. Anterior ambulacrum: Sunken, bordered by irregularly arranged tubercles set with long spines that do not form an adapical tuft; well-developed pore pairs and specialized

A A

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 235

tube feet; the adjacent interambulacral plates form a more or less distinct keel. Paired petals: Flush, posterior pair shorter than anterior pair, wide at the internal fasciole. Oral side: Flattened, most of the surface naked; very long and narrow labral plate, sternal plates also narrow, only posterior part tuberculated; adjacent ambulacra broad. Periproct: Submarginal, not sunken. Fascioles: Internal fasciole long and narrow; subanal fasciole bilobed. Primary spines: Few set on large tubercles on the upper side, arranged in distinct, but well separated, horizontal rows; areoles deeply sunken. Distribution: L. subcarinata is known from East Africa to the Gulf of Siam, the Java Sea, and to Hong Kong, China. It is not known east of the Timor Sea. It lives at depths of 10 to 35 m. Biology: This species lives in more muddy bottoms than the other species of Lovenia (Fig. 3.180).

C

D

Fig. 3.180: (A) Lovenia lata. Test length 66 mm, Sagami Bay, Japan. Sketches after the photos in Shigei (1986). (B–D) Lovenia subcarinata. Test length 67 mm; Hong Kong. ZMH: (B) Aboral side: the apical system is positioned posteriorly and, consequently, the posterior petals are shorter than the anterior ones. (C) Oral side: not only the posterior ambulacra but also the anterior part of the plastron and broad areas at the sides and in front of the peristome are naked. (D) Side view.

◂ Fig. 3.179: (A–D) Lovenia gregalis. Test length 63 mm, Philippines. Photos courtesy of H. van Noordenburg; (A) Aboral side. (B) Oral side. (C) Side view. (D) Specimen with spines; test length 104 mm, Buton Strait, Sulawesi, Indonesia. ZMUC. Side view. (E, F) Lovenia hawaiiensis. Young specimen, test length 18 mm, Honolulu, Hawaii. ZMUC Holotype ECH 167: (E) Aboral side. (F) Oral side.

236 

 3.3 Superorder Atelostomata von Zittel, 1879

Lovenia triforis Koehler, 1914 Test: Heart-shaped and low vaulted, posterior interambulacrum keeled, sloping regularly toward the truncated, overhanging posterior side; edges sharp; length may reach 55 mm. Apical system: Three gonopores, only a few hydropores. Anterior ambulacrum: Sunken toward the more or less deep V-shaped frontal notch. Paired petals: Short and narrow, pore pairs subconjugate. Fascioles: Internal fasciole disappearing near the anterior ambulacrum, subanal fasciole with up to four pore pairs per column. Peristome: Kidney-shaped, area below the periproct depressed, but not funnel-like. Primary spines: Faintly banded. Distribution: Known from the Bay of Bengal and the Malayan region over the Philippines to Japan; bathymetric range 80 to 250 m (Fig. 3.181). Genus Pseudolovenia A. Agassiz & H.L. Clark, 1907 Pseudolovenia hirsuta A. Agassiz & H.L. Clark, 1907 Test: Low, anterior part well domed, posterior part lower; conspicuous frontal notch; posterior end subvertically truncated; maximum length 60 mm. Apical system: Anterior, four gonopores. Paired petals: Pore pairs in the posterior ambulacra gradually continue without any sharp break into the ambital pore pairs beyond the actual petal. Oral side: Sternal plates tuberculated only posteriorly, length of labrum variable. Fascioles: Internal fasciole only 20 mm long with rounded and broad posterior end; subanal fasciole wide. Primary tubercles: Several primary tubercles with sunken areoles in the posterior column of the anterolateral and in the anterior column of the posterolateral interambulacrum. Distribution: This species is endemic to the Hawaii Islands at depths of 350 to 1,265 m (Fig. 3.182).

3.3.2.8 Family Maretiidae Lambert, 1905 The Maretiidae differs from the spatangids by its narrow, elongated, wedge-shaped labral plate and the relatively short and, on the whole, triangular sternum. They differ from the Loveniidae by the lack of an internal fasciole and by having a shield-shaped subanal fasciole (mostly) rather than a bilobed one. In some maretiid genera, an

additional peripetalous fasciole is developed. The apical system is ethmolytic with three or four gonopores. The family Maretiidae includes seven genera: Granobrissoides Lambert, 1920. Southern Australia; two species. Gymnopatagus Döderlein, 1901. Indo-West Pacific; two species. Hemimaretia Mortensen, 1950. East Africa; one species. Homolampas A. Agassiz, 1874. Caribbean and Malayan Archipelago; two species. Maretia Gray, 1855. Indo-West Pacific; four species. Nacospatangus A. Agassiz, 1873 (syn. with Pseudomaretia Koehler, 1914 and Gonimaretia H.L. Clark, 1917). IndoPacific; seven species. Pycnolampas A. Agassiz & H.L. Clark, 1907. Hawaii Island; one species. Spatagobrissus H.L. Clark, 1923. South Africa and South Australia; two species. Literature: Serafy 1974; David & De Ridder 1989; Baker & Rowe 1990; Rowe & Gates 1995. Genus Granobrissoides Lambert, 1920 Two extant species are described: Granobrissoides dyscritus H.L. Clark, 1938 Granobrissoides hirsutus Mortensen, 1950 Granobrissoides dyscritus H.L. Clark, 1938 The holotype of G. dyscritus, at least 85 mm in length, lacks the entire oral and posterior side. The shape of the plastron and the subanal fasciole are not known. There are no large primary tubercles anywhere. Found probably in Victoria, Australia. Baker & Rowe (1990) described two specimens from Southern Australia as cf. G. dyscritus, with no primary tubercles within the peripetalous fasciole, but in length only 25 mm. They show a subcircular test with an arched aboral side, a high, truncated posterior side, a rather short labrum, and an inflated posterior plastron. Found in Western and South Australia (Fig. 3.183). Granobrissoides hirsutus Mortensen, 1950 Test: Oval, anterior side convex, aboral side regularly arched with rounded margin, fairly large forms. Apical system: Ethmolytic with four gonopores. Anterior ambulacrum: Narrow, flush and set with inconspicuous pore pairs. Paired petals: Lanceolate, pore pairs in anterior column of anterior pair reduced.

▸ Fig. 3.181: Lovenia triforis. (A–C) Test length 46 mm; Mactan Island, Philippines: (A) Aboral side. (B) Oral side. (C) Side view. (D–F) Specimen with spines; test length 45 mm, Cebu, Philippines: (D) Aboral side. (E) Oral side. (F) Side view.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

B

C

D

E

F

 237

238 

 3.3 Superorder Atelostomata von Zittel, 1879

A

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Fig. 3.182: Pseudolovenia hirsuta. Length 60 mm, Hawaii Islands. From H.L. Clark (1917). (A) Aboral side. (B) Side view. (C) Oral side.

A

B

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E

D

F

Fig. 3.183: Granobrissoides hirsutus. (A) Ethmolytic apical system and anterior ambulacrum with small pore pairs. Scale bar is 4 mm. (B) Labrum and adjacent ambulacral plates. Scale bar is 6 mm. Both after Mortensen (1951). (C–F) Test length 75 mm, South of Lake Entrance, Victoria, Australia. ZMUC ECH 182 (as Lissospatangus hirsutus). (C) Aboral side: there are no primary tubercles and spines on the aboral interambulacra. (D) Oral side: the tubercles are surrounded by large areoles and set with strong, curved spines. (E) Left side: the upper side is covered by fine, uniform-sized spines. (F) Right side: the distinct peripetalous fasciole encircles the petals closely. There are no indentations in the interambulacra and it continues across ambulacrum III.

Oral side: Flattened, naked areas relatively narrow; peristome kidney-shaped. Periproct: At the undercut posterior side. Fascioles: Peripetalous fasciole well-developed, subanal fasciole shield-shaped. Primary tubercles: None on the aboral side; spines fine and uniform; oral primary tubercles with conspicuous areoles, spines slender, robust, and curved. Distribution: The species is only known from Victoria and Tasmania, Australia; at 25 to 40 m.

Genus Gymnopatagus Döderlein, 1901 Test: Very fragile, especially the oral side; outline elongated heart-shaped with conspicuous frontal sulcus; in profile very low and with sharp margin. Apical system: Anterior, ethmolytic, four gonopores positioned on the inner part of genital plates. Anterior ambulacrum: Depressed increasingly toward the margin; pore pairs small, uniserially arranged and with simple podia. Paired petals: Subequal and bowed, narrowing distally.



Oral side: Flat, peristome D-shaped; periproct on short, truncated posterior side; labrum long and narrow, straight cut. Fascioles: Peripetalous fasciole without lateral indentations; shield-shaped subanal fasciole with two pore pairs to each side. Tubercles: Heterogenous within the peripetalous fasciole; sharp contrast between the long spines within the fasciole and the short, fine, densely distributed small ones on the rest of the aboral surface; tubercles crenulated and not sunken. The genus includes two species: Gymnopatagus magnus A. Agassiz & H.L. Clark, 1907. Indo-West Pacific. Gymnopatagus valdiviae Döderlein, 1901. East coast of Africa. The two species differ in the distribution of the primary tubercles and spines within the peripetalous fasciole. In G. magnus, both the lateral interambulacra and the posterior interambulacrum are set with primary tubercles; in G. valdiviae, the primary tuberculation is restricted to the lateral interambulacra. Gymnopatagus magnus A. Agassiz & H.L. Clark, 1907 Main characteristics of the genus: Test: Low in profile, with the highest point at the apical system, gently sloping to the anterior margin and less steeply to the low, shortly truncated posterior end; maximum length recorded 111 mm. Anterior petals: Anterior pore zones curved like a shallow S; pore pairs barely conjugate; pore zones distally almost closed. Oral side: A low, but distinct posterior keel; labrum long and narrow, sternum fully tuberculated. Fasciole: Subanal fasciole shield-shaped, with two pore pairs on each side. Tubercles: Orally rather uniform; naked areas in posterior ambulacra broad, continue into the anterior groove. Distribution: The species is known from the Indian Ocean to Japan, at depths of 780 to 2,350 m (Fig. 3.184). Gymnopatagus valdiviae Döderlein, 1901 Test: In outline heart-shaped with deep anterior sulcus; upper side lowly arched, margin sharp, posterior end short and obliquely truncated; length up to 77 mm. Apical system: Slightly anterior with four gonopores. Anterior ambulacrum: Sunken toward the margin, depression does not continue onto the oral side. Paired petals: Bowed, length about half the test length, pore zones distally closed, pore pairs distinctly conjugate. Oral side: Flat, posterior end keeled, sternum fully tuberculated; naked area on posterior ambulacra very broad,

3.3.2 Order Spatangoida L. Agassiz, 1840 

 239

leaving a laterally narrow, oblique area covered by larger tubercles set with straight, slender spines. Fascioles: Peripetalous fasciole thin, no lateral indentation, indistinct on the anterior ambulacrum; subanal fasciole broad. Primary tubercles: Aborally up to eight enlarged primary tubercles in anterolateral interambulacra, up to fifteen in the posterolateral pair, no enlarged primary tubercles in interambulacrum 5. Distribution: Coast of Somalia, East Africa; depth between 741 and 1,362 m (Fig. 3.185). Genus Hemimaretia Mortensen, 1950 The genus is monospecific. Hemimaretia elevata Döderlein, 1906 The only known specimen has a length of 30.5 mm, it is now totally broken. Test: Very fragile, in outline oval, anterior notch shallow, but distinct; in profile moderately low, aboral side only slightly arched, posterior interambulacrum keeled; anterior side steeply sloping, posterior end obliquely truncated. Apical system: Conspicuously anterior, ethmolytic, three gonopores. Anterior ambulacrum: Aborally depressed, narrow with small, simple pores. Paired petals: Rather short, flush with the test, pore pairs small, nonconjugate; lacking proximally. Oral side: Flat, phyllodes very short: only the two ambulacral plates nearest the peristome are set with penicillate tube feet; peristome D-shaped; periproct on truncated posterior side, transversely oval. Labrum long and narrow, sternum also tuberculated only on the posterior part. Fasciole: Subanal fasciole shield-shaped with two pore pairs to each side; no peripetalous fasciole. Primary tubercles: Large, on the aboral side fairly numerous, distributed in lateral interambulacra, absent in the posterior one; sunken and internally associated with the conspicuous ampullae (as in some species of Lovenia); tuberculation otherwise fine and dense; oral large tubercles also with sunken areoles and parapets. Primary spines: Curved, thorny. Occurrence: Off Kenya, East Africa, at a depth of 693 m. Genus Homolampas A. Agassiz, 1874 Test: Very fragile; in outline oval with a more or less distinct frontal groove; small to large forms. Apical system: Ethmolytic, three or four gonopores. Anterior ambulacrum: Narrow and flush, pores rudimentary, podia simple. Paired ambulacra: Nonpetaloid and flush; pore pairs small, toward ambitus smaller and more distant.

240 

 3.3 Superorder Atelostomata von Zittel, 1879

A

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3.3.2 Order Spatangoida L. Agassiz, 1840 

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Fig. 3.185: (A–D) Gymnopatagus valdiviae. (A, B) Test length 76 mm, Somalia, East Africa. Aboral side: there are several longer primary spines aborally. (B) Oral side. Test length 59 mm, Somalia, East Africa. From Döderlein (1906). The naked areas are very broad. (C) Test without spines, length 40 mm, Somalia, East Africa. From Döderlein (1906). (D) Aboral side. East Africa. Courtesy of B. Ruthensteiner, ZS München. A few greatly enlarged primary tubercles are developed in the lateral interambulacra, but not in the posterior interambulacrum 5. Scale bar is 40 mm. (E–H) Hemimaretia elevata. Test length 30.5 mm, Kenya, East Africa. From Döderlein (1906). (E) Aboral side: the fairly numerous, large primary spines form a dense coat. (F) Oral side. (G) Side view. (H) Ethmolytic apical system with three gonopores. Scale bar is 1 mm. After Mortensen (1951).

Oral side: Flat, anteriorly sunken toward the peristome; periproct on the short and truncated posterior end; labrum narrow and wedge-shaped; sternal plates triangular, tuberculated only on the posterior part.

Fascioles: Well-developed subanal fasciole mostly bilobed, peripetalous fasciole more or less distinct or even absent. Primary tubercles: Few large, noncrenulate primary tubercles aborally with more or less distinct sunken

◂ Fig. 3.184: Gymnopatagus magnus. (A–C) Test length 111 mm, Philippines. ZMUC. (A) Aboral side: the anterior row of pore zones in the anterior petals is curved like a shallow S. (B) Oral side: the naked areas in the posterior ambulacra are broad, the sternal plates being fully tuberculated. (C) Side view: the posterior end is low. (D) Specimen with spines, test length 105 mm, Philippines, ZMUC. Aboral side: numerous long primary spines are distributed within the peripetalous fasciole. Outside the fasciole, the spines are fine and uniform. (E) Apical system: the gonopores are clustered in the center. Scale bar is 2 mm. (F) Labrum and adjacent ambulacral plates. Scale bar is 6 mm. (G) Distal part of anterior petal. Scale bar is 6 mm, After Mortensen (1951).

242 

 3.3 Superorder Atelostomata von Zittel, 1879

ampullae, general tuberculation fine and uniform; large, lateral tubercles on oral side with more or less sunken areoles and parapets (as in Lovenia). Primary spines: Serrated. Two species are described: Homolampas fragilis (A. Agassiz, 1869). Caribbean. Homolampas lovenioides Mortensen, 1948. Sulawesi, Indonesia. Homolampas fragilis (A. Agassiz, 1869) Main characteristics of the genus: maximum length may reach 21 mm. Test: Upper side slightly keeled, frontal groove broad and shallow; three gonopores. Fascioles: Subanal fasciole twice as broad as long, distinctly bilobed; no peripetalous fasciole. Primary tubercles: Large aborally, confined to the posterior column of the anterior interambulacra and the anterior column of the posterior interambulacra, no tubercles in interambulacrum 5. Primary spines: Aborally about half the test length; one side serrated. Color: In life, grayish with purple tinge or pale brown, the test often mottled with darker spots. Distribution: This species is known from the Caribbean Sea and the Florida Strait. Bathymetric range 550 to 3,550 m (Fig. 3.186). Homolampas lovenioides Mortensen, 1948 Main characteristics of the genus: Test: Narrowing toward the posterior end; frontal notch broad and shallow; aboral side raised, rounded keel in interambulacrum, sloping gently toward vertical posterior side. Apical system: With three or four gonopores. Oral side: Flat, peristome deeply sunken. Fascioles: Subanal fasciole bilobed, three times as wide as long, no peripetalous fasciole. Occurrence: Gulf of Boni, Sulawesi, Indonesia, at a depth of 987 m. Genus Maretia Gray, 1855 The genus is characterized by a low test. Apical system: Ethmolytic, four gonopores; anterior ambulacrum with simple pores, paired petals lanceolate and flush, anterior series of anterior petal with rudimentary pore pairs adapically. Oral side: Peristome wider than long; periproct on shortly truncated posterior side; labral plate long and narrow, sternal plates narrow and triangular, tuberculated only posteriorly.

Fasciole: Subanal fasciole shield-shaped. Tubercles: Heterogeneous aborally, larger primary tubercles sunken; oral lateral tubercles in series, with parapet and areoles slightly sunken. Primary spines: Long and serrate. Maretia differs from Eupatagus in having only a subanal fasciole, whereas the latter also has a peripetalous fasciole. Lovenia is distinguished from Maretia in having an internal fasciole in addition to the subanal one. It is distinguished from Nacospatangus by the number of gonopores; Maretia has four whereas Nacospatangus has three. Literature: Bolau 1873; Rowe & Gates 1995; Miskelly 2002. Four species are included in the genus: Maretia carinata Bolau, 1873. South Sea. Maretia cordiformis Mortensen, 1948. Malayan Archipelago. Maretia planulata (Lamarck, 1816). Indo-West Pacific. Maretia tuberculata A. Agassiz & H.L. Clark, 1907. Japan. Only a single immature specimen from the Korea Strait is known. Neither Nisiyama (1968) nor Shigei (1986) mention this species as occurring in Japan (Fig. 3.187). Maretia carinata Bolau, 1873 This species differs from M. planulata according to the following characteristics: The upper side is relatively high and arched, the posterior end rounded. The petals are shorter, and the pore pairs are less numerous, especially in the anterior row of the anterior petals. There are only eight pore pairs against thirteen in M. planulata at the same size. The plastron is broader, and the tuberculated part of the sternum larger. Distribution: M. carinata is recorded from the South Pacific Islands (Fig. 3.188). Maretia cordiformis Mortensen, 1948 Test: Fragile; aboral side lowly arched, oral side flattened; in outline broadly heart-shaped with a distinct broad anterior notch, as wide as long, short posterior end vertically truncated; length may reach 40 mm. Anterior ambulacrum: Flush adapically, depressed toward the margin. Paired petals: Long, relatively broad, bowed, pore pairs large and conjugate. Oral side: On the whole flat, anterior ambulacrum sunken, posterior end slightly elevated; naked area very broad with almost parallel sides; peristome subpentagonal, periproct at the truncated posterior face; phyllodes in anterior pair of ambulacra long, short in posterior pair; labral plate narrow and elongated, just contacting sternal plates; sternum set with small tubercles only on the very posterior part. Fasciole: Subanal fasciole shield-shaped and slightly reniform, broader than long, one or two pore pairs to each side; enclosing a single cluster of tubercle.

3.3.2 Order Spatangoida L. Agassiz, 1840 



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 243

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F

G

Fig. 3.186: (A–E) Homolampas fragilis. (A–C) Test length 21 mm, West Indies. ZMUC. (A) Aboral side. (B) Oral side: there is a deep depression in front of the peristome. The sternum is tuberculated only on the posterior part, the naked area being broad. (C) Side view: the posterior part of the plastron is strongly elevated toward the rear. (D) Peristomial region. (E) Part of aboral side: there are a few large primary tubercles with deeply sunken ampullae. Scale bar is 4 mm. After Mortensen (1950). (F, G) Homolampas lovenioides. (F) Labrum and adjacent ambulacral plates. Scale bar is 5 mm. (G) Serrate spines from the sternum (left) and from the frontal notch (right). Scale bar is 2 mm. After Mortensen (1950).

244 

A

 3.3 Superorder Atelostomata von Zittel, 1879

B

C

D

E

Fig. 3.187: (A–C) Species of Maretia: labral plate and adjacent ambulacral plates. After Mortensen (1951). (A) Maretia carinata: the labrum is relatively broad. Scale bar is 4 mm. (B) Maretia cordiformis: the labrum is straight cut at the peristomial edge. Scale bar is 4 mm. (C) Maretia planulata: the labrum is very elongated and arched at the peristomial edge. Scale bar is 5 mm. (D, E) M. carinata. Test length 30 mm; Viti Levu, Fiji Island. ZMH. (D) Aboral side: the petals are shorter than in M. planulata. (E) Side view: the test is gently vaulted on the upper side.

Tuberculation: Aborally heterogeneous, large primary tubercles in all interambulacra. Primary spines: Long, very slender, curved and serrated on one side; primaries of the subanal tuft straight and as long as the test length. Color: Test yellowish-white to pinkish-red and light brown, spines sometimes faintly banded. Distribution: The species is recorded from the entire Malayan region and the Philippines. Bathymetric range 50 to 150 m. Maretia planulata (Lamarck, 1816) Test: Thin and lightweight; oval in outline with very faint frontal notch and tapering to the truncated rear; upper side flattened, posterior interambulacrum keeled; maximum length 70 mm. Apical system: Anterior; four gonopores; madreporite strongly expanding to the posterior. Anterior ambulacrum: Narrow and flush with the test, pore pairs small, tube feet simple. Paired petals: Long and lanceolate, almost flush; pore pairs in anterior row of anterior petals rudimentary.

Oral side: Flattened, with well-developed phyllodes around the kidney-shaped peristome; posterior interambulacrum and adjacent ambulacra raised to form a sharp border to the lateral interambulacra, tuberculation restricted to the posterior end of the plastron and around the peristome; periproct on the undercut posterior end, pointed above and below. Subanal fasciole: Shield-shaped. Spination: Aborally heterogeneous with coarse tubercles scattered over the paired interambulacra set with long spines; areoles of primary tubercles sunken; long lateral spines on oral side arranged in rows. Color: The long spines are whitish, the short aboral spines in a symmetrical pattern of whitish and light to dark brown, which remain visible on the dry test. Bare tests are white. Biology: The species lives buried in sand or silt but can move swiftly over the sea bottom. Distribution: M. planulata is known in the Indo-Pacific from East Africa to Fiji and Gilbert Island, and from Sagami Bay, Japan to Sydney, East Australia. It lives from the ebb zone to a depth of 60 m (Fig. 3.189).

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

B

 245

C

D

E

F

Fig. 3.188: Maretia cordiformis. (A–C) Test length 28 mm, width 28 mm, Flores Sea, Indonesia. ZMUC. (A) Aboral side: all interambulacra are set with scattered coarse primary tubercles. The primary spines are rather slender. (B) Oral side: the naked area is very broad. (C) Side view: the upper side is slightly arched, the posterior end being vertically truncated. (D) Specimen with spines, length of test 33 mm; Palawan, Philippines. (D) Aboral side: the spines of the subanal tuft may be as long as the test length. (E, F) Young specimen of 13 mm length. Negros, Philippines. (E) Aboral side: the test is wider than long. (F) Oral side.

Maretia tuberculata A. Agassiz & H.L. Clark, 1907 Only a single immature specimen from the Korea Strait is known. Neither Nisiyama (1968) nor Shigei (1986) mention this species as occurring in Japan. Genus Nacospatangus A. Agassiz, 1873 It differs from Maretia by the number of gonopores; Maretia has four whereas Nacospatangus has three. Test: In outline oval, no or only shallow anterior notch; aboral side lowly arched, posterior interambulacrum keeled. Apical system: Anterior, ethmolytic, three gonopores. Anterior ambulacrum: Flush or only slightly sunken, narrow with simple pores. Paired petals: Flush, pore zones closed distally; anterior series in anterior pair reduced adapically; pore pairs conjugate. Oral side: Flat; plastron keeled toward the posterior; peristome D-shaped; periproct at the steep posterior end with or without deep invagination; labral plate narrow, sternal

plates also rather narrow; posterior part tuberculated, naked areas in posterior ambulacra narrow and almost parallel. Fasciole: Subanal fasciole shield-shaped. Tuberculation: Uniform and fine aborally; larger tubercles bordering the anterior ambulacrum; in some species, a few large primary tubercles in interambulacra 1 and 4 near the ambitus; on the oral side larger tubercles. Remarks: Mortensen (1951) suggested that the genera Pseudomaretia Koehler, 1914 and Gonimaretia H.L. Clark, 1917 should be synonymized with Nacospatangus. Thus, this genus consists of seven species: Nacospatangus altus (A. Agassiz, 1864). Indo-West Pacific. Nacospatangus depressus H.L. Clark, 1917. South California, possibly Gonimaretia laevis. Nacospatangus gracilis A. Agassiz, 1873. Juan Fernandez Island, off the coast of Chile. Nacospatangus interruptus (Studer, 1880). Northwest ­Australia.

246 

A

 3.3 Superorder Atelostomata von Zittel, 1879

B

C

D

E

Fig. 3.189: Maretia planulata. (A, B) Test length with spines 45 mm; Siquijor, Philippines. (A) Oral side: the plastron and the adjacent ambulacra are nearly naked, only the posterior part of the sternal plates and the labral plate are tuberculated and bear spines. (B) Side view: the aboral primary spines are laid back over the test. An anal tuft is developed within the subanal fasciole. (C–E) Specimen without spines; test length 41 mm; Siquijor, Philippines. (C) Aboral side: the primary tubercles are scattered over the paired interambulacra. (D) Oral side: the raised plastron is sharply bounded by the series of large lateral tubercles. (E) Side view: the test is depressed and the rear of the plastron is inflated.

Nacospatangus laevis (H.L. Clark, 1917). Southern California, USA, Baja California. Nacospatangus oblongus (Mortensen, 1950). Galapagos Islands. Nacospatangus tylotus (H.L. Clark, 1917). Malayan region and Philippines (Fig. 3.190). Nacospatangus altus A. Agassiz, 1864 Main characteristics of the genus: test length may reach 54 mm. Test: Moderately high, aboral side gently arched from the rather sharp anterior margin to the truncated posterior side; test oval in outline with a shallow anterior sulcus, narrowing toward the posterior. Paired petals: Pore pairs rudimentary in the adapical half of length. Labrum: Very narrow, elevated at the peristome. Periproct: In a deeply sunken invagination on the posterior face. Subanal fasciole: Shield-shaped with three or four pore pairs to each side. Primary tubercles: Up to five or six in the posterolateral interambulacra near the ambitus with depressed areoles.

Color: Test in living specimens deep purple, preserved specimens are gray-white with a more or less distinctly purplish tinge. Distribution: The species is recorded in the Indo-West Pacific from Mauritius to central Japan and to the Banda Sea, Indonesia; from the littoral to a depth of more than 200 m. Nacospatangus depressus H.L. Clark, 1917 Possibly synonymous with G. laevis after A.B. Smith. The holotype is 11.5 mm long; the only paratype is even smaller. Occurrence: Off south California. Nacospatangus gracilis A. Agassiz, 1873 The holotype is an immature specimen of 17 mm length. Occurrence: Southeast Pacific, Juan Fernandez Islands. Nacospatangus interruptus (Studer, 1880) Main characteristics of the genus: this species resembles N. altus in having a deep anal invagination. N. interruptus differs by the broader and shorter labrum. Furthermore, there are no primary tubercles on the aboral side and the

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

B

 247

C

D

G E

F

H

Fig. 3.190: Nacospatangus altus. (A–D) Test length 49 mm; Noto Peninsula, Japan. (A) Aboral side: the anterior ambulacrum is bordered by larger tubercles. There are only a few primary tubercles in the lateroposterior interambulacra near the ambitus. (B) Oral side: the naked areas in the posterior ambulacra are narrow. (C) Side view: the aboral side is gently arched, the ambitus being rather sharp. (D) Posterior side viewed obliquely: above the subanal fasciole, there is a deeply sunken invagination. The anal opening itself is positioned at its upper side. (E, F) Specimen with spines, length of test 54 mm; Noto Peninsula, Japan. (E) Aboral side. The general spination is short and uniform, the large primary spines are lost. (F) Oral side. (G) Anterior ambulacrum. Scale bar is 4 mm. (H) Peristome, labrum, and adjacent ambulacral plates: the labrum is long and narrow. Scale bar is 6 mm. After Mortensen (1951).

sternum is almost fully tuberculated, whereas in N. altus, it is naked adorally. Distribution: Northwest coast of Australia, at depths of 5 to 55 m (Fig. 3.191). Nacospatangus laevis (H.L. Clark, 1917) Test: Regularly oval, no frontal notch; profile low, aboral side flattened, posterior interambulacrum elevated; test length may reach 42 mm. Apical system: Slightly anterior. Anterior ambulacrum: Flush, bordered by irregularly arranged larger tubercles.

Paired petals: Anterior row of anterior pair with strongly reduced pore pairs, posterior petals also rudimentary near the apical system. Oral side: Flat, labrum relatively short, sternum almost fully tuberculated; periproct at the truncated posterior side, not at all sunken. Fasciole: Subanal fasciole with three tube feet to each side. Primary tubercle: None or only one on the aboral side in the lateroposterior interambulacra near the ambitus; other tuberculation small and dense, and longer on the oral side.

248 

A

 3.3 Superorder Atelostomata von Zittel, 1879

B

C

E

F

D

G

H

I

J

K

Fig. 3.191: (A–G) Nacospatangus interruptus. (A–D) Test length 30 mm, Broome, Western Australia, from Studer (1880) (as Lonchophorus interruptus). (A) Oral side. (B) Subanal region. (C) Aboral side. (D) Side view, posterior is to the left. (E) Anterior ambulacrum. (F) Peristome, labrum, and adjacent ambulacral plates: the labrum is relatively short and broad. Scale bar is 4 mm. After Mortensen (1951). (G) Right anterior ambulacrum: the anterior row is rudimentary. Scale bar is 3 mm. After Mortensen (1951). (H–K) Nacospatangus laevis. Test length 38 mm; Cortez Bank, south of the Gulf of California, Mexico. ZMUC. (H) Aboral side: only a few pore pairs are developed in the anterior row of the anterior pair of petals. (I) Oral side: the sternum is almost fully tuberculated. (J) Side view. (K) Posterior side of test; width 21 mm. The periproct is not at all sunken. From H.L. Clark (1917).

Distribution: This species is known from the Channel Islands, Southern California, USA to Baja California and the Gulf of California, Mexico; 5 to 300 m deep. Nacospatangus oblongus (Mortensen, 1950) Main characteristics of the genus: maximum length 20 mm. Test: In outline elongated oval; in profile flattened, but distinctly keeled toward the posterior on the aboral and oral side; posterior side obliquely truncated. Anterior pair of petals: In anterior series, only one or two pore pairs developed. Periproct: Not at all sunken. Primary tubercles: Only one or two in lateral interambulacra just above the ambitus. Color: Brownish with silvery spines.

Distribution: This species is likely to be endemic to the Galapagos Islands, approximately 38 to 70 m. Remarks: N. oblongus is nearly related to N. laevis from California. In both species, the periproct is not sunken. N.  oblongus differs from that species by the relatively longer test and by the less-developed pore pairs in the anterior series of the anterior petals (Fig. 3.192). Nacospatangus tylotus (H.L. Clark, 1917) Test: In outline regularly oval, large specimens sometimes angular, anterior side straight, posterior side truncated; in profile gently arched from the rather sharp anterior edge to the vertical posterior end; posterior interambulacrum with a rounded keel; maximum length 43 mm. Apical system: Central, three gonopores.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

B

 249

C

D

E

F

G

H

I

Fig. 3.192: (A–C) Nacospatangus oblongus. Test length 20 mm, Galapagos Islands. ZMUC ECH 553. (A) Aboral side: only very few pore pairs are developed in the anterior rows of the anterior petals. (B) Oral side: the naked area in the posterior ambulacra and in front of the peristome is relatively large. (C) Side view. (D–I) Nacospatangus tylotus. Test length 41 mm; Kei Islands; Indonesia. ZMUC. (D) Aboral side: there are two to four large primary tubercles arranged in a row just above the ambitus. (E) Oral side: the phyllodes are short. (F) Side view: the test is gently arched and the plastron strongly inflated. (G) Posterior side with the subanal fasciole. (H) Anterior ambulacrum: the plates are narrow, the larger tubercles being confined to the adapical part. Scale bar is 4 mm. (I) Peristome, labrum, and adjacent ambulacral plates: the sternum is tuberculated except in the adoral part. Scale bar is 6 mm. After Mortensen (1951).

250 

 3.3 Superorder Atelostomata von Zittel, 1879

Anterior ambulacrum: Flush and conspicuously narrow, pores indistinct. Paired petals: With rudimentary pore pairs adapically, especially in the anterior row of the anterior petals; in posterior pair of petals adapically converging rows of pore pairs. Oral side. Flat, except in the strongly inflated plastron; labral plate relatively short, sternal plates tuberculated except adorally; periproct at the posterior face, not sunken. Subanal fasciole: Shield-shaped with two or three tube feet. Primary tubercles: Large, in a series of two to four near the lateral margin; larger tubercles bordering the anterior ambulacrum only adapically; general tuberculation fine; large interambulacral tubercles on the oral side with deep areoles, associated with distinct internal swellings. Color: The test is of uniform light brown with a pinkish tint, the aboral primary spines are white. Distribution: The species is recorded in the Malayan region and the Philippines; bathymetric range 235 to 1,460 m (Fig. 3.193). Genus Pycnolampas A. Agassiz & H.L. Clark, 1907 The genus is monospecific. Pycnolampas oviformis A. Agassiz & H.L. Clark, 1907 Only a couple of immature specimens are known. They measure between 11 and 22 mm. Test: Very delicate, in outline ovoid, slightly narrowing posteriorly, no anterior notch; in profile depressed, posterior interambulacrum slightly keeled. Apical system: Anterior, young specimens, no gonopores developed. Anterior ambulacrum: Flush and narrow, minute pores. Paired ambulacra: Flush, subpetaloid. Oral side: Flat, plastron forming a keel posteriorly; peristome not at all sunken, periproct on posterior truncated side; labral plate long, wedge-shaped, reaches to the third adjacent ambulacral plate; sternal plates triangular and almost fully tuberculated. Fascioles: Peripetalous fasciole thin, laterally not indented, but pointed in the posterior interambulacrum; subanal fasciole broad with three pore pairs to each side. Tuberculation: Heterogeneous, with some larger tubercles scattered inside the peripetalous fasciole.

Occurrence: This species is only known from the Hawaiian Islands. Remarks: Mortensen (1950) suggested that these specimens might be juveniles of Linopneustes. Genus Spatagobrissus H.L. Clark, 1923 Two species are known and are rather similar: Spatago­ brissus mirabilis is recorded from the South African coast, whereas Spatagobrissus incus is from Southern Australia. Spatagobrissus mirabilis H.L. Clark, 1927 Test: Oval in outline with a slightly pointed posterior end; depressed in profile, upper side slightly sloping toward the anterior; thick margin; length may reach 110 mm. Apical system: Subcentral, four gonopores, madreporite expanding to the posterior. Anterior ambulacrum: Perfectly flush, microscopic pore pairs. Paired petals: Of approximately the same length, interporiferous zone well-developed, distally closed; pore zones slightly depressed. Oral side: Faintly convex, peristome semilunate, tuberculated sternum triangular, the adjacent ambulacra hardly flexed; periproct low on the short undercut posterior end below the margin. Fascioles: Peripetalous fasciole surrounds the distal ends of the petals almost parallel to the margin; subanal fasciole on oral side, shield-shaped, rounded and anteriorly pointed, no pores enclosed. Spination: Numerous primary spines within peripetalous fasciole being distributed over the entire anterior part of the test, short and fine over the lower part of the posterior half, conspicuously indicated by small and densely distributed tubercles. Color: In life, the spines are brownish; the bare test is white. Distribution: S. mirabilis is restricted to South Africa, known only from the Cape Peninsula eastward to Walker Bay in shallow water (Fig. 3.194). Spatagobrissus incus Baker & Rowe, 1990 This species differs from S. mirabilis as follows: The apical system and consequently the area enclosed by the peripetal fasciole lies more anterior. The columns of pore pairs are reduced adapically, mainly in the anterior

▸ Fig. 3.194: Spatagobrissus mirabilis. (A–D) Test length 51 mm; Cape Town, South Africa. (A) Aboral side: the peripetalous fasciole follows the distal ends of the petals nearly parallel to the margin. (B) Oral side: the periproct is fully visible in oral view. (C) Side view: the upper side is flattened, sloping obliquely to the thick anterior margin. (D) Apical system. The madreporite do not far expand to the posterior. Scale bar is 1 mm. After Mortensen (1951). (E–H) Specimen with spines; length 59 mm; Cape Town, South Africa. (E) Aboral side: larger spines are distributed around the anterior half of test and within the whole peripetalous fasciole. (F) Oral side. (G) Side view. (H) The subanal fasciole is shield-shaped, pointed toward the peristome. Scale bar is 5 mm.

3.3.2 Order Spatangoida L. Agassiz, 1840 



 251

B

A C

Fig. 3.193: Pycnolampas oviformis. Test length 22 mm; Hawaiian Islands, Central Pacific. From H.L. Clark (1917). (A) Aboral side: the peripetalous fasciole is very thin, being pointed at its posterior end. (B) Oral side: the labral plate is narrow and wedge-shaped, extending to the third adjacent ambulacral plate. (C) Posterior side: the shield-shaped subanal fasciole is well-developed.

A

B

C

D

E

F

G

H

252 

 3.3 Superorder Atelostomata von Zittel, 1879

pair of petals. The periproct is positioned at the top of the somewhat rounded posterior end. The shield-shaped subanal fasciole is wider than long and angular rather than rounded. Maximum length more than 80 mm. Distribution: Spatagobrissus incus is restricted to Southern Australia, known only from the Bass Strait to the Spencer Gulf at depths of 10 to 110 m. Biology: This species lives buried in coarse, gravelly sand in a depth of approximately 100 mm below the sea floor. It ploughs forward slowly in its burrow swallowing large quantities of sediment. It digests the organic particles and organisms encrusting the sand grains (Fig. 3.195).

3.3.2.9 Family Spatangidae Gray, 1825 The family is characterized by an ethmolytic apical disc; lanceolate and at most slightly depressed petals. Usually,

there is only a subanal fasciole (+peripetalous fasciole in Plethotaenia), and no anal branches. The aboral side shows scattered primary tubercles. Two genera are included in this family. Plethotaenia H.L. Clark, 1917. Caribbean. Two species. Spatangus Gray, 1825. Worldwide. Fourteen species or more. Literature: Chesher 1968. Genus Plethotaenia H.L. Clark, 1917 This genus is characterized by a distinct frontal notch and an irregular peripetalous fasciole, generally doubled or in parallel or net-like branches surrounding the petaloid area. In these features, it differs from the otherwise rather similar Spatangus. Test: Strong, nearly as wide as long, deep frontal notch, more or less high. Apical system: Subcentral, ethmolytic, four gonopores.

A

B

C

D

Fig. 3.195: Spatagobrissus incus. (A–C) Test length 61 mm; Spencer Gulf, South Australia. (A) Aboral side. (B) Oral side. (C) Side view. (D) Diagrammatic sketch of angular subanal fasciole.



Anterior ambulacrum: Broad, within peripetalous fasciole set with simple, nonpenicillate tube feet and weakly sunken, continuing into deep notch. Paired petals: Bowed, closed distally, pore pairs reduced adapically, anteriorly longer than posteriorly. Oral side: On the whole flat, peristome anterior, facing forward, overarched by lip; labrum short and wide, sternal plates long and parallel-sided, paired episternal. Periproct: On posterior truncated side, subcircular to transversely oval. Fascioles: Subanal fasciole shield-shaped or oval, pore pairs inconspicuous; peripetalous fasciole variable from a rudimentary, single line to a multiple one forming an irregular network; not indented laterally. Primary tubercles: Distinct within peripetalous fasciole and bordering the anterior ambulacrum and the posterior median ridge. Spines: Short (10% of length), sharply pointed. Chesher (1968) has described two species: Plethotaenia angularis Chesher, 1968. Restricted to the Caribbean region. Plethotaenia spatangoides (A. Agassiz, 1883). From New England to the Antilles. Biology: The two species can be found together in the same habitat, muddy to sandy sediments, at depths between 150 and 620 m. They differ in shape: in P. spatangoides, the test is high with a rounded margin; the posterior end is almost vertically truncated. In P. angularis, the test is flat, sloping to a sharp margin, and the posterior end is oblique and undercut. As they have not developed funnel-building tube feet, they live shallowly burrowed, the apical spines break through the sediment. With its high test, P. spatangoides moves slowly through the substrate, whereas P. angularis has a flat test, which enables the animal to move faster (Chesher 1968). Plethotaenia angularis Chesher, 1968 Main characteristics of the genus: Test: Low, length reaching 134 mm. This species differs from P. spatangoides according to the following features: Test is lower with overhanging, truncated posterior side. Frontal notch is deeper than wide. Outline is more angular. Aboral primary tubercles are mainly restricted to the area within the peripetalous fasciole. Color: Grayish-purple. Distribution: Caribbean from the Straits of Florida to Barbados, at a depth of 150 to 623 m (Fig. 3.196). Plethotaenia spatangoides (A. Agassiz, 1883) Main characteristics of the genus: maximum length may reach 112 mm.

3.3.2 Order Spatangoida L. Agassiz, 1840 

 253

The test is strongly inflated, in outline rounded; frontal notch shallow, wider than deep. The labral plate is fully tuberculated. Peripetalous fasciole net-shaped; subanal fasciole encircles a slight projection. Primary tubercles: Aborally numerous and extending onto interambulacra 1 and 4 beyond the petals. Color: Grayish-purple. Distribution: The species is known in the West Atlantic from Martha’s Vineyard to the Lesser Antilles. Bathymetric range 150 to 623 m. Genus Spatangus Gray, 1825 There are 14 Recent species distributed in all oceans, predominantly in deeper water. The members of this genus look very similar, there are few features for differentiation. They are characterized by a heart-shaped test with a more or less deep frontal notch; aboral side domed with rounded margin. Apical system: Ethmolytic, four gonopores. Anterior ambulacrum: Small, simple pores, no tube feet for funnel-building. Paired petals: Lanceolate, flush with the test; pore pairs large and conjugate; however, anterior row of pore pairs in anterior pair of petals adapically rudimentary. Oral side: Rather flat, peristome wider than long, overarched by lip; labrum anvil-shaped, sternal plates broad and tuberculated. Periproct: On short undercut posterior side. Fasciole: Subanal fasciole well-developed, shield-shaped or bilobed; no peripetalous fasciole. Tuberculation: Heterogeneous, enlarged primary tubercles along the anterior ambulacrum, on interambulacrum 5 and, to a varying degree, scattered on the lateral interambulacra. Color: The test is usually purple to grayish pink. Biology: The species of Spatangus are found from the sublittoral to the deep sea at 1,000 m depth living buried in the sediment. They do not build respiratory funnels to the surface, but construct one or two sanitary tubes for waste matter using the specialized tube feet and spines developed within the subanal fasciole. Spatangus altus Mortensen, 1907. China Sea. Spatangus beryl Fell 1963. New Zealand. Spatangus californicus H.L. Clark, 1917. Gulf of California. Spatangus capensis Döderlein, 1906. Around South Africa. Spatangus diomedeae Fell, 1963. Philippines. Spatangus inermis Mortensen, 1913. Mediterranean. Spatangus luetkeni A. Agassiz, 1872. Japan and Hawaii. Spatangus mathesoni Mc Knight, 1968. Restricted to New Zealand.

254 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

C

D

F

G

E

Fig. 3.196: (A–D) Plethotaenia angularis. Test length 95 mm; north of Cuba. ZMUC. (A) Aboral side: the primary tubercles are mainly restricted to the area within the peripetalous fasciole. (B) Oral side. (C) Profile. (D) Side view, test 116 mm long. The test is usually low, with an overhanging and truncated posterior side. Sketch after a photo in Chesher (1968). (E–G) Plethotaenia spatangoides. Test length 112 mm long. (E) Side view. The test is highly vaulted, the posterior side rounded. (F) Upper side: the peripetalous fasciole passes in several branches around the petals. Length approximately 100 mm (primary tubercles omitted). (G) Test length 112 mm; north of Cuba. Aboral side: the primary tubercles extend laterally beyond the peripetalous fasciole. Sketch after a photo in Chesher (1968).

Spatangus multispinus Mortensen, 1925. Restricted to New Zealand. Spatangus pallidus H.L. Clark, 1908. Japan. Spatangus paucituberculatus A. Agassiz & H.L. Clark, 1907. Southeast coast of Australia till Juan Fernandez Islands.

Spatangus purpureus O.F. Müller, 1776. North Atlantic and the Mediterranean. Spatangus raschi Lovén, 1869. North East Atlantic. Spatangus thor Fell, 1963. Restricted to New Zealand.

3.3.2 Order Spatangoida L. Agassiz, 1840 



 255

Tab. 3.11: The most common species of Spatangus. S. californicus

S. capensis

S. purpureus

S. raschi

Subanal fasciole shield-shaped

Subanal fasciole shield-shaped

Subanal fasciole broader than long, bilobed

Subanal fasciole shield-shaped

Primary tubercles in upper interambulacra within fasciole

Primary tubercles numerous, not Primary tubercles in upper found in ambulacra below petals interambulacra within fasciole

Posterior end undercut to almost Posterior end undercut vertical

The species shows a certain degree of variation, and they are sometimes extremely difficult to distinguish. Even the pedicellariae give no reliable differentiation. Remarks: The subgenus Granopatagus was erected by Lambert (1915) for S. inermis and S. paucituberculatus, ­differing from Spatangus in lacking primary tubercles on the aboral lateral interambulacra. However, there are intermediates recorded in the number of primary tubercles, which seem to make this distinction no longer valid (Tab. 3.11). Shigei (1986) synonymized S. luetkeni with S. paucitu­ berculatus and S. pallidus after having compared numerous specimens with differing numbers of primary tubercles. Baker and Rowe (1990) did not go as far as Shigei. They examined many specimens including the type material and defined the two species, S. luetkeni and S. paucitubercula­ tus. However, there occur specimens, especially around New Zealand, which show such a great range of variation in number of aboral primary tubercles and general shapes, that their identification is unclear. Literature: Döderlein 1905; Mortensen 1907; 1925, 1927, 1951; Clark 1917; Fell 1963; McKnight 1968, 1969; Shigei 1986; Baker & Rowe 1990; Rowe & Gates 1995; Mironov 2006; van Noordenburg 2008. Spatangus altus Mortensen, 1907 This very rare species recalls in shape Spatangus raschi and S. capensis, but it differs in the number of pore pairs within the subanal fasciole: in S. altus, there are three pore pairs to each side; in the two other species, there are only two pore pairs. There are regular transverse series of primary tubercles down to the ambitus in the lateral interambulacra, but none adapically. Occurrence: China Sea, depth unknown (Fig. 3.197). Spatangus beryl Fell, 1963 Main characteristics of the genus: test more than 120 mm long, robust. Anterior ambulacrum: Notch shallow.

Posterior end undercut

Primary tubercles numerous, distributed also in ambulacra below petals Posterior end undercut

Periproct: Overhung by anal rostrum, visible from below. Subanal fasciole: With a posterior re-entrant angle or reentrant curve. Primary tubercles: Larger ones on the posterior interambulacrum and to both sides of the frontal ambulacrum; only one or two whitish larger tubercles in the paired interambulacra. Color: Deep reddish purple. Distribution: Southeastern area, Foveaux Strait, New Zealand, in moderate depth and coarse sediments. Spatangus californicus H.L. Clark, 1917 The species is very much like S. purpureus. It differs in the steeply truncated posterior end, the shieldshaped subanal fasciole protrudes slightly, and the plastron is keeled. Distribution: The species is reported from Southern ­California, USA, Baja California, and the Gulf, Mexico, from depths of 55 to 410 m. Spatangus capensis Döderlein, 1905 Main characteristics of the genus: length may reach 120 mm. Resembles S. raschi with parallel plastron and shield-shaped subanal fasciole, which is not very broad. Primary tubercles: In zigzag rows in aboral lateral interambulacra; contrary to S. raschi no, primary tubercles in ambulacra beyond petals. Posterior end: Obliquely truncated, periproct visible from below. Distribution: Around South Africa from off East London to Saldanha Bay; 70 to 500 m. Spatangus diomedeae Fell, 1963 Main characteristics of genus: Test: Fragile; anterior notch deep and broad. Oral side: Peristome overhung by labrum. Primary tubercles: All lateral interambulacra naked; inconspicuous tubercles in posterior interambulacrum and along anterior ambulacrum. Distribution: Philippines.

256 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

C

D

Fig. 3.197: (A) Spatangus altus. Length 85 mm; China Sea. ZMUC Holotype ECH 35. There are no primary tubercles adapically. (B) Spatangus californicus. Test length 63 mm; Baja California, Mexico. Aboral side: the tubercles follow the line of the plate sutures in the interambulacra. (C, D) Spatangus capensis. Test length 92 mm; Simon’s Bay, South Africa. ZMH. (C) Aboral side: the primary spines are very numerous. (D) Test length 92 mm, Simons Bay, South Africa. MfN Berlin, ECH 7266, syntype. Aboral side.

Spatangus inermis Mortensen, 1913 Main characteristics of the genus: test length up to 120 mm. Periproct: Not overhung, visible neither from below nor above.

Primary tubercles: In the odd interambulacrum, it varies from 0 to 26; in the posterior pair of interambulacra, there are none. Color: Light yellowish to light violet.



Distribution: Gulf of Naples, approximately 50 m; Banyuls Sea, 20 to 25 m; Northeast Atlantic Seamounts Ampere, Great Meteor, Irving and Seine, 60 to 313 m (Mironov 2008). Spatangus luetkeni A. Agassiz, 1872 Main characteristics of the genus: maximum length 133 mm. Test: Low, upper side regularly domed; posterior interambulacrum slightly elevated or keeled, in outline broadly heart-shaped with a deep frontal notch; posterior side obliquely truncated or nearly vertical. Petals: Bowed, distally nearly closed. Primary tubercles: A few or more adapically in interambulacra 1 and 4, somewhat concentrated along the midline; additional larger tubercles irregularly scattered along the median line of the posterior interambulacrum and in interambulacra 2 and 3, bordering the anterior ambulacrum. Oral side: Labrum protruding and with lip, sternal plates relatively narrow. Subanal fasciole: With two pore pairs on each side, shape varies from heart-shaped to bilobed. Distribution: From Japan and Hawaiian Islands to southeast Australia and New Zealand; bathymetric range 19 to 999 m (Baker & Rowe 1990).

3.3.2 Order Spatangoida L. Agassiz, 1840 

 257

Color: The test has a deep rich violet, fading out after preservation. Distribution: Bass Strait, Australia and New Zealand, at depths of 36 to 1,006 m. Biology: This species is recorded as living in fine sediments (McKnight 1969) (Fig. 3.198). Spatangus pallidus H.L. Clark, 1908 Main characteristics of the genus: Shigei (1986) felt that it was synonymous with S. luetkeni. All the specimens were young. They differ from S. luetkeni according to the following features: Oral side: Strongly keeled sternum. Fasciole: Subanal fasciole broad and bilobed; two pores to each side. Color: Light color. Distribution: Only found at Sagami Bay, Japan; depth 90 to 100 m.

Spatangus mathesoni McKnight, 1968 Main characteristics of the genus: length may reach up to 110 mm. Test: Broad heart-shaped, deep frontal notch, shallow notch at the posterior end. Oral side: Periproct not visible from below or above. Subanal fasciole: Reniform or transversely oval. Primary tubercles: Six to eight enlarged and sunken tubercles in the posterolateral ambulacra. Color: Deep reddish-brown in life and purple in its preserved state. Distribution: Off New Zealand, North Island, Tasman Sea, and Chatham Rise; depth 650 to 1,050 m.

Spatangus paucituberculatus A. Agassiz & H.L. Clark, 1907 Main characteristics of the genus; length up to 132 mm. Test: Large, aboral side convex sloping to a rather sharp margin anteriorly and to the overhanging, truncated posterior side, very deep anterior notch. Oral side: Labrum almost naked with only very small tubercles; sternum slightly bowed; periproct transversely oval. Subanal fasciole: Shield-shaped, pointed below, upper band straight or slightly indented. Primary tubercles: Bordering the anterior ambulacrum within the deeply sunken sulcus; some scattered primary tubercles along the posterior midline, no primary tubercles in lateral interambulacra. Color: In the living echinoid, the test is deep purple, the spines are white to silvery. Distribution: This species is recorded in the entire western and central Pacific from Indonesia and the Philippines to the Hawaiian Islands, and also at Juan Fernandez Island off the Chilean coast. Bathymetric range 468 to 604 m.

Spatangus multispinus Mortensen, 1925 Main characteristics of the genus: length may reach 100 mm. Test: Low, conspicuous frontal notch; posterior end vertically truncated or concave. Aboral surface with numerous primary tubercles arranged in zigzag lines in all interambulacra, smaller irregular ones bordering the anterior ambulacrum; a few primary tubercles also in the lateral ambulacra beyond the petals. Oral side: Sternum nearly parallel-sided, tuberculation continues onto the labrum (contrary to S. raschi, in which there is a naked area). Subanal fasciole: Heart-shaped. Primary spines: Long, erect, and fairly dense.

Spatangus purpureus O.F. Muller, 1776 Test: Strong, outline heart-shaped with conspicuous anterior notch, narrowing posteriorly; in profile not high, but more or less regularly convex; posterior interambulacrum raised; length up to 120 mm. Apical system: Slightly anterior or subcentral, four gonopores, madreporite extending to the posterior. Anterior ambulacrum: Grooved, depth increasing toward the margin; adjacent interambulacra somewhat keeled; pore pairs small and not specialized. Paired petals: Broadly lanceolate, pore pairs large and connected by a small furrow; pore pairs rudimentary in the anterior-most row.

258 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

C

D

E

G

F

H

I

Fig. 3.198: (A) Spatangus luetkeni. Test length 48.5 mm, Hakodate, Northern Japan. Photo ZSM, München, courtesy of B. Ruthensteiner. (A) Aboral side: there are a few distinct primary tubercles in interambulacra 1 and 4 close to the apical system. Additionally, some larger tubercles are scattered on the posterior interambulacrum and bordering the anterior ambulacrum. (B) Spatangus mathesoni. Paratype, length 94 mm, Tasman Sea. Sketch after McKnight (1968). Aboral side. (C–E) Spatangus multispinus. Test length 80 mm, New Zealand. ZMUC. (C) Aboral side: the surface is covered by primary tubercles arranged in short oblique series (D) Oral side. The sternum is fully tuberculated. (E) Profile: the truncated posterior end is slightly concave. (F) Spatangus pallidus, young specimen, from H.L. Clark (1917). (G–I) Spatangus paucituberculatus. Test length 102 mm, South China Sea. (G) Aboral side (H) Oral side. (I) Profile.

3.3.2 Order Spatangoida L. Agassiz, 1840 



Oral side: Flattened; plastron tuberculated from the labrum to the rear; adjacent ambulacra slightly bowed; labral plate covering the kidney-shaped peristome. Periproct: On the undercut, truncated posterior end, wider than long. Subanal fasciole: Broader than long, bilobed, not pointed toward the peristome. Tubercles: Primary tubercles in all interambulacra, often in zigzag series along the plate sutures. Spines: Primaries long, the others short and dense; short on plastron, longer laterally. Color: In life, the urchin is deep purple, the primary spines are lighter or whitish. The bare test is white.

A

B

 259

Distribution: S. purpureus is known from the North Cape, Norway to Iceland, along the European coasts, in the Mediterranean Sea, and along the coast of West Africa southward to Senegal. It lives from the lowermost tide limit to a depth of approximately 900 m (Fig. 3.199). Spatangus raschi Lovén, 1869 This species resembles S. purpureus, but differs in the following distinct features: Primary tubercles: Smaller and scattered all over the aboral side. They also extend over the ambulacra between the end of the petals and the margin. Paired petals: Narrower than in S. purpureus.

D

C

E

F

G

H

Fig. 3.199: (A–D) Spatangus purpureus. (A–C) Test length 101 mm; Greece. (A) Aboral side: the petals are broad and the series of pore pairs arched. (B) Oral side: the plastron is fully tuberculated, the adjacent ambulacra slightly flexed. (C) Side view: the test is domed above and truncated at the posterior end. (D, E) Specimen with spines; length of test 72 mm; Iceland. (D) Aboral side. The echinoid is purple. (E) Oral side: the plastron is entirely covered by short spines. (F–H) Spatangus raschi. Test length 91 mm; North Norway; ZMH. (F) Aboral side: the lanceolate paired petals are narrow. (G) Oral side: the fully tuberculated plastron is narrow and parallel-sided, the adjacent ambulacra are broader. (H) Side view: the rather small primary tubercles are distributed over the aboral side, not only in the interambulacra but also in the ambulacra between the distal end of the petals and the margin.

260 

 3.3 Superorder Atelostomata von Zittel, 1879

Oral side: Plastron is parallel-sided and narrower; labrum covers the peristome to a larger degree. Subanal fasciole: Shield-shaped, may be reduced or even lacking in adults. The test may be high domed or rather low; maximum length of test approximately 100 mm. Color: The living sea urchin is, as in Spatangus purpura­ tus, deep purple. Distribution: S. raschi is found from Northern Norway and south of Iceland to the Strait of Gibraltar, in depths of 150 to 1,000 m. It has not been recorded from the American coast. Spatangus thor Fell,1963 Main characteristics of the genus: length up to 120 mm or more. This species is found at the same place as S. beryl. It differs from that species by the number of primary tubercles, S. beryl is naked. Subanal fasciole: Heart-shaped with two pores to each side, another pair just outside. Primary tubercles: Fifteen to seventeen enlarged whitish tubercules in each posterolateral ambulacrum in short zigzag series; vertical series of tubercles along dorsal midline of posterior unpaired interambulacrum and on every side of the frontal ambulacrum. Distribution: Southeastern region, Foveaux Strait, New Zealand, at depths of 29 to 34 m and in coarse sediments.

3.3.2.10 Family Macropneustidae Lambert, 1895 A.B. Smith listed several fossil genera in this family, the very rare and poorly known genus Argopatagus (= Phrissocystis) is added tentatively. The family is characterized by an ethmolytic apical system, with and without a subanal fasciole. Genus Argopatagus A. Agassiz, 1879 Argopatagus is almost equal to the genus Phrissocystis A. Agassiz, 1898, but it differs in having developed a subanal fasciole, whereas in Phrissocystis (P. aculeata and P. multispina), a fasciole is lacking. Mortensen (1950) believed the two genera synonymous: Test: Very delicate and fragile; oval outline, low profile; maximum length more than 120 mm. Apical system: Four gonopores, probably fused genital plates, madrepores extending to the rear of the posterior ocular plates. Ambulacra: All apetaloid and flush, pore pairs with tube feet only on the four to five uppermost plates, the others display only minute single pores. Oral side: Flat, slightly sunken toward the peristome, phyllodes with three to four penicillate podia, posterior ambulacra naked; labral plate elongate, sternum not surely known; peristome overarched by lip, wider than long; periproct marginal.

Subanal fasciole: May or may not be present. Primary tubercles: Aboral ambulacra and interambulacra with primary tubercles concentrated in the middle of the plates. Spines: Aborally thorny, oral spines smooth; length up to 35 mm. The genus consists of four species: Argopatagus aculeata A. Agassiz, 1898. Cocos Island. Argopatagus multispina A. Agassiz & H.L. Clark, 1907. ­Probably Hawaii Islands. Argopatagus planus A. Agassiz & H.L. Clark, 1907. ­Kagoshima Gulf, Japan. Argopatagus vitreus A. Agassiz, 1879 (= Phrissocystis humilis de Meijere, 1903). Indo-West Pacific. Argopatagus aculeata A. Agassiz, 1898 Main characteristics of the genus: only fragments are known. Length more than 120 mm, subanal fasciole absent; two to eight primary tubercles in the midzone of the aboral plates. Color: Yellowish brown. Distribution: Known only from the Cocos Islands, at a depth of 1,820 m (Fig. 3.200). Argopatagus multispina A. Agassiz & H.L. Clark, 1907 Characteristics of the genus: length approximately 100 to 150 mm. This species differs from Argopatagus aculeata by the more numerous primary spines on the aboral plates having 8 to 12 in the midzone, fewer adapically. Color: Deep red-brown. Occurrence: Probably from Hawaii, depth unknown. Argopatagus planus A. Agassiz & H.L. Clark, 1907 Only one young and immature specimen with a length of 28 mm is known. Main characteristics of the genus: Test: Anteriorly rounded and very low, no frontal sulcus, posterior part elevated, posterior face vertically truncated. Apical system: Slightly posterior. Oral side: Posterior interambulacra not contacting the peristome. Subanal fasciole: Well-developed, bilobed. Occurrence: Found only in the Kagoshima Gulf, Southern Japan, at a depth of 790 m. Argopatagus vitreus A. Agassiz, 1879 (Phrissocystis humilis de Meijere, 1903) This is the best known species of the genus. Characteristics of the genus: length up to 120 mm. Apical system: Slightly posterior; four gonopores. Ambulacra: Pore pairs only close to the apical system, with black tube feet forming a dark star, following pores microscopic and simple.

3.3.2 Order Spatangoida L. Agassiz, 1840 

A

C

B

 261

E

D

F

G

H

I

Fig. 3.200: (A–E) Argopatagus aculeata. Test length 120 mm; found only east of the Cocos Islands, East Pacific, 1,820 m. From A. Agassiz (1904). Only fragments are known: (A) Part of aboral side. (B) Enlargement of apical system. (C) Peristome overarched by the lip. (D) Periproct. (E) Posterior end. (F) Argopatagus multispinus. From H.L. Clark (1917). Adapical part of aboral side with thorny primary spines. Maximum length of spines approximately 40 mm. (G–I) Argopatagus planus. Test length 28 mm, Kagoshima Gulf, Southern Japan. From Clark (1917). (G) Aboral side. (H) Oral side. (I) Side view.

Oral side: Flat, labrum and sternum tuberculated. Subanal fasciole: Bilobed and distinct. Primary spines: On the aboral side coarse and thorny, length up to 36 mm; on the sternum spatulate. Distribution: Widely known in the Indo-West Pacific from the Maldives to Hawaii and from Japan to the Arafura Sea; bathymetric range 460 to 2,250 m. Remarks: A. vitreus differs from A. planus in its sloping posterior end and in its oral interambulacral plates, which contact the peristome. In A. planus, the interambulacral plates are occluded from the peristome (Fig. 3.201).

into the anal funnel. Additionally, a peripetalous fasciole may be developed on the aboral side. Two families belong to this suborder: Aeropsidae Lambert, 1896. One genus. Micrasteridae Lambert, 1920. Ethmophract apical system, two extant genera.

Suborder Micrasterina Fischer, 1966

Genus Aeropsis Mortensen, 1907 Mortensen thought this genus to be a primitive one. However, the contrary has been proven to be correct: Aerop­ sis is a highly specialized deep sea echinoid. It belonged to

The micrasterine echinoids are typically characterized by the subanal fasciole, serving to sweep the waste water

3.3.2.11 Family Aeropsidae Lambert, 1896 Only one genus is included in this family, Aeropsis. It is characterized by a strongly elongated test and a peripetalous fasciole, which passes obliquely from the aboral to the oral side surrounding the frontal part of the test.

262 

A

 3.3 Superorder Atelostomata von Zittel, 1879

C

B

D

G E

F

Fig. 3.201: Argopatagus vitreus. (A–C) Test length 50 mm, Southern Japan. From H.L. Clark (1917). (A) Aboral side. (B) Oral side: the plates of the posterior paired interambulacra contact the peristome. (C) Side view: the test is low, the aboral side being slightly arched. (D) Apical system and adjacent plates. Scale bar is 5 mm. After Mortensen (1950). (E, F) Specimen with spines, length 93 mm, Flores Sea, Indonesia. From de Meijere (1903) as Phrissocystis humilis. (E) Aboral side: the primary spines are long, coarse, and thorny. (F) Oral side: the spines on the sternum are spatulate. (G) Fragment of the aboral side. Flores Sea, Indonesia. ZMUC. The adapical area is rather naked, but toward the margin, the number of primary tubercles, concentrated in the middle of each plate, increases. The tubercles are perforated and crenulated.

the Hemiasterida but it is now a member of the Micrasterina (Kroh & Smith 2010). Two very rare species are known: Aeropsis fulva (A. Agassiz, 1898). East and North Pacific. Aeropsis rostrata (Thomson, 1877). Northern Atlantic. Aeropsis fulva (A. Agassiz, 1898) Test: Very elongated, sides compressed, aboral side flattened and sloping to the anterior margin, no anterior sulcus, oral side convex with a high posterior margin; length up to 63 mm.

Apical system: Anterior, ethmophract, two gonopores in two fused genital plates to four gonopores in four genital plates, long genital papillae. Anterior ambulacrum: Broad, slightly depressed adapically, with two series of large pore pairs, set with very conspicuous tube feet, which end in a wide disk supported by numerous delicate, radiating leaf-like processes. Paired ambulacra: Apetaloid with small, round pore pairs. Oral side: Peristome slightly anterior, round or oval; phyllodes quite rudimentary; periproct on the posterior upper side.



Peripetalous fasciole: Passing from the aboral side obliquely down to the anterior oral side. Spines: Within the fasciole slender and simple, others more or less spatulate, most distinct on the plastron. Distribution: This species is recorded from the Pacific, from the Bering Sea in the north, off Peru in the east, and the Arafura Sea in the southwest. Bathymetric range 1,465 to 5,200 m (Fig. 3.202). Aeropsis rostrata (Thomson, 1877) This species is very similar to A. fulva, but its test is distinctly shorter. In both, the number of gonopores in the ethmophract apical system varies from two to four. A. ros­ trata has likewise large, blossom-shaped tube feet in the anterior ambulacrum and spatulate spines. The two species differ in geographic distribution: A. rostrata is recorded only in the northwest Atlantic from the Davis Strait to off Cape Hatteras and Cape Cod, USA; bathymetric range 2,550 to 3,125 m.

3.3.2.12 Family Micrasteridae Lambert, 1920 The genera of this family were very common and dated back to the Upper Cretaceous. Only two species of Cyclaster survive in New Zealand and in the Western Pacific to East Australia. Usually, the family is characterized by a subanal fasciole and an ethmophract apical system. Cyclaster Cotteau, 1856. West Pacific and New Zealand. Isopatagus Mortensen, 1948 & Desor, 1858. West Pacific. Genus Cyclaster Cotteau, 1856 The genus has developed three gonopores, the pore in genital plate 2 is lacking. It consists of two species: Cyclaster recens Mortensen, 1950. West Pacific. Cyclaster regalis Baker, 1969. Endemic to New Zealand. Cyclaster recens Mortensen, 1950 Test: Trapezoid elongated, anteriorly broadest, narrowing toward the posterior; shallow frontal depression; highly vaulted, aborally arched, sloping to the anterior margin; posterior side vertically truncated; maximum length approximately 70 mm. Apical system: Ethmophract, three gonopores, none in the extended madreporite. Anterior ambulacrum: Slightly sunken; pore pairs very distinct, oblique, close set adapically, more distant toward the margin, in which they almost disappear. Paired petals: Paired petals sunken, narrow, with parallel sides, distally open. Oral side: Slightly convex, peristome small, at the end of a shallow furrow and a little distant from the margin,

3.3.2 Order Spatangoida L. Agassiz, 1840 

 263

small lip, phyllode inconspicuous; labral plate long and tuberculated; periproct small, round, high on the posterior side. Fascioles: Peripetalous fasciole not reaching frontal ambulacrum; subanal fasciole well-developed, almost rectangular in outline; three pore pairs on each side. Tuberculation: Aborally small, dense and uniform, orally small and sparser. Distribution: This species is recorded from off Vietnam. Remarks: C. recens differs from C. regalis by its shallower frontal sulcus and by the peristome positioned at a distance from the anterior margin. Cyclaster regalis Baker, 1969 Test: High, upper side convex, anteriorly sloping, toward the posterior side slightly keeled, at the rear truncated; length barely 90 mm. Apical system: Small, ethmophract with three gonopores, none in the madreporite; in females gonopores larger than in males. Anterior ambulacrum: Sunken, deeply notched at the ambitus; set with differentiated pore pairs. Paired petals: Short and depressed. Oral side: Peristome very anterior, covered by a prominent lip; labrum elongated; plastron inflated and densely set with tubercles. Fascioles: Peripetalous fasciole present, subanal fasciole bilobed. Primary spines: On aboral side approximately 9 mm long, longer on oral side, spatulate on the plastron, and often covered by horny envelope. Color: The live echinoid is reddish brown with pinkish spines; the bare test is yellowish-gray. Distribution: C. regalis is restricted to New Zealand in depths of 150 to 500 m. Remarks: Cyclaster recens Mortensen, 1950, rarely recorded in the West Pacific and East Australia, differs in having a less depressed anterior furrow and a rectangular subanal fasciole (Fig. 3.203). Genus Isopatagus Desor, 1858 The genus consists of a single extant species. Isopatagus obovatus Mortensen, 1948 Test: Ovoid, no frontal notch; aboral side highly vaulted; length up to 102 mm. Apical system: Ethmophract with three gonopores. Petals: All five with low, dense plates, outer pore larger than inner one; pore pairs continue beyond petals distantly spaced toward peristome; anterior petal half the length of paired petals.

264 

 3.3 Superorder Atelostomata von Zittel, 1879

A

C

B

E

G

D

F

H

I

J

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

C

B

D

F

 265

E

G

H

Fig. 3.203: (A–E) Cyclaster recens. Test length 68 mm, Vietnam. ZMUC Holotype ECH 278. (A) Aboral side: there are only three gonopores. (B) Oral side: the frontal sulcus is shallow, leading to the peristome, which is distant from the margin. (C) Side view. (D) Ethmophract apical system: the enlarged madreporite extends toward the anterior. It has no gonopore. Scale bar is 2 mm. (E) Peristome and labrum. Scale bar is 6 mm, after Mortensen (1951). (F–H) Cyclaster regalis. Test length 77 mm, height 46 mm; Auckland Island, south of New Zealand, approximately 450 m. (F) Aboral side: the short petals are surrounded by the peripetalous fasciole. (G) Oral side. The plastron is prominent. (H) Side view. ◂ Fig. 3.202: (A–E) Aeropsis fulva. Test length 49 mm. Panama, Pacific coast; 3,680 m deep. ZMUC. (A) Aboral side. The podia in the anterior ambulacrum are strongly developed. They are folded here. (B) Oral side: the peristome is more or less oval. (C–E) Drawing, test length 35 mm; off Peru. From Agassiz (1904). (C) Aboral side with periproct. (D) Oral side with peristome: the peristome is elongated oval, the labral plate and the sternal plates being long and narrow. (E) Side view: the peripetalous fasciole encircles the anterior side obliquely from the aboral to the oral surface. (F, G) A. fulva. (F) Apical system: the number of gonopores varies from two to four. (G) The large tube foot in the anterior ambulacrum looks like a flower. Scale bar is 2 mm. After Mortensen (1950). (H–J) Aeropsis rostrata. Test length 15 mm, Davis Strait, Northern Atlantic. ZMUC. (H) Aboral side: the periproct is positioned high on the posterior aboral side. The fasciole passes from the aboral to the oral side encircling the sloping anterior side. (I) Oral side: the peristome is an elongated oval. (J) Apical system: the number of gonopores vary from two to four. Scale bar is 3 mm. After Mortensen (1950).

266 

 3.3 Superorder Atelostomata von Zittel, 1879

Oral side: Rather flat, plastron slightly elevated, peristome small, periproct inframarginal; labrum with prominent lip, sternal plates symmetrical. Fasciole: Lacking. Tubercles: Small; spines fine and short; length 5 mm. Occurrence: Only reported from the Sulu Sea at a depth of 890 m (Fig. 3.204).

Suborder Paleopneustina Markov & Soloviev, 2001 The suborder is characterized by having a marginal and a peripetalous fasciole in early ontogeny, which may be lost partly or entirely during development (Fig. 3.205). Literature: Chesher, 1968 and Markov & Soloviev, 2001. Six families belong to this suborder: Paleopneustidae A. Agassiz, 1904. Three extant genera. Pericosmidae Lambert, 1905. Two extant genera. Marginal and peripetalous fasciole coalesced anteriorly, the latter forming a branch over the test between the apical system and the periproct, no occluded ambulacral plates. Prenasteridae Lambert, 1905. Three extant genera. Schizasteridae Lambert, 1905. Twelve extant genera. Paleopneustina incertae sedis A. A single extant genus. Paleopneustina incertae sedis B. Five extant genera.

3.3.2.13 Family Paleopneustidae A. Agassiz, 1904 The family is characterized by large to very large tests, mostly without a frontal notch. Their petals are long, parallel-sided and flush or only slightly sunken. The pore zones are more or less distinctly open. The marginal and peripetalous fascioles are usually absent in adults. Literature: de Meijere 1904; Chesher 1968; David & De Ridder 1989; van Noordenburg 2008. The family consists of three genera: Paleopneustes A. Agassiz, 1873. Caribbean. One species. Peripatagus Koehler, 1895. North Atlantic, and off New Zealand. One species. Plesiozonus de Meijere, 1903. West Pacific. Three species. The genera differ in the length of the labrum; in Paleopneustes, it extends to the third adjoining plates; in Plesiozonus, it reaches only the second ambulacral plate. Genus Paleopneustes A. Agassiz, 1873 The genus is monospecific. Paleopneustes cristatus A. Agassiz, 1873 Test: High and regularly domed, with rather sharp edge; posterior end short and obliquely truncated; in outline ovoid without a frontal notch; length may reach 150 mm.

Apical system: Anterior, ethmolytic, three gonopores. Anterior ambulacrum: Flush, pores small, tube feet simple. Paired petals: Flush or only slightly sunken, pore series long, parallel-sided, distally open with occluded plates; interporiferous zone twice as broad as pore zones. Oral side: Flat, posterior ambulacra naked; peristome sunken; phyllodes short but distinct; periproct on short posterior face, visible from below; labrum protruding over the peristome, extending posteriorly to the third adjacent ambulacral plate, the two first plates being very low; sternal plates narrower than the episternal plates. Fascioles: Peripetalous and marginal fasciole present in juveniles, absent in adults. Primary tubercles: Small, irregularly scattered over the test. Spines: Short, straight, fragile and thorny, tuft of longer spines just in front of the apical system. Distribution: This species is widely spread in the Caribbean from Florida and the Gulf of Mexico to Colombia and Barbados; bathymetric range 76 to 805 m (Fig. 3.206). Genus Peripatagus Koehler, 1895 The genus is monospecific. Peripatagus cinctus Koehler, 1895 Test: Very fragile, virtually no frontal notch; ovoid in outline; upper side low arched, posterior end slightly truncated; oral side convex; nearly as long as wide, length may reach 20 mm. Apical system: Anterior, four or three genital pores that are lacking in the madreporite; few madrepores; ethmophract. Ambulacra: Not at all petaloid, a few single tiny pores, double only adapically; all ambulacra of same shape. Oral side: Peristome large, slightly anterior, labrum not prominent and not extending posteriorly beyond the middle of the first adjoining ambulacral plate; phyllodes poorly developed; periproct large, high on the posterior side. Fasciole: Marginal fasciole narrow, bending downward anteriorly, on the posterior side passing below the periproct. Tubercles: Short and uniform; irregularly scattered all over the test, those on the plastron somewhat larger carrying spatulate spines. Distribution: North Atlantic, Azores; depth 880 to 1,494 m; off Northeast New Zealand; 808 to 924 m. Genus Plesiozonus de Meijere, 1903 Test: High, upper side rounded or subconical; sharp margin; in outline broadly ovoid with shallow or no frontal notch; large to very large forms. Apical system: Anterior to subcentral, slightly depressed; ethmolytic with three gonopores.

3.3.2 Order Spatangoida L. Agassiz, 1840 



B

A

C

 267

D

E

F

Fig. 3.204: (A–C) Isopatagus obovatus. Test length 99 mm; Sulu Sea. ZMUC ECH 253. (A) Aboral side: the pore pairs in the petals are densely set, the ambulacral plates being very low. (B) Oral side: labrum and sternal plates are fully tuberculated. (C) Side view. The pore pairs beyond the petals are distinct, but distant-spaced. (D–F) Isopatagus obovatus. (D) Apical system. Scale bar is 2 mm. (E) Labrum and adjacent plates. (F) Petal in A V. Scale bar is 8 mm; after Mortensen (1951).

Fig. 3.205: Fascioles of Paleopneustidea. The marginal fasciole (green band) evolves early in ontogeny. While the periproct shifts toward the posterior, the marginal fasciole “moves” downward (adorally). Later, the peripetalous fasciole (orange band) evolves between the apical system and the periproct (adapically). In extant paleopneustins, the two fascioles are separated into a complete marginal fasciole positioned below the periproct (adoral) and a complete peripetalous fasciole above the latter and adapical of the periproct. These fascioles may become lost during ontogeny: It is likely that the young animal lives burrowed in the sediment, whereas the adult lives rather on the sea surface. Modified after Markov & Soloviev (2001).

Anterior ambulacrum: Narrow, slightly sunken or flush, small and simple pore pairs. Paired petals: Flush or slightly depressed, long and ­parallel-sided, distally open; occluded plates. Oral side: Flat, peristome kidney-shaped, facing forward; periproct inframarginal; posterior ambulacra naked; labral plate reaching to second adjacent ambulacral plate, sternal plates elongated. Fascioles: Peripetalous and marginal fascioles occur only in juveniles; sometimes, short parts of the fascioles are traceable in adults, e.g., below the anterior petals or below the periproct. Tubercles: Crenulate and perforate; small and uniform, scattered over the aboral surface; spines short (Tab. 3.12).

268 

 3.3 Superorder Atelostomata von Zittel, 1879

B

A

D

C

E

F

G

Fig. 3.206: (A–D) Paleopneustes cristatus. Test length 110 mm; off Barbados. ZMUC ECH 113. (A) Side view. (B) Aboral side: just in front of the apical system, some longer spines form a short tuft. (C) Oral side. (D) Apical system, after Mortensen (1950b). (E,F) Peripatagus cinctus. Specimen in the USNM collection. Test length scarcely 20 mm. Georgia, Cumberland Island. Photos courtesy of A. Kroh, NMW. (E) Aboral side. (F) Oral side. (G) Holotype from off the North Island of New Zealand, 19.6 mm; NZOI F868. Photo courtesy of O. Anderson.

3.3.2 Order Spatangoida L. Agassiz, 1840 



 269

Tab. 3.12: A comparison of three species from the Genus Plesiozonus: P. diomedeae

P. hirsutus

P. tenuis

Narrow interporiferous zones

Broad interporiferous zones

Broad interporiferous zones

Labrum adoral straight-cut

Labrum strongly protruding

Labrum moderately protruding

Petals distinctly sunken adapically, anterior pair longer

Petals with slight median groove, equal-sized

Petals flush, equal-sized

Outer pore of pair elongated

Outer pore of pair elongated

Pore pairs equal and round

Three species belong to this genus: Plesiozonus diomedeae Mortensen, 1948. Philippines. Plesiozonus hirsutus de Meijere, 1903. Philippines. Plesiozonus tenuis David & De Ridder, 1989. Philippines. Plesiozonus diomedeae Mortensen, 1948 Main characteristics of the genus: the length may reach 138 mm. Test: Strongly inflated, aborally arched or subconical; broadly ovoid in outline, with shallow but distinct frontal sulcus. Paired petals: Narrow, adapically sunken becoming flush toward the margin. Oral side: Flat, peristome sunken opening into a deep furrow toward the anterior; periproct positioned at the short, vertical, or obliquely truncated posterior face; labral plate straight-cut at the peristomial edge. Fascioles: Traces of a peripetalous fasciole may be present, e.g., at the end of the petals, parts of a very thin marginal fasciole sometimes occur adorally of the periproct. Distribution: This species is known only around the ­Philippines; bathymetric range 180 to 325 m (Fig. 3.207). Plesiozonus hirsutus de Meijere, 1903 Main characteristics of the genus: maximum length 140 mm. Test: Oval to ovoid, anteriorly rounded; aboral side strongly inflated, oral side convex. Paired petals: Flush, only along the median suture slightly depressed; broad interporiferous zone, wider than the pore zones, pore pairs large with the outer pore of a pair elongated. Oral side: Peristome kidney-shaped, strongly overarched by the labrum; shallow furrow toward the anterior margin; periproct positioned on the oral side. No traces of fascioles (Fig. 3.208). Plesiozonus tenuis David & De Ridder, 1989 Main characteristics of the genus: maximum length recorded only 76 mm. Typical for this species are the broad petals, flush with the test and with round or slightly transverse oval pores of equal sizes. The interporiferous zones

are wide. The labrum protrudes only moderately over the peristome. Distribution: Known only from the Philippines at a depth of 200 to 250 m.

3.3.2.14 Family Pericosmidae Lambert, 1905 The pericosmids differ from the paleopneustids by having shorter petals sunken throughout their length. In the Paleopneustidae the petals are longer, flush to slightly sunken adapically and distally more or less open. In the pericosmids the fascioles are well-developed, whereas they are absent in adult paleopneustids. The family consists of two extant genera: Faorina Gray, 1851. Andaman Islands, Sulu Sea to Southern China. One species. Pericosmus L. Agassiz, 1840. Indo West Pacific. Ten species. The two genera are distinguished in the following features: In Faorina, the labral plate is elongated and pointed toward the sternum; in Pericosmus, the labrum is stout broadly touching the sternum. Furthermore, in Faorina, the peripetalous fasciole passes closely around the distal ends of the petals but is not sharply indented in the lateral interambulacra as in Pericosmus (Fig. 3.209). Genus Faorina Gray, 1851 Chesher (1968) and Smith (2005) showed that the monospecific Faorina belongs to the family Pericosmidae, as in juveniles a peripetalous and a marginal fasciole are present. Faorina chinensis Gray, 1851 Test: Thick and rather strong, highly domed, highest point behind the apical system; outline heart-shaped with shallow frontal groove, posterior side truncated; test length up to 96 mm. Apical system: Slightly anterior with three gonopores. Anterior ambulacrum: Narrow and depressed; pore pairs uniserial.

270 

 3.3 Superorder Atelostomata von Zittel, 1879

B

A

C

D

E

Fig. 3.207: Plesiozonus diomedeae. (A–C) Test length 130 mm, Mindanao, Philippines. ZMUC ECH 137. (A) Aboral side: the interporiferous zone is narrow, the midline indented. (B) Oral side: the labral plate ends straight at the peristomial edge. (C) Side view: the test is regularly arched, the posterior face being short and oblique. Laterally, there are traces of the marginal fasciole. (D) Test length 92 mm, Aliguay, Philippines. Aboral side: the test is subcircular and the apical system is positioned anterior. (E) Specimen with spines; test length 98 mm, Philippines. Side view: the vertex lies anterior of the center, the posterior end is vertically truncated.

Paired petals: Sunken, narrow with parallel sides, posterior pair slightly shorter than anterior pair; naked, depressed midlines leading from petals around the margin to oral side. Oral side: Convex; peristome kidney-shaped, facing obliquely forward; labral plate pointed posteriorly and

projecting the peristome; sternal and episternal plates densely tuberculated, as are parts of the outer plates in A I and A V; periproct on the vertical posterior end. Fascioles: Peripetalous fasciole broad, passing very closely around the distal ends of the petals; anteriorly

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

 271

C

B

D

E

F

G

H

Fig. 3.208: (A, B) Plesiozonus: Labral plate and adjacent ambulacral plates: (A) P. diomedeae. (B) P. hirsutus. Scale bars are 10 mm. After David & De Ridder (1989). (C–F) Plesiozonus hirsutus. (C) Specimen with spines, length 109 mm, Siquijor, Philippines. Aboral side: the spines are short and straight. (D–F) Specimen without spines; test length 124 mm, Philippines. (D) Aboral side: the petals are long with the interporiferous zones being broader than the pore zones. (E) Oral side: the labrum overarches the peristome. The large periproct is positioned on the oral side. (F): Side view. (G,H) Plesiozonus tenuis. (G) Distal part of the right posterior petal with the occluded plates (shaded) alternating with nonoccluded plates. Scale bar is 5 mm. (H) Labral plate (shaded) with adjacent ambulacral plates. The labrum is slightly to moderately overarching the peristome. Scale bar is 10 mm. Redrawn after David & De Ridder (1989).

272 

A

 3.3 Superorder Atelostomata von Zittel, 1879

C

B

D E

Fig. 3.209: (A, B) Differences of Faorina and Pericosmus in oral plating: In Faorina the plate in IA 4 is connected to a pair of plates as in Pericosmus, but there follows only a single plate in IA 1. The labral plate (lab) is elongated and pointed toward the sternal plates. In Pericosmus the interambulacral plates next to the peristome in IA 1 and IA 4 are followed by a pair of plates. The labral plate (lab) is stout. After A.B. Smith (2005). (C–E) Faorina chinensis. Length of test 73 mm; East India. ZMH. The specimen has been attacked by predatory snails, there are three holes and the traces of several attempts. (C) Aboral side: the peripetalous fasciole is well-developed. (D) Oral side: the plastron is densely tuberculated, as are parts of the outer plates in the oral ambulacra A I and A V. (E) Profile: only a deep anal curve below the periproct remains of the marginal fasciole in adults.

with a double band; marginal fasciole often incomplete or lacking in adults, mostly present as a deep curve below the periproct. Spines: Short, thin and fragile; longer along the petals, at plastron with spoon-shaped ends.

Color: The spines and the test are dark purplish to almost black, which usually remains in bare tests. Distribution: This species is known from the coast of ­Southern China and Vietnam to the Sulu Sea, and from Burma and the Andamans in depths from 50 to 180 m.



Genus Pericosmus L. Agassiz, 1840 Test: Strong, highly vaulted with more or less distinct frontal notch; rounded margin. Apical system: Ethmolytic with three gonopores. Anterior ambulacrum: More or less sunken, pores differentiated and set with suckered podia. Paired petals: Short, narrow, parallel-sided, distally with occluded plates and closed; sunken along their whole length. Oral side: Flat; peristome sunken; periproct marginal on truncated posterior face; phyllodes short; labral plate projecting over the peristome; stout, extending to second ambulacral plate; more or less tuberculated; sternal plates long and symmetrical, fully tuberculated. Fascioles: Marginal and peripetalous fasciole present. Tuberculation: Small and uniform, abruptly becoming denser below the marginal fasciole at the sides. Spines: Fine and only a few millimeters long; aborally curved, those along the petals bent over the ambulacra; oral spines widened distally. Color: Test and spines are usually deep purple, more rarely white. The well cleaned test may be a light purple. The surface is glossy. Ten extant, rather similar species are included in the genus: Pericosmus akabanus Mortensen, 1939. Gulf of Aqaba and Arabian Red Sea. Pericosmus bidens Mortensen, 1950. Mauritius. Pericosmus cordatus Mortensen, 1950. Southern Japan. Pericosmus keiensis Mortensen, 1950. Malayan Archipelago and Philippines. Pericosmus macronesius Koehler, 1914. Indo-West Pacific. Pericosmus mauritianus Mortensen, 1950. Mauritius. Pericosmus melanostomus Mortensen, 1948. Malayan region to China Sea. Pericosmus oblongus Mortensen, 1950. Malayan region and Philippines. Pericosmus porphyrocardius McNamara, 1984. Northwest and northeast coast of Australia. Pericosmus tenuis Mortensen, 1950. Mauritius. Remarks: According to Mortensen the various species of Pericosmus are difficult to distinguish except by means of the pedicellariae. He described most of his new species on the base of three specimens or less, and/or of young individuals with a length between 10 and 55 mm. That is problematic, for the shape of the test and the development of the fascioles change considerably with age. Pericosmus akabanus Mortensen, 1939 Test: High, upper side variable: shortly flattened and anteriorly almost vertical to subconical with the vertex at the apical system or slightly behind; in outline ovoid with

3.3.2 Order Spatangoida L. Agassiz, 1840 

 273

distinct anterior notch; midline of posterior interambulacrum may be sunken adorally; length up to 74 mm. Anterior ambulacrum: Deeply depressed toward the margin, small tube feet, and minute tubercles. Paired petals: Relatively long and sunken; interporiferous zone approximately as broad as a pore zone; distal two or three plates occluded. Oral side: Peristome conspicuously sunken, periproct at the short, slightly concave, truncated posterior face. Fasciole: Peripetalous fasciole very thin and difficult to trace, sometimes totally lacking, sometimes with two or three transverse bands anteriorly, hardly visible: marginal fasciole more conspicuous, but may also be hard to detect anteriorly, where it usually bends abruptly downward to the edge of the test. Distribution: The species lives in the Arabian Red Sea, including the Gulf of Aqaba; 272 to 486 m (Fig. 3.210). Pericosmus bidens Mortensen, 1950 Test: Hemispherically domed, slightly truncated at the posterior side; in outline ovoid, fairly deep anterior notch; maximum length 55 mm. Anterior ambulacrum: Sunken toward the margin; small pore pairs vertically positioned. Paired petals: Short, narrow and depressed; the anterior being longer than the posterior; occluded plates at the end. Oral side: Flat, labrum overarching the peristome; sternal plates narrow, fully tuberculated, tubercles continue onto labral plate; periproct transverse on the concave posterior side. Fascioles: Peripetalous and marginal fascioles rather thin, disappearing anteriorly. Larger tubercles: Some in the anterior interambulacrum bordering ambulacrum III. Occurrence: Mauritius, Indian Ocean. No depth recorded. Pericosmus cordatus Mortensen, 1950 Test: Heart-shaped with a deep frontal notch; hemispherically domed, posterior end obliquely concave and distinctly notched; maximum test length 83 mm. Apical system: Central, ethmolytic, madreporite hardly extending beyond ocular plates. Anterior ambulacrum: Increasingly sunken toward the margin, pores small and associated with a peripodium. Paired petals: Narrow, short, shallowly sunken, anterior pair a little longer than the posterior one. Oral side: Flat, peristome obliquely positioned, phyllodes not well-developed; labrum slightly overarching the peristome, sparsely tuberculated, sternum broad, fully tuberculated. Periproct: At the posterior face on top of the anal notch.

274 

 3.3 Superorder Atelostomata von Zittel, 1879

B

A

C

D

F

E

G

H

I

Fig. 3.210: (A–E) Pericosmus akabanus. Test length 65 mm, Gulf of Aqaba, Red Sea. ZMUC ECH 67. (A) Aboral side: the petals are broad, long and distinctly sunken. (B) Oral side. (C) Side view. The test is very high. Obviously the echinoid died after being attacked by a boring snail. (D) Apical system: the ocular plates are conspicuously indented in the madreporite. The extension of the madreporite toward the posterior may be variable. Scale bar is 3 mm. (E) Labrum and adjacent ambulacral plates: the labral plate is stout reaching only to the second ambulacral plate. The lip overarches the peristome. Scale bar is 6 mm. After Mortensen (1951). (F–I) Pericosmus bidens. Test length 55 mm; Mauritius, Indian Ocean. ZMUC ECH 92. (F) Aboral side: the petals are short, narrow and sunken. (G) Oral side: the sternum is narrow and parallel-sided. (H) Side view. (I) Distal end of posterior left petal with occluded plates. Scale bar is 5 mm. After Mortensen (1951).

3.3.2 Order Spatangoida L. Agassiz, 1840 



Peripetalous fasciole: Thin, laterally sharply, but not deeply indented. Marginal fasciole: Running just above the margin with a bent in the anterior ambulacrum and below the periproct. Tuberculation: Very fine, spines short and uniform, 2 to 3 mm long. Color: Uniformly dark purple.

 275

Distribution: This species is known from the Malayan region over the Philippines to Japan at a depth of 80 to 250 m (Fig. 3.211). Pericosmus keiensis Mortensen, 1950 Main characteristics of the genus: test length may reach 65 mm. This species is characterized by its distinctly sunken

A

B

C

D

E

Fig. 3.211: Pericosmus cordatus. (A–C) Test length 76 mm, Goto Island, Southern Japan. ZMUC ECH 118. (A) Aboral side: the peripetalous fasciole is not much indented laterally. (B) Oral side: the sternum is broad. (C) Side view. (D) Specimen with spines; test length 63 mm, Philippines. Aboral side. The spines are short. (E) Labral plate and adjoining ambulacral plates. Scale bar is 10 mm. After Mortensen (1951).

276 

 3.3 Superorder Atelostomata von Zittel, 1879

anterior ambulacrum set with pores, which are encircled by a peripodium and associated to differentiated tube feet. The posterior end is high and vertical. The aboral posterior interambulacrum is raised, the midline conspicuously depressed and naked. Color: Brownish-purple. Distribution: This species is known from the Malayan region and the Philippines, at 80 to 250 m. Remarks: The convex oral side and the well-developed fascioles indicate that it lived burrowed in the sediment (Fig. 3.212). Pericosmus macronesius Koehler, 1914 Test: Thick-shelled, high domed; outline heart-shaped with distinct anterior notch; length up to 70 mm. Apical system: With three gonopores; slightly anterior. Anterior ambulacrum: Depressed, increasing in depth toward the margin. Paired petals: Short, narrow, and parallel-sided; sunken. Oral side: Flattened; peristome anterior and sunken; periproct on the short, truncated, slightly concave posterior side; projecting labrum; labral plate stout; sternal plates long. Fascioles: Peripetalous fasciole with deep indentations in the interambulacra; marginal fasciole passes around the margin, shallow bend below the periproct: Tuberculation: Short and uniform. Color: In life, the spines are purple, the bare test is grayishwhite. Distribution: Recorded from the Indo-West Pacific at depths of 80 to 500 m (Fig. 3.213). Pericosmus mauritianus Mortensen, 1950 The species is characterized by a short labrum, reaching only the first adjoining ambulacral plates. One specimen of 13 mm; Mauritius, 220 m. Pericosmus melanostomus Mortensen, 1948 The tube feet in the anterior ambulacrum are differentiated having a well-developed sucking disc. The peristome and the periproct are almost black, contrasting to the whitish test and spines. Three specimens between 9 and 18 mm, Malayan region over the Philippines to the South China Sea; bathymetric range 18 to 70 m.

Pericosmus oblongus Mortensen, 1950 Mortensen (1950) described a specimen of 10 mm length as the holotype, which has no frontal notch. The welldeveloped fascioles and the convex oral side are typical features of juveniles. Malayan region at a depth of approximately 200 m. Pericosmus porphyrocardius McNamara, 1984 Test: Relatively low; broadly cordate, wider than long, anterior notch deep and wide, posterior end transversely truncated; maximum length 106 mm. Apical system: Anterior, slightly sunken; three gonopores. Anterior ambulacrum: Adapically flush, increasingly depressed toward the notched margin, pore pairs minute. Paired petals: Shallow, narrow and straight, anterior pair longer than posterior pair; interporiferous zone very narrow. Oral side: Flat, peristome lunate and deeply sunken; periproct sunken, subcircular or transversely oval; labrum slightly overarching the peristome, sternum narrow: Fascioles: Peripetalous fasciole with two branches anteriorly, may be discontinuous on frontal ambulacrum; marginal fasciole slightly above the ambitus. Spines: Very short, maximum length of 5 mm in anterior ambulacrum on the aboral side. Distribution: The species is recorded off the northwest coast and along the northeast coast of Australia at depths of 140 to 420 m. Pericosmus tenuis Mortensen, 1950 Characteristics of the genus. Length of holotype 37 mm. Test: Ovoid, shallow frontal depression, aboral side regularly arched. As the gonopores are not yet developed, the species may reach a considerably larger size. Anterior ambulacrum: Flush adapically with distinct pore pairs, simple tube feet without sucking disc. Paired petals: Remarkably short, almost flush with the test, interporiferous zone as broad as a pore zone. Fascioles: Peripetalous fasciole thin, obliterated in anterior ambulacrum; marginal fasciole distinct throughout. Occurrence: Mauritius, Indian Ocean; depth 230 m. Remarks: This is a juvenile specimen without gonopores. The oral side is convex and the fascioles distinct except in the anterior ambulacrum.

▸ Fig. 3.212: Pericosmus keiensis. (A–C) Test length 45 mm, Kei Islands, Indonesia. ZMUC Holotype ECH 192. (A) Aboral side. The peripetalous fasciole crosses the anterior ambulacrum with two branches. (B) Oral side. (C) Side view: the marginal fasciole runs relatively high around the test with a sharp bend below the periproct. The posterior side is vertically truncated. (D) Specimen with spines. Test length 53 mm, Philippines. Photo courtesy of H. van Noordenburg. Side view: this white specimen is a rarity. The spines below the marginal fasciole are longer and more densely distributed (the posterior side is to the left). (E) Pericosmus keiensis set by a coral: the test of the dead echinoid laid half-burrowed in the sediment and was settled by a coral serving as hard ground. Photo courtesy of H. van Noordenburg. (F) Distal end of left posterior petal. Scale bar is 4 mm. (G) Apical system. Scale bar is 2 mm. After Mortensen (1951).

3.3.2 Order Spatangoida L. Agassiz, 1840 



B

A

C D

E

F

G

 277

278 

A

 3.3 Superorder Atelostomata von Zittel, 1879

C

B

D

F

E

G

Fig. 3.213: (A–D) Pericosmus macronesius. (A) Specimen with spines. Test length 72 mm; Port Louis, Mauritius. ZMUC. (A) Aboral side: the peripetalous fasciole is strongly indented between the petals. (B–D) Specimen without spines; test length 68 mm, Leyte, Philippines. (B) Aboral side. (C) Oral side. (D) Side view. (E) Pericosmus porphyrocardius. Test length 94.4 mm, off Port Hedland, Western Australia. (A) Aboral side. (B) Side view. (C) Posterior view. Sketches after the photos in McNamara (1984). (F,G) Pericosmus tenuis. Test length 37 mm, Mauritius. ZMUC Holotype ECH 320. The specimen has not yet developed gonopores. (F) Aboral side: the petals are short. (G) Side view.

3.3.2.15 Family Prenasteridae Lambert, 1905 The family differs from the Schizasteridae by the course of the marginal fasciole: It passes several plates below the distal ends of the anterior petals whereas in schizasterids the marginal fasciole runs immediately around the end of the petals.

The family consists of three extant genera: Agassizia Valenciennes in Agassiz & Desor, 1847. Tropical Americas. Two species Prenaster Chesher, 1968. Bahamas, West Atlantic. Monospecific.



Tripylus Philippi 1845. Antarctic to subantarctic waters, brood-protecting. Monospecific. Genus Agassizia Valenciennes in L. Agassiz & Deor, 1847 Two nearly related species are known, which differ mainly in the arrangement of the posterior petals. Agassizia excentrica A. Agassiz, 1869 Agassizia scrobiculata Valenciennes, 1846 It seems that the posterior petals are more reduced adapically in Agassizia scrobiculata than in A. excentrica. However, there is much variation in both species. The geographical range is a good distinction: A. scrobiculata is recorded from the west coast of tropical America whereas A. excentrica is from the Caribbean. Test: Almost globular, upper side strongly convex, underside less so, outline a little longer than wide, no anterior notch; length up to 55 mm. Apical system: Central, plates fused, madreporite expanding behind the posterior ocular plates; four gonopores; ethmolytic. Anterior ambulacrum: Almost flush with the test; pore pairs small, carrying specialized tube feet. Paired petals: Slightly depressed; anterior pair long, with only one series of pore pairs, the anterior row inconspicuous; posterior pair approximately one-third the length of anterior pair. Oral side: Large D-shaped peristome near the anterior margin; labral plate short and wide, slightly projecting; periproct on vertical, truncated posterior end; almost naked area below the periproct within the bend of the fasciole; two or three subanal penicillate podia at each side. Fascioles: Peripetalous fasciole closely surrounding posterior petals, indented behind the anterior petals, crossing the anterior ambulacra on the oral side; lateroanal branch separating behind the lower end of the anterior petals, curving below the periproct in a deep bend. Spines: Fine and densely coated, with more or less differentiated larger ones on oral and aboral sides. Biology: The genus seems to burrow to a depth of approximately 20 cm below the sea bottom (Fig. 3.214).

3.3.2 Order Spatangoida L. Agassiz, 1840 

 279

Distribution: The species occurs along the west coast of America from the Gulf of California, Mexico, to Peru and the Galapagos Islands. It lives from the ebb zone to a depth of approximately 20 m. Genus Prenaster Lambert, 1911 This genus includes one extant species, described on the basis of a single specimen.

Agassizia excentrica A. Agassiz, 1869 The anterior series of pore pairs in the posterior petals is less developed than the posterior row. Distribution: The species is recorded from Florida to Barbados in a depth of 45 to 900 m.

Prenaster enodatus Chesher, 1968 Test: Fragile, in outline ovoid without anterior notch, tapering toward the posterior; upper and lower sides almost parallel, steeply rounded anteriorly, sharply truncated at posterior end. Apical system: Very anterior, ethmolytic with four small gonopores; genital plates small, not fused; ocular plates relatively large, deeply recessed into the test; pores of about the same size as gonopores: Anterior ambulacrum: Flush with the test on aboral side, narrow and with tiny, slit-like single pores. Paired ambulacra: Narrow and flush; subpetaloid with small, obliquely positioned pore pairs; rows of pore pairs parallel with narrow interporiferous zone; distally open; anterior pair divergent at almost 180°, distally slightly flexed; anterior row of pore pairs reduced; posterior pair also narrow and faintly flexed; longer than anterior pair. Oral side: Labrum short and broad, extending to first adjacent ambulacral plates; sternal plates long, broad and symmetrical; periproct very large, sunken, occupying most of the posterior face. Fascioles: Peripetalous branch very narrow, not coalescing the marginal fasciole disappearing behind the anterior petals, not indented laterally; marginal fasciole complete with lateroanal branch, crossing ambulacra II and IV far below the distal ends of the petals, passing anteriorly below the ambitus. Tuberculation: Uniform, slightly enlarged along the anterior ambulacrum and on the plastron being spatulate; spines short and fine, covering also the oral and aboral ambulacra. Color: In the living specimen light brown with the tube feet pigmented in a darker red-brown, the cleaned test being white. Occurrence: Only recorded from the Bahamas at a depth of 366 m. Biology: It is likely the species lives buried in mud or fine sand as other spatangoids (Chesher 1968) (Fig. 3.215).

Agassizia scrobiculata Valenciennes, 1846 Both series of pore pairs in the posterior petals are equally well-developed.

Genus Tripylus Philippi, 1845 This genus is monospecific. It is characterized by having a more or less complete marginal fasciole with lateroanal

280 

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 3.3 Superorder Atelostomata von Zittel, 1879

B

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D

G E

F H

Fig. 3.214: (A–D) Agassizia excentrica. Test length 31 mm; Florida, USA. (A) Aboral side: the anterior series of pore pairs are almost invisible in the anterior paired petals, in the posterior petals they are poorly developed. (B) Oral side: the plastron is broad. (C) Side view: the test is almost globular. (D) Posterior side: the lateroanal fasciole forms a deep bend below the periproct, the area within this bend is naked. (E–H) Agassizia scrobiculata. Test length 31 mm, Isla el Requeson, Mexico. Photo by Bas van der Steld. (E) Aboral side. (F–H) Specimen with spines. Test length 31 mm; Bahia Concepcion, California, USA. Photo courtesy of H. v. Noordenburg. (F) Aboral side: larger spines cover the sunken petals and the anterior ambulacrum. (G) Oral side: the anterior part of the peripetalous fasciole passes very low and is visible on the underside. The spines of the plastron are spatulate. Left anterior petal A IV: (H) Only the posterior column is well-developed, the anterior series is inconspicuous. Scale bar is 3 mm. After Mortensen (1951).

branch. The peripetalous branch, which mostly coalesces with the former behind the anterior petals, may continue as a separated branch toward the anterior side. The fasciole passes two or three plates beyond the distal end of the anterior petals. The development is nonpelagic and brood-protecting. Tripylus excavatus Philippi, 1845 Test: Subcircular, moderately raised aborally; shallow, but distinct anterior notch; length may reach 50 mm. Apical system: Anterior to subcentral; ethmolytic with three gonopores.

Paired petals: Narrow, deeply sunken in the females forming marsupia, less depressed in males; plate pattern of oral side variable. Oral side: Labrum short, extending to first adjoining ambulacral plates; sternal plates very broad, laterally strongly bowed; peristome kidney-shaped, facing forward, periproct on truncated posterior face. Fascioles: Distinct and complete; anterior branch distant from the end of the anterior petals; Peripetalous branch indented laterally, joining the marginal fasciole behind the anterior petals. Color: Test and spines in living specimens are pinkish-violet.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

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C

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Fig. 3.215: (A–C) Prenaster enodatus. Test length 60 mm, Bahamas. Sketches after photos in Chesher (1968). (A) Side view: the test is box-shaped. (B) Posterior side: the periproct is very large. (C) Aboral side: the paired ambulacra are subpetaloid, the parallel rows of pore pairs being very narrow. Photo courtesy of Doris J. Pawson. (D–I) Tripylus excavatus, female. Test length 37 mm; Tierra del Fuego, Argentina. ZMH. (D) Aboral side. (E–G) Male specimen. Test length 31 mm; Magellan Strait, Chile. ZMUC. (E) Aboral side: the fascioles are broad and conspicuous. Test length 35 mm. (F) Oral side. (G) Side view. (H, I) Drawing of male specimen. (H) Aboral side. (I) Oral plate pattern. The sternum is broad, the sides being strongly bowed. Scale bar is 10 mm. Modified after Madon-Senez (1998).

Distribution: T. excavatus is recorded along the coast of Fireland and in the South Georgia Islands. Depth range: from the littoral to 113 m. Remarks: There is a great deal of variation in the course of the fasciolar bands (Fig. 3.216).

3.3.2.16 Family Schizasteridae Lambert, 1905 The two families differ in the position of the marginal fasciole: In the schizasterids it passes immediately below the end of the anterior petals, in the prenasterids it is more distant, running two or more ambulacral plates below the end of the petals.

282 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

E

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H

Fig. 3.216: (A–D) Fascioles of schizasterids: (A) The marginal fasciole (green band) runs around the test and evolved first. The periproct (pp) is always positioned above the fasciole. (B,C) The peripetalous fasciole (orange band) crosses the aboral side like a bow evolving later in ontogeny and the periproct always below the fasciole. Both are coalesced anteriorly. (D) The remaining fasciole appears to be a single one, but actually it is composed of the peripetalous bow and the anterior branch of the marginal fasciole. Modified after Markov & Soloviev (2001). (E–H) Abatus cavernosus. Juvenile specimens of 5.3 and 5.5 mm length. From A. Agassiz (1904). (E) Aboral side with spines and podia. (F) Posterior side. (G) Side view. (H) Aboral side with plate pattern. The marginal fasciole is still completely developed, passing behind the periproct. The peripetalous bow runs between the apical system and the periproct over the aboral side. Later in ontogeny, the lateroanal branch of the marginal fasciole disappears.

Smith (2005) recently restricted the family Schizasteridae to 11 genera, based on morphological features such as plate architecture. In The Echinoid Directory, he separated several genera into still unnamed groups, including within others the Antarctic brood-protecting genera Amphipneustes, Brachysternaster and Genicopatagus. Tripylus, a brooder as well, was placed into the family Prenasteridae. The six broodprotecting genera are closely related, despite their morphological differences, and they share a similar evolutionary scenario. Féral et al. (1994) showed by RNA analyses that Abatus and the four above named genera are monophyletic, displaying nonpelagic development and brood-protecting ability. Eleven genera are included in the family Schizasteridae: Abatus Troschel, 1851. Antarctic and Subantarctic. Eleven species, brood-protecting.

Aceste Thomson, 1877. Worldwide. Three species. Brisaster Gray, 1855. Worldwide. Eight species. Diploporaster Mortensen, 1950. Indian Ocean. Two species. Hypselaster H.L. Clark, 1917. Worldwide. Nine species. Moira A. Agassiz, 1872. Worldwide. Five species. Moiropsis A. Agassiz, 1881. Caribbean. Monospecific. Ova Gray, 1825. Worldwide. Five species. Protenaster Pomel, 1883. Southern Australia. Monospecific. Schizaster L. Agassiz, 1836. Six extant species. Tripylaster Mortensen, 1907. Subantarctic waters. Monospecific, not brood-protecting. Many of the genera are very similar: Apical system: With two, three, or four gonopores. Anterior ambulacrum: More or less deeply sunken, pore pairs associated with specialized, suckered tube feet.



Paired petals: Sunken, posterior pair distinctly shorter than anterior pair. Periproct: High on posterior truncate side. Fascioles: Marginal fasciole coalesced anteriorly with the peripetalous fasciole; lateroanal (branch) fasciole present, rudimentary or entirely lacking in adults. The differentiation of the genera in the family Schizasteridae was and is still the subject of discussions. Usually, the schizasterids are distinguished by the number of gonopores and by the development of the lateroanal fasciole. However, both features are not stable in the genera or even on the species level: The lateroanal fasciole may be developed to different degrees, or it becomes completely lost in ontogeny. Additionally, some species (e.g., Schizaster compactus) show variably two, three, or four gonopores. Literature: Mortensen 1910; Koehler 1926; Fischer 1966; Chesher 1972; McNamara & Philipp 1980; Pearse & ­McClintock 1990; Lockhart et al. 1994; McNamara 1995; Rowe & Gates 1995; Madon-Senez 1998; 2002; David et al. 2000 (CDRom), 2005 (with a comprehensive bibliography); Pearse et al. 2008 (comprehensive bibliography). All Antarctic members are brood-protecting, which means that the young do not run through a pelagic larval stage with following metamorphosis, but develop from large yolk-rich eggs directly into small juvenile. Their development is nonpelagic, being sheltered in brood pouches on the female.

Why are so many brooding species concentrated in Antarctic and subantarctic waters? It is not the cold and harsh environment because in the Arctic, which has equal conditions, only a single species with nonpelagic development is known. Instead, the fossil records show that the nonpelagic development of the Antarctic echinoids arose long before the cooling of the Southern Ocean began. A fossil echinoid, assigned to Abatus, was found dating from the Late Eocene of Seymour Island (approximately 36 million years ago). Careful data analyses in the last 20 years revealed that the highest proportion of Antarctic species with nonpelagic development is found in the region of the Scotia Arc, not in the subantarctic islands or around the Antarctic continent. The reason for this phenomenon is the powerful Antarctic Circumpolar Current (ACC) that has been flowing through the Drake Passage for more than 30 million years. The individuals of species with nonpelagic development are rafted infrequently to other downstream habitats and become established to form isolated populations and then new species. Moreover, in the Scotia Arc, new habitats are continually formed by tectonic activities,

3.3.2 Order Spatangoida L. Agassiz, 1840 

 283

and the populations are repeatedly fragmented (ACC Hypothesis; Pearse et al. 2008). In that way, the brooding genera are divided into numerous, often very similar species, spreading over the islands of the Scotia Arc to the Bouvet and Kerguelen Islands, down the east side of the Antarctic Peninsula and into the eastern part of the Weddell Sea. And some of them found their way around the Antarctic continent to the Ross Sea or even into the Bellingshausen Sea. This extraordinary evolutionary development has happened in only two clades of echinoids, in members of the Cidaridae, where the juveniles are housed around the peristome, and in the paleopneustids. Also, in other groups of marine organisms, e.g., the crustaceans, few, but species-rich clades with nonpelagic development are recorded around Antarctica with most of the species in the Scotia Arc (Pearse et al. 2008). Genus Abatus Troschel, 1851 Test: More or less broadly oval, slight anterior notch; aboral side more or less domed. Apical system: Subcentral, ethmolytic, three or two gonopores; in females larger than in males. Anterior ambulacrum: Deeply sunken from the apical system toward the indented margin, a single row of pore pairs to each column, associated with specialized tube feet. Paired petals: Deeply sunken, in females very deeply depressed forming marsupia, anterior petals longer than posterior pair. Oral side: More or less convex, peristome large, kidneyshaped, facing forward; covered by labrum; periproct elongated, on vertically truncated posterior side; anal tube feet usually present; labral plate short and wide; in most species, reaching to first adjacent ambulacral plate, sternal plates in broad contact with labrum, symmetrical, fully tuberculated. Fascioles: Margino-peripetalous fasciole very distinct, lateroanal branch present in juveniles and lost in adults. Tuberculation: Dense and more or less uniform, spines bordering the sunken ambulacra longer and bending over pore zones; plastronal spines often spatulate. Color: Test and spines are brownish. The genus contains 11 species and is exclusively distributed in Antarctic and Subantarctic waters. Abatus agassizii Pfeffer, 1889 Abatus beatriceae Larrain, 1985 Abatus bidens Mortensen, 1910 Abatus cavernosus (Philippi, 1845) Abatus cordatus Verrill, 1876. Endemic to the Kerguelen Islands, southern Indian Ocean.

284 

 3.3 Superorder Atelostomata von Zittel, 1879

Abatus curvidens Mortensen, 1936 Abatus koehleri Thiéry, 1909 Abatus ingens Koehler, 1926 Abatus philippii Loven, 1871 Abatus shackletoni Koehler, 1911 Abatus (Pseudabatus) nimrodi Koehler, 1911 It is often very difficult to distinguish the species because they are rather similar and there is much variation in appearance (Fig. 3.217).

A

F

Abatus agassizii Pfeffer, 1889 Main characteristics of genus: length may reach 65 mm. Test: In outline circular, no anterior notch; posterior side vertically truncated. Apical system: With three gonopores. Paired petals: Of equal length. Oral side: Labral plate short, extending to first adjoining ambulacral plates, sternal plates very broad, forming an oval plastron.

B

C

D

E

G

Fig. 3.217: (A–C) Abatus agassizii, female. Test length 36 mm, South Georgia. ZMUC. Male specimen; scale bar is 10 mm. Redrawn after Madon-Senez (1998). (A) Aboral side. (B) Plate pattern of aboral side. (C) Peristome, labrum and sternum. (D,E) Abatus beatriceae, male. Scale bar is 10 mm. Redrawn after Madon-Senez (1998). (D) Aboral side. (E) Peristome, labrum, and sternum: the sternal plates are long and narrow. (F,G) Abatus bidens, female. Test length 26 mm, South Georgia. ZMUC. Rather young specimen. (F) Aboral side. (G) Side view.



Margino-peripetalous fasciole: Nearly pentagonal, passing well above the margin. Distribution: Recorded from the northern part of the Antarctic Peninsula to the Falkland Islands and South Georgia, and in the Eastern Weddell Sea; bathymetric range from the littoral to 592 m, but mostly living above 200 m. Abatus beatriceae Larrain, 1985 Main characteristics of the genus: length of medium size. Test: Elongate, anterior side well rounded, tapering conspicuously toward the narrow posterior end. Posterior petals: Of same length or slightly longer than anterior pair, but not extending to the fasciole. Oral side: Labrum long, reaching the third or fourth adjoining ambulacral plates, sternum long and narrow. Fasciole: Margino-peripetalous fasciole marginal anteriorly, not indented laterally; lateroanal branch present only in very young specimens. Occurrence: Only a few specimens were collected in the Drake Passage; 3,514 to 3,806 m. Abatus bidens Mortensen, 1910 Main characteristics of the genus: length reaches approximately 40 mm. Test: In outline ovoid with distinct anterior notch. Paired petals: Of equal length, anterior pair at an angle of at least 120°. Oral side: Labrum short, extending to first adjacent ambulacral plates, protruding over peristome; periproct at vertical posterior side, not visible from above. Fasciole: Anterior part of the margino-peripetalous fasciole low on the test and narrow. Occurrence: South Georgia; depth between 64 and 270 m. Remarks: In Kroh & Mooi (2017), it is synonymous with A. cavernosus. Abatus cavernosus (Philippi, 1845) Main characteristics of the genus: length may reach 50 mm. Test: Oval with distinct anterior notch, posterior side vertically truncated; shape of test very variable. Paired petals: Anterior pair slightly longer than posterior pair. Oral side: Labrum short, extending to the first adjacent ambulacral plates, largely covering the peristome; sternum long and broad. Fascioles: Anterior branch of fasciole passing close to the ambitus, visible from above; margino-peripetalous fasciole broad, indented laterally. Distribution: Recorded widely around Antarctica from the Magellanic region over South Shetland and South Georgia to the Bouvet Island, and off Mac Robertson

3.3.2 Order Spatangoida L. Agassiz, 1840 

 285

Coast, Adélie Coast, and King George V Coast. Bathymetric range: 10 to 3,806 m, most commonly between 500 and 1,000 m (Fig. 3.218). Abatus cordatus Verrill, 1876 Main characteristics of the genus: maximum length approximately 45 mm. Test: Oval with shallow anterior and posterior notch. Oral side: Labrum partly covering the peristome, extending to the first or second adjoining ambulacral plates; no anal tube feet developed (distinct from all other Abatus species). Fasciole: Margino-peripetalous fasciole indented laterally, very broad in the paired ambulacra, narrower on the interambulacra. Distribution: This species is restricted to the Kerguelen Island and Heard Island at depths of 0 to 585 m, but is most common in very shallow water of 0 to 3 m. Biology: The biology of reproduction of the brooding (marsupiate) schizasterids is still poorly known. Only A.  cordatus Verrill, 1876, endemic to the Kerguelen Islands in the South Indian Ocean and living abundantly in shallow waters between 1.50 and 200 m, is very well studied. At these subantarctic islands, it is possible to obtain data throughout the year and the sea urchins are in “direct reach” of the scientists. According to locality and adapted to the seasonally changing resources of nutrition, A. cordatus reproduces annually, with the development from fertilized egg to long-spined juvenile ready to live independently taking 250 days. The offspring in the pouches are always of about the same age. After releasing the young, the female rests for approximately 100 days (Schatt & Féral 1996). In contrast to this cycle, the paleopneustids of high-Antarctic waters are likely to reproduce throughout the year (Fig. 3.219). Abatus curvidens Mortensen 1936 Main characteristics of the genus: maximum length 51 mm. Test: Regularly oval with faint anterior sulcus. Oral side: Labrum prominent, extending to the second or third adjoining ambulacral plates. Fasciole: Margino-peripetalous fasciole thin (relative to other species of Abatus), anteriorly passing around the ambitus, slightly indented in the lateral interambulacra. Distribution: This species is recorded only from the northern end of the Antarctic Peninsula, in the Palmer Archipelago, South Shetland Islands, and Graham Land; bathymetric range 20 to 769 m. Abatus ingens Koehler 1926 Main characteristics of the genus: length usually reaches 60 to 70 mm, maximum length recorded 88 mm.

286 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

C

D G

E

F

Fig. 3.218: Abatus cavernosus. (A) Female, test length 37 mm; Weddell Sea. AWI. Aboral side: the brood pouches (marsupia) are very deep. (B–D) Male specimen. Test length 38 mm, South Georgia. ZMUC. (B) Aboral side: the ambulacra are depressed, but not as deep as in the females. (C) Oral side. (D) Side view. (E) Aboral side with the marginal fasciole. The peripetalous fasciole joins the marginal fasciole just behind ambulacrum IV. The marginal fasciole does not continue toward the posterior forming a lateroanal branch. (F) Peristome, labrum, and sternum: the sternal plates are broad. Scale bar is 10 mm. Modified after Madon-Senez (1998).

Test: Rounded anteriorly without depression, conspicuously attenuated toward the posterior; periproct visible in oral sight. Oral side: Labrum usually extending to the fourth adjoining ambulacral plates; overarching the peristome.

Fasciole: Fasciolar band sinuous, indenting in all interambulacra. Color: In life, very dark, almost black. Distribution: This species is recorded from Prydz Bay to the east coast of the Ross Sea, at a depth of 20 to 271 m.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

B

C

D

E

 287

F

Fig. 3.219: Abatus cordatus. (A–C) Female, test length 32 mm, Kerguelen Islands. ZMUC. (A) Aboral side: there are four deeply depressed brood pouches. (B) Oral side. (C) Side view. (D) Female specimen with spines; test length 45 mm. Kerguelen Islands. ZMH. (E) Aboral side of test. The anterior notch is shallow. (F) Peristome, labrum and sternum: the sternal plates are broad. Scale bar is 10 mm. Modified after Madon-Senez (1998).

Biology: This species is observed ploughing through the sediment with only the lowest part of the test buried (David et al. 2005) (Fig. 3.220). Abatus koehleri Thiéry, 1909 Synonymous to A. elongatus Koehler, 1908; see The World Echinoidea Database. WoRMS.

Main characteristics of the genus: length may reach 65 mm. Test: Ovoid, anterior side rounded without depression, posterior part narrowing to the truncated posterior face. Anterior ambulacrum: Flush. Anterior petals: Strongly diverging at an angle of approximately 160°.

288 

 3.3 Superorder Atelostomata von Zittel, 1879

B

A

C

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E F

H

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I

K

Fig. 3.220: (A–D) Abatus ingens, female. Test length 87 mm, Ross Sea, Antarctica. Male specimen. ZMUC. (A) View into the internal test from the oral side: the four marsupia are strongly developed. (B) Aboral side. (C) Aboral side: there is no anterior notch. The fasciole is strongly sinuous. (D) Peristome, labrum, and sternum: the labrum extends to the fourth adjoining ambulacral plates, the sternal plates being relatively straight. Scale bar is 10 mm. Modified after Madon-Senez (1998). (E, F) Abatus koehleri female. Test length 39 mm. McMurdo Sound, Ross Sea, Antarctica. ZMUC; as A. elongatus. (E) Aboral side. (F) Side view. (G–K) Abatus philippii. (G) Male specimen, length 28 mm. (H) Female, test length 27 mm; off Argentina. ZMUC. In the female, only the anterior ambulacra are deeply depressed forming brood pouches. (I) Specimen with spines. Test length of male 41 mm; Weddell Sea, depth 175 m; ZMH. (J, K) Male specimen. Plate pattern; scale bar is 10 mm. Modified after Madon-Senez (1998). (J) Aboral side: there is a faint anterior notch. (K) Peristome, labrum, and sternum: the labrum extends to the first adjoining ambulacral plates, the sternal plates are broad and bowed.



3.3.2 Order Spatangoida L. Agassiz, 1840 

 289

Oral side: Labrum short, extending to the first or second adjoining ambulacral plates; periproct on posterior truncated face, bordered by two characteristic tufts of spines. Fasciole: Fasciolar band very thin, three to five miliaries in width. Distribution: Recorded from the South Orkney Island and in McMurdo Sound, Ross Sea, Antarctica. Bathymetric range 11 to 804 m.

Oral side: Labrum large, reaching to second or fourth adjoining ambulacral plates, more or less protruding over the peristome; sternum triangular with straight sides. Color: Dark brown or black. Distribution: Recorded abundantly along the Antarctic coast from 60°E (Prydz Bay) to 180°E (Ross Sea), at a depth of 13 to 732 m.

Abatus philippii Lovén, 1871 Main characteristics of the genus: length may reach 40 mm. This species is unique in the genus Abatus in having only the anterior pair of petals modified to marsupia. The posterior ambulacra are not deepened in the females. Test: High, posterior interambulacrum slightly raised, vertically truncated at the rear; in outline oval to hexagonal with a faint frontal notch. Paired petals: Posterior pair distinctly shorter than anterior pair. Labrum: Short, only slightly covering the peristome. Fasciole: Fasciolar band broad, posteriorly well above the margin. Distribution: Recorded at South Shetland Islands and along the Patagonian coast to the mouth of the La Plata, also circumantarctic off South Georgia Islands, Sabrina Coast, and in the Ross Sea. Bathymetric range 27 to 804 m.

Genus Aceste Thomson, 1877 The genus is extraordinary in having an extremely broad, long, and deeply sunken anterior ambulacrum. The large pores are associated with very conspicuous podia, which have a flower-like distal disc. The genus consists of three species: Aceste bellidifera Thomson, 1877. Atlantic from the USA over the Caribbean to the Subantarctic. Aceste ovata A. Agassiz & H.L. Clark, 1907. Indo-Pacific to off Peru. Aceste weberi Koehler, 1914. Timor Sea, Indonesia. Mortensen (1950) has doubts about the validity of these three species because the range of variation in general shape and form of the anterior ambulacrum is large. He considered that at least the two Indo-Pacific species may be conspecific (Fig. 3.222).

Abatus shackletoni Koehler, 1911 Main characteristics of the genus: length may reach 67 mm, commonly approximately 40 mm. Test: More or less ovoid, frontal side rectilinear or with only a very slight depression; posterior side truncated. Labrum: Long and stout, extending to the third or fourth adjoining ambulacral plates; not much projecting over the peristome. Fasciole: Of moderate width, passing close to the entire ambitus, no indentations laterally. Color: Brown to grayish purple. Distribution: This species is recorded circumpolar from the Antarctic Peninsula to Enderby Land, the Adélie Coast, and McMurdo Sound; depth range from the littoral to 631 m (Fig. 3.221). Abatus (Pseudabatus) nimrodi Koehler, 1911 Main characteristics of the genus: length may reach 60 mm, mostly 30 to 40 mm. This species is well distinguished from the other species of Abatus: In A. (P.) nimrodi, the depressed part of the petals begins distant from the apical system; in the females, seven to nine ambulacra plates. The petals are relatively narrow. Test: Regularly ovoid with rounded anterior side, apical area flattened; posterior end shortly truncated.

Aceste bellidifera Thomson, 1877 Test: Elongated cordate, anterior notch very deep; wedgeshaped in profile with the vertex posteriorly; length 20 to 30 mm. Apical system: Very posterior, just within the posterior corner of the fasciole; ethmolytic; two gonopores; plates more or less fused. Anterior ambulacrum: Deeply sunken, arranged uniserially; with large podia specialized for funnel-building, discs with large rosettes. Paired ambulacra: Nonpetaloid and flush with the test; pore pairs more or less rudimentary; posterior pair extremely short. Oral side: Convex; peristome vertical at anterior margin, opening facing toward the front into the anterior notch, covered by the labrum when seen from below; periproct below the low posterior margin; labral plate elongate and narrow, reaching the first adjoining ambulacral plates, broadly contacting the sternum; sternal plates broad, fully tuberculated, extending to fifth adjacent ambulacral plates. Fasciole: Margino-peripetalous fasciole well-developed, lateroanal branch lacking. Spines: Aborally uniform; much longer along the large anterior ambulacrum and distally widened; plastronal spines spoon-shaped, other oral spines stout and densely

290 

 3.3 Superorder Atelostomata von Zittel, 1879

A

C

B

D

E

F

G

H

Fig. 3.221: (A, B) Abatus shackletoni, male. (A) Aboral side: the anterior margin is hardly depressed. (B) Peristome, labrum, and sternum: the sternal plates are straight and on the whole triangular. Scale bar is 10 mm. Modified after Madon-Senez (1998). (C–H) Abatus (Pseudabatus) nimrodi, male. Test length 65 mm, Ross Sea, Antarctica. ZMUC. (C) Aboral side: In males, the petals are narrow and hardly depressed. (D) Oral side: the sternum is triangular, the sides straight or even slightly concave. (E) Side view. (F, G) Male specimen; scale bar is 10 mm. Modified after Madon-Senez (1998). (F) Aboral side: there is no anterior notch. (G) Peristome, labrum, and sternum: the labrum extends to the fourth adjoining ambulacral plates, the sternal plates being straight and on the whole triangular. (H) Female specimen. Aboral side: the marsupia are short and distant from the apical system. Grayish line: Supposed course of fasciole. Scale bar is 10 mm. Sketch after photo in David et al. (2005).



distributed; those encircling the periproct long, curved, and pointed. Distribution: Atlantic from the northeast coast of the USA over the Canaries to subantarctic waters in the south; 1,135 to 3,475 m.

3.3.2 Order Spatangoida L. Agassiz, 1840 

 291

Brisaster antarcticus (Döderlein, 1906). Antarctic and ­Subantarctic. Brisaster capensis (Studer, 1880). Southernmost South Africa. Brisaster fragilis (Düben & Koren, 1844). North Atlantic from North Norway to Florida. Aceste ovata A. Agassiz & H.L. Clark, 1907 Brisaster kerguelensis H.L. Clark, 1917. Endemic at the KerThis species strongly resembles A. bellidifera. Mortensen guelen Islands. was unable to find a reliable difference because the speci- Brisaster latifrons (A. Agassiz, 1898). Southeast Alaska to mens from the Atlantic as well as those from the Indothe Gulf of California. Pacific show a wide range of variation in general shape, in Brisaster moseleyi (A. Agassiz, 1881). Off the west to the the form of the anterior ambulacrum and in the spines. east coast of Southern South America. Distribution: A, ovata is widely spread in the Indo-­ Brisaster owstoni Mortensen, 1950. Northern Japan from Pacific: from East Africa over India, Japan and Hawaii to Sagami Bay to Hokkaido. the Coast of Peru, South America. Bathymetric range 435 Brisaster tasmanicus McKnight, 1974. New Zealand. to 4,755 m. Brisaster townsendi (A. Agassiz, 1898) North Pacific from Southeast Alaska to Panama. Aceste weberi Koehler, 1914 Literature: Döderlein 1906; McKnight 1974; Shigei 1986; Very much like A. bellidifera and A. ovata: length may Hood & Mooi 1998; David et al. 2005. reach approximately 25 mm. Occurrence: Timor Sea, Indonesia; depth 918 m. Brisaster antarcticus (Döderlein, 1906) Hood & Mooi (1998) felt B. antarcticus synonymous with Genus Brisaster Gray, 1855 B. kerguelensis H.L. Clark, 1917 and B. tasmanicus McKnight, Brisaster is characterized by having a deep anterior groove, 1974. a low test, and three gonopores. It is distinguished from Main characteristics of the genus: the holotype is only a Schizaster, Ova, and especially from the rather similar and fragment. closely related Tripylaster by the typical globiferous pedi- Paired petals: Rather narrow, anterior column of anterior cellariae, which show only a single terminal tooth. The petals with rudimentary pore pairs; anterior pair slightly others have more than one. flexed outward; posterior pair short and strongly divergent. Four species of the worldwide distributed genus Fasciole: Lateroanal fasciole restricted to a small part Brisaster and the genus Tripylaster are restricted to the below the periproct or wholly lacking. Subantarctic. They are not brooders, being phylogeneti- Distribution: This species lives in Subantarctic and Antcally the sister group to the species with nonpelagic devel- arctic waters around the Bouvet Islands, the Crozet, the opment (Féral et al. 1994). Heard and the Kerguelen Islands, South Atlantic and Test: Subcircular to oval, deep anterior notch; in profile South Indic. anteriorly rather low, posterior end truncated. Apical system: Ethmolytic with three gonopores, posi- Brisaster capensis (Studer, 1880) tioned more or less posteriorly. Main characteristics of the genus: test length may reach Anterior ambulacrum: Depressed from apical disc; pore more than 50 mm. pairs well-developed, uniserial and set with penicillate Test: Subcircular with narrow, deep anterior notch. tube feet specialized for funnel-building. Apical system: Relatively distant from the posterior margin. Paired petals: Sunken, anterior pair more than double Anterior ambulacrum: Also rather narrow. the length of the posterior pair. Paired petals: Straight, not flexed distally. Oral side: Peristome anterior and forward-facing into Fascioles: Complete marginal and peripetalous fasciole anterior furrow; covered by the labrum; periproct on the well-developed. posterior side; labral plate short and mushroom-shaped, Distribution: Known only from off South Africa, mainly extending to first adjacent ambulacral plates; sternal restricted to the Cape Peninsula; 215 to 350 m. plates symmetrical. Fascioles: Margino-peripetalous fasciole present, passing Brisaster fragilis (Düben & Koren, 1844) around the distal ends of the petals; lateroanal branch Main characteristics of the genus: length may reach 90 mm present in some species, in others only present in juveniles. in the north, but commonly 60 mm.

292 

 3.3 Superorder Atelostomata von Zittel, 1879

B

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H



Test: Fragile; wedge-shaped, rising gently toward the highest point, a keel immediately behind the apical disc. Paired petals: Anterior pair three times as long as posterior pair. Fascioles: Peripetalous fasciole broad, lateroanal branch narrow, inconspicuous or lacking laterally; distinct curve below the periproct. Distribution: This species is known in the Northeast Atlantic from the Shetland Islands along the Norwegian Coast to Finmark, off Iceland, and in the Northwest Atlantic from the Davis Strait to Florida, USA, at depths of 40 to 1,300 m (Fig. 3.223). Brisaster kerguelensis H.L. Clark, 1917 Main characteristics of the genus: see the remarks in B. antarcticus. Test length of holotype reaches 55 mm. Occurrence: Kerguelen Islands, Indian Ocean; depth range 200 m. Brisaster latifrons (A. Agassiz, 1898) Main characteristics of the genus: maximum length of test 77 mm. There is much confusion about the differentiation between this species and B. townsendi, which has the same geographical range. Both show considerable variations. Usually, B. townsendi has longer and broader posterior petals, but specimens also occur, such as B. latifrons, with long posterior petals. Hood & Mooi (1998) emphasize that it is not the length of the petals but the width, which may rather help to distinguish the two species. In B. townsendi, the petals are usually broader. Lateroanal fasciole: None developed. Distribution: Northern Pacific from Alaska to the Gulf of California, deep sea (Fig 3.224). Brisaster moseleyi (A. Agassiz, 1881) Main characteristics of the genus: test length may reach 75 mm. Test: Aborally convex, vertex subcentrally; oral side also convex. Apical system: Relatively distant from the posterior margin. Paired petals: Anterior pair distally flexed outward, posterior pair straight.

3.3.2 Order Spatangoida L. Agassiz, 1840 

 293

Fascioles: Margino-peripetalous fasciole well-developed, lateroanal branch much reduced, at most only visible in the subanal area. Distribution: This species is known from off Chile to the Falkland Islands and off Patagonia. However, Hood & Mooi (1998) found that the specimens from the west coast of South America are distinct from those found off the east coast. Bathymetric range: 110 to 1,425 m. Brisaster owstoni Mortensen, 1950 Main characteristics of the genus: length may reach 47 mm. Typical for this species is the comparatively high test with the vertically truncated posterior end and the very short posterior petals. However, in larger specimens, the test may be relatively lower. The lateroanal fasciole is restricted to a bow below the periproct. Distribution: Japan from Sagami Bay to Northern Hokkaido, Okhotsk Sea, deep sea. Remarks: Shigei (1986) synonymized this species with B. latifrons. However, the morphometric analysis of Hood & Mooi (1998) supported the distinctiveness of both species, although they admit strong similarities (Fig. 3.225). Brisaster tasmanicus McKnight, 1974 Holotype 22 mm; 1,184 to 1,193 m deep. Southeast of New Zealand at 37 to 2,530 m. Hood & Mooi (1998) felt that B. antarcticus was synonymous with B. kerguelensis H.L. Clark 1917 and B. tasmanicus McKnight, 1974. Brisaster townsendi (A. Agassiz, 1898) This species is very much like B. latifrons, which partially overlaps in its distribution. Young specimens are impossible to distinguish, and even in adult specimens it may be difficult, as the range of variation is large. The posterior petals reach more than half the length of the anterior pair. The anterior ambulacrum may be narrower than in B. latifrons. Distribution: Southeast Alaska to the Gulf of Panama including the Gulf of California, at depths of 35 to 1,900 m (Fig. 3.226).

◂ Fig. 3.222: (A–D) Aceste bellidifera. Test length 16.8 mm; off Liberia, West Africa. Photo courtesy of C. Neumann, MfN Berlin. (A) Aboral side: the fasciole is very conspicuous. The tube feet in the large, sunken anterior ambulacrum have a flower-like disc. (B) Oral side. The plastronal spines are spoon-shaped. (C) Drawing from Agassiz (1881). Test length 27 mm, off Canaries. (a) Aboral side, (b) oral side, (c) side view, (d) posterior side, (e) anterior side. (D) Apical system: there are only two gonopores. The plates are fused. Scale bar is 1 mm. From Mortensen (1950). (E–H) Aceste ovata. Test length 25 mm, off the coast of Kenya, East Africa, depth 4,940 m. ZMUC. (E) Aboral side: in living specimens, the deeply sunken anterior ambulacrum is covered by long spines. (F) Oral side: the plastronal spines are long, the distal ends being spoon-shaped. (H) Side view: most of the aboral side is occupied by the deeply sunken anterior ambulacrum. (G) Another specimen; test length 20 mm, East Africa. ZMUC. Aboral side.

294 

A

 3.3 Superorder Atelostomata von Zittel, 1879

B

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E

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G

Fig. 3.223: (A, B) Brisaster capensis. Test length 50 mm, Cape Peninsula, South Africa. ZMUC. (A) Aboral side: the paired petals are straight. (B) Side view: the lateroanal fasciole is well-developed. (C) Apical system of B. fragilis. Scale bar is 2 mm. After Lovén (1872). (D–G) B. fragilis. Test length 43 mm; Norway. (D) Aboral side: the anterior pair of petals is about three times as long as the posterior pair. (E) Oral side: the short labral plate projects over the mouth, which faces forward into the deep anterior groove. (F) Side view: the broad peripetalous fasciole coalesces with the distinct marginal fasciole, the lateroanal branch is thin. (G) Specimen with spines. Test length 39 mm; Northern Norway. Aboral side: long spines cover the sunken anterior ambulacrum with the funnel-building tube feet.

▸ Fig 3.224: (A–D) Brisaster kerguelensis. Test length 55 mm, Kerguelen Islands. ZMUC. (A) Aboral side: larger spines extend over the sunken anterior ambulacrum. (B) Oral side: the plastron is fully tuberculated. (C) Side view: the lateroanal branch is strongly reduced in adults. (D) Specimen with spines. Test length 50 mm, Kerguelen Islands. ZMUC. Aboral side. (E,F) Brisaster latifrons. Test length 40 mm, St. Cruz Island, CA, USA. ZMUC. (E) Aboral side: the posterior petals are short. (F) Side view: there is no lateroanal fasciole.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

B

C

D

E

F

 295

296 

A

 3.3 Superorder Atelostomata von Zittel, 1879

B

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F

Fig. 3.225: (A–C) Brisaster moseleyi. Test length 60 mm, off Chile. ZMUC. (A) Aboral side: the anterior paired petals are flexed outward, the posterior ones are straight. (B) Oral side. (C) Side view: the lateroanal fasciole is only visible on the posterior side. (D–F) Brisaster owstoni. Test length 23 mm, Tokyo Bay, Japan. (D) Aboral side: the posterior petals are very short and strongly divergent. (E) Oral side. (F) Side view: the test is relatively high. The height depends on the animals’ age; similarly, larger tests may be lower.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

 297

B

C

D

E

Fig. 3.226: (A–C) Brisaster townsendi. Test length 31 mm; Baja California, Mexico. (A) Aboral side: the posterior petals reach more than half the length of the anterior petals. (B) Oral side. (C) Side view: the underside is convex. (D, E) Specimen with spines. Test length 35 mm; Southern California, USA. (D) Aboral side: the spines are short and fine. (E) Oral side: the spination is short and scattered over the surface.

Genus Diploporaster Mortensen, 1950 The genus differs from Ova in having four gonopores. The pore pairs in the anterior ambulacrum are arranged biserially, two rows per column. Ova has only two gonopores and the anterior pore pairs form a single series per column.

There are two extant species: Diploporaster barbatus Mortensen, 1950. East Africa. Diploporaster savignyi (Fourtau, 1904). Red Sea. The two species are distinguished by shape: in Diploporaster savignyi, the test is wedge-shaped, the posterior end

298 

 3.3 Superorder Atelostomata von Zittel, 1879

being raised. The anterior ambulacrum is overarched by sharply keeled interambulacra. In Diploporaster barbatus, the posterior end is low, and the anterior interambulacra are raised, but not overarching the sunken ambulacrum III. Diploporaster barbatus Mortensen, 1950 Test: Slightly elongated with a deep frontal notch; aboral side slightly arched; posterior end truncated; length up to 59 mm. Apical system: Posterior, ethmolytic with four gonopores, the anterior pores may be smaller. Anterior ambulacrum: Broad, deeply sunken; set with pore pairs associated with specialized tube feet, arranged in two series. Paired petals: Sunken, posterior pair much shorter than anterior pair; diverging, anterior pair flexed outward distally. Oral side: Slightly convex, peristome overarched by the labrum; periproct small, on truncate posterior face; labral plate short and wide, broadly contacting sternal plates. Fascioles: Complete marginal fasciole, but lateroanal branch much thinner, peripetalous fasciole broad and deeply indented in the lateral interambulacra. Tuberculation: Dense and uniform, larger along the ambulacra. Spines: Larger ones covering the ambulacra; plastronal spines on oral side stronger and with distal “spoon”; there are two tufts of spines at the posterior end, giving the echinoid a beard-like appearance. Distribution: The species is known from the East African Coast off Zanzibar, Indian Ocean; no depth is stated (Fig. 3.227). Diploporaster savignyi (Fourtau, 1904) Test: Elongate with very distinct anterior notch; anteriorly low, rising toward highest point immediately behind the apical system, posterior interambulacrum keeled, at rear truncated; length 45 mm. Apical system: Posterior of center, four gonopores, the anterior pair reduced in size. Anterior ambulacrum: Broad and deeply depressed, becoming more shallow toward the margin, adjacent interambulacra overhanging; pore pairs in double rows, associated tube feet specialized for funnel-building and food-gathering. Paired petal: Depressed and distally flexed, posterior pair less than half the length of anterior pair.

Oral side: Plastron flattened, sides rising to the margin; sternum broad, adjacent ambulacra narrow; periproct at the posterior end about half way up. Fascioles: Peripetalous fasciole well-developed, running closely along the petals in interambulacra IA 1 and IA 4; lateroanal fasciole with deep bend below the periproct. Tubercles: Uniformly small and dense, larger toward the anterolateral margin and at the oral side. Color: The bare test is whitish. Distribution: The species is known from the Red Sea and the Arafura Sea, living in shallow water. Literature: A.M. Clark & Rowe 1971. Genus Hypselaster H.L. Clark, 1917 Mortensen (1951) stated that the genus Hypselaster is rather unsatisfactorily defined. It is characterized by having two gonopores, the lateroanal fasciole being rudimentary or lacking. Test: More or less inflated, posterior end truncated; in outline oval with deep anterior notch. Apical system: Subcentral with two gonopores, lacking in anterior plates, ethmolytic. Anterior ambulacrum: Deeply sunken, pore pairs associated with specialized tube feet. Paired petals: Also deeply depressed, posterior pair shorter than anterior pair, which is distally flexed outward. Oral side: Convex, peristome kidney-shaped and more or less overarched by the distinct lip of the labrum; phyllodes more or less developed; periproct small and marginal at the posterior side; labral plate elongated, extending to the first or second adjacent ambulacral plates; sternal plates symmetrical, in broad contact with labral plate. Fascioles: Margino-peripetalous fasciole well-developed, lateroanal branch present in more or less distinct traces on the subanal area or entirely lacking. Tuberculation: Fine and more or less uniform, spines slender, on plastron distally spoon-shaped. The genus contains 10 species, which are rather similar. Some might be synonymous. Hypselaster affinis Mortensen, 1948. Malayan region and Philippines. Hypselaster brachypetalus H.L. Clark, 1917. Grenada, West Indies. Hypselaster dolosus H.L. Clark, 1938. West Australia. Hypselaster fragilis (A. Agassiz & H.L. Clark, 1907). Japan.

▸ Fig. 3.227: (A–C) Diploporaster barbatus. Test length 59 mm, Zanzibar. ZMUC ECH 91. (A) Aboral side. The anterior ambulacrum is wide and deeply sunken. (B) Oral side: the strong plastronal spines are spoon-shaped. Toward the posterior end, two tufts of finer spines are developed. (C) Side view: the oral side is convex, the plastron being set with solid spines. (D–F) Diploporaster savignyi. Test length 37 mm; Sorong, New Guinea. (D) Aboral side: the anterior ambulacrum is deeply depressed with a broad notch at the margin. (E) Oral side: the sternum is wide, the adjacent ambulacra rather narrow. (F) Profile: the highest point of the test lies immediately behind the apical system.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

B

C

D

E

F

 299

300 

 3.3 Superorder Atelostomata von Zittel, 1879 Paired petals: Posterior pair less than half the length of anterior pair. Labral plate: Forms a prominent lip and extends to first adjoining ambulacral plates. Fascioles: Margino-peripetalous fasciole only slightly indented between the petals; lateroanal branch distinct only as a straight line far below the periproct, continuing for a short distance upward along the posterior edge of the test. Distribution: Malayan Archipelago and Philippines, at depths of 915 to 1,630 m. Remarks: The paratype in Copenhagen is now broken (Fig. 3.228).

Hypselaster jukesii (Gray, 1855). Northern Australia to Queensland. Hypselaster kempi Koehler, 1914. Laccadive Sea and Bay of Bengal. Hypselaster limicolus (A. Agassiz, 1878). Gulf of Mexico. Hypselaster maximus (A. Agassiz & H.L. Clark, 1907). Fragments(?), Hawaiian Islands approximately 500 m. Hypselaster rotundus (A. Agassiz & H.L. Clark, 1907). Southern Japan. Literature: H.L. Clark 1917; A.M. Clark & Rowe 1971; ­McNamara & Philipp 1980. Hypselaster affinis Mortensen, 1948 Test: Oval, narrowing toward the posterior truncated side, shallow anterior notch; elevated profile, largest height behind the peripetalous fasciole, posterior interambulacrum keeled; anterior interambulacra forming a rather sharp keel adjacent to the anterior ambulacrum; length 48 mm. Apical system: Distinctly posterior, madreporite not extending beyond posterior ocular plates. Anterior ambulacrum: Shallowly sunken.

A

E

B

Hypselaster brachypetalus H.L. Clark, 1917 Main characteristics of the genus: holotype 25 mm, it may be a young specimen. Anterior ambulacrum: Not much sunken. Occurrence: Grenada, West Indies; depth 1,750 m. Hypselaster dolosus H.L. Clark, 1938 Main characteristics of the genus: length approximately 50 mm. Very much like H. jukesii. Thought to be synonymous.

D

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G

H

I

J Fig. 3.228: (A, B) Hypselaster affinis. Test length 48 mm, Philippines. ZMUC ECH 63, from Mortensen (1951). (A) Aboral side: the test is tapering toward the posterior. (B) Oral side: the posterior end is concavely. (C, D) Drawings after Mortensen (1951). (C) Peristome and adjacent plates: the labrum shows a distinct lip. Scale bar is 6 mm. (D) Apical system: scale bar is 4 mm. (E, F) Hypselaster brachypetalus. Test length 25 mm. Peristome plates and apical system. Scale bar is 2 mm; after H.L.Clark (1917). (G–J) Hypselaster fragilis. Young specimen of 16 mm, Southern Japan. From H.L. Clark (1917). (G) Aboral side. (H) Oral side. (I) Posterior side. (J) Side view.

3.3.2 Order Spatangoida L. Agassiz, 1840 



 301

Occurrence: Western Australia from Broome to the Augustus Islands in the littoral.

Occurrence: This species is only recorded from the Kashika Islands, Southern Japan; depth 700 m (Fig. 3.229).

Hypselaster fragilis (A. Agassiz & H.L. Clark, 1907) In the only known young specimen, the apical system is anterior. On the oral side, the lateral interambulacra are excluded from the peristome. The margino-peripetalous fasciole is well-developed, but there is no trace of a lateroanal branch.

Hypselaster jukesii (Gray, 1855) According to Mortensen (1951), the apical system in H. dolosus is positioned posterior of the center and in H. jukesii anterior of the center. However, Mortensen’s photos of H. jukesii clearly show a posterior apical system. Also, the supposed differences in the shape of the petals and the

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Fig. 3.229: (A–C) Hypselaster jukesii. Test length 40 mm, Broome, Western Australia. ZMUC (as H. dolosus). (A) Aboral side: the peripetalous fasciole is very broad. (B) Oral side. (C) Side view. (D, E) Photos in Mortensen (1951). (D) Aboral side: the apical system is clearly posterior of center. (E) Side view: the posterior part of the test is strongly raised and vertically truncated. (F) Peristome plates. Scale bar is 5 mm; after Mortensen (1951), as H. dolosus.

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 3.3 Superorder Atelostomata von Zittel, 1879

valves of the globiferous pedicellariae are not borne out by Mortensen’s illustrations. A.M. Clark & Rowe (1971). Therefore, H. dolosus and H. jukesii are thought to be synonymous. Main characteristics of the genus: maximum length 60 mm. Test: With distinct anterior notch, raised toward the ­posterior. Apical system: Posterior of center. Anterior ambulacrum: Deeply sunken; with regular series of pore pairs. Oral side: Labrum extending to middle of first ambulacral plates; lip; oral interambulacra 1 and 4 in contact with peristome. Fascioles: No subanal traces of lateroanal fasciole. Distribution: The species is recorded along the north coast of Australia to Brisbane in the east; bathymetric range 0 to 50 m. Hypselaster kempi Koehler, 1914 Main characteristics of the genus: length of the holotype 43 mm. Test: Oval with shallow anterior notch; wedge-shaped with elevated posterior part. Apical system: Slightly posterior. Anterior ambulacrum: Shallowly depressed, pore pairs in regular series. Fascioles: Margino-peripetalous fasciole broad, lateroanal branch rudimentary. Distribution: Indian Ocean, Bay of Bengal and Laccadive Sea, at depths of 1,360 to 1,935 m. Hypselaster limicolus (A. Agassiz, 1878) Main characteristics of the genus: Test: High and compact, posterior end almost vertically truncated; in outline, subcircular with shallow anterior notch, largest width at the middle of the anterior petals, narrowing toward the posterior. Apical system: Subcentral. Anterior ambulacrum: Depressed aborally. Labrum: With distinct lip, extending to second adjacent ambulacral plates. Fascioles: Margino-peripetalous fasciole well-developed, lateroanal branch present in traces or absent. Distribution: This species is recorded from the Gulf of Mexico; Caribbean, bathymetric range 125 to 260 m (Fig. 3.230). Hypselaster maximus (A. Agassiz & H.L. Clark, 1907) Length of test approximately 110 mm. It was described on the basis of a fragment, which showed some larger tubercles within the peripetalous fasciole. Distribution: Hawaii Islands, USA; depth 385 to 565 m.

Hypselaster rotundus (A. Agassiz & H.L. Clark, 1907) It was described on the basis of a single specimen; the labrum extending to the second adjacent ambulacral plates. Distribution: Southern Japan; depth 71 m. Genus Moira A. Agassiz, 1872 Test: Highly vaulted with concavely truncated posterior side, subanal area more or less projecting; in outline elongated with anterior notch and more or less roundly pointed posterior end. Apical system: Ethmolytic, two gonopores, anterior gonopores lacking. Anterior ambulacrum: Deeply depressed adapically, becoming shallow toward the margin; widening below the protruding adjoining interambulacra; pore pairs set with specialized tube feet, arranged uniserially. Paired petals: Also deeply depressed, forming widened cavities below the protruding adjoining interambulacra, leaving slit-like openings on aboral surface; anterior petals longer than posterior pair. Oral side: Slightly convex, peristome large, wider than long, more or less overarched by the labrum; periproct small, marginal at the subvertical or concave posterior side; labral plate short and wide, extending to first adjacent ambulacral plates; broadly contacting the large, symmetrical sternal plates. Fascioles: Margino-peripetalous fasciole well-developed, passing immediately along the edges of the petals; lateroanal fasciole thin, strongly bent below the periproct. Tuberculation: Aborally and orally small, dense and uniform; spination fine and slender, plastronal spines with spoon-shaped distal end. Remarks: This genus differs from all other schizasterids in having deeply sunken ambulacra each forming a cavity with slit-like openings toward the aboral surface. Burrowing and feeding behavior of a spatangoid (Moira atropos): This species lives in the Caribbean region in shallow water. The sea urchin digs itself into the sediment using the spines at the sides. As it descends and the distance to the surface increases, the spines of the apical tuft continually deposit mucus onto the wall of the funnel, which leads to the open water, to consolidate it (Fig. 3.231). Long tube feet with large discs ending in numerous papillae extend to the surface of the substrate. They are efficient rakers for organic material, which then falls down the tube onto the test of the sea urchin. There, the particles are incorporated into a mucus string moving down the bottom of the anterior ambulacrum and around the anterior notch to the mouth. The large tube feet in the phyllodes, set with numerous finger-like digits, take parts of the string and lead them to the mouth. If the surface of the substrate is detritusrich, the animal may remain stationary for several days.

3.3.2 Order Spatangoida L. Agassiz, 1840 



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Fig. 3.230: (A, B) Hypselaster kempi. Test length 43 mm, Indian Ocean. From Koehler (1914). (A) Aboral side. (B) Side view: the test rises strongly toward the posterior. (C–E) Hypselaster limicolus. Test length 51 mm, Gulf of Mexico, Caribbean. ZMUC. (C) Aboral side: the test is almost as wide as long. (D) Oral side: the lip of the labrum overarching the peristome is conspicuous. (E) Side view: the test is high with an almost vertical posterior face. (F) Labrum and adjacent plates. Scale bar is 5 mm. After Mortensen (1951).

Otherwise, M. atropos moves forward by means of the strongly spatulate spines on the sternum. The uppermost podia in the anterior ambulacrum assimilate mucus from the fasciolar spines and excavate with short, quick strokes a new funnel and plastering the walls. The old funnel is abandoned. While the new tube is being built and little or no food is coming down from the surface, the phyllodial podia selectively gathers organic particles in the sediment in front and below the test and passes them directly to the mouth. Sensory tube feet continuously examine the substrate. The funnel is also used for bringing fresh water into the burrow. A water current, created by the cilia of the fasciolar spines, flows over the petals providing the respiratory podia with fresh, oxygenated water; and also sweeps the waste to the rear of the test. Literature: Nichols 1960; Chesher 1963, 1967, 1969; McNamara & Philipp 1980; Hendler et al. 1995

Five species are included in the genus: Moira atropos (Lamarck, 1816). From Cape Hatteras, USA, through the Caribbean to Sao Paulo, Brazil. Moira clotho Michelin, 1855. West coast of the Americas from the Gulf of California to Ecuador. Moira lachesinella Mortensen, 1930. Southern Japan and East China Sea. Moira lethe Mortensen, 1930. Australia, from Bowen around the south coast to Broome in the west. Moira stygia (A. Agassiz, 1872). Red Sea and East Africa to the Malayan Archipelago. Biology: The genus is very well adapted to muddy bottoms, where it may burrow to a depth of 15 cm, but it can also live in sands or coral gravel (Fig. 3.232). The American species differ from the three Indo-West Pacific species in the shape of the posterior side: more or less vertically truncated in the former, with a subanal projection and a concave profile in the latter.

304 

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 3.3 Superorder Atelostomata von Zittel, 1879

apical tuft of spines

spatulate spines

aboral food-gathering tube feet funnel-building tube feet

phyllodal food-gathering tube feet

sensory tube feet

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aboral food-gathering tube feet funnel-building tube feet

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spatulate spines

phyllodal food-gathering tube feet

sensory tube feet

Fig. 3.231: (A, B) Diagrammatical longitudinal section of the progressive stages I and II in burrow building of the genus Moira. The animal moves to the right. Modified after Chesher (1967).

Moira atropos (Lamarck, 1816) Test: Very thin, highly vaulted; outline rounded anteriorly with a distinct notch, slightly pointed at the posterior end; in profile anterior side arched, posterior end vertical, length up to 57 mm. Apical system: Posterior, depressed and with two ­gonopores. Anterior ambulacrum: Deeply depressed, pore pairs uniserial, associated tube feet penicillate and strongly expandable.

Paired petals: Deeply sunken, overarched by adjacent interambulacra. Oral side: Convex, anterior furrow leading to the large mouth; lip projecting, phyllodes short. Fascioles: Peripetalous fasciole broad, running mostly immediately around the edge of the petals and the anterior ambulacrum; lateroanal fasciole narrow. Spines: Aborally short, becoming longer toward the anterolateral margin; sternal spines spatulate.

3.3.2 Order Spatangoida L. Agassiz, 1840 



 305

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posterior

spatulate spines

Fig. 3.232: (A–C) Moira atropos. Test length 41 mm, Florida, USA. (A) Aboral side: the paired petals are almost overbridged by the adjacent interambulacra; in the anterior ambulacrum, the opening is broader to leave access for the specialized tube feet. The peripetalous fasciole runs immediately beside the petals. (B) Oral side: the peristome is large and kidney-shaped, in the phyllodes, only a few pores with tube feet are developed. (C) Profile: the posterior end is vertically truncated, there is no projection below the periproct. (D–F) Specimen with spines. Test length 45 mm; Puerto Rico. ZMH. (D) Aboral side: A tuft of spines is developed around the apical system and used for funnelbuilding. (E) Oral side: the locomotory spines of the plastron are spatulate, the spines at the sides long and arched, serving for burrowing. (F) Profile: the spines of the apical tuft maintain the funnel walls with mucus, the spines at the sides are used for burrowing, and the spatulate spines of the base for locomotion in the sediment.

306 

 3.3 Superorder Atelostomata von Zittel, 1879

Color: In life, the sea urchin is light brown to yellowishwhite, often with horseshoe-shaped, dark red-brown pigmented fasciole near the apical system. Distribution: The species is known from Cape Hatteras, NC, USA and the Bermudas throughout the Caribbean to Sao Paulo, Brazil. It lives buried in the intertidal zone and to a depth of 450 m, but is most common in less than 50 m. Moira clotho Michelin, 1855 Main characteristics of the genus: length of test may reach 50 mm.

Test: High, compact, posterior side subvertically truncated. Apical system: Posterior, abruptly sunken, two gonopores distant from each other. Oral side: Convex, labrum slightly projecting the ­peristome. Larger spines: Tuft in front of the apical system. Distribution: This species is recorded from the northern end of the Gulf of California to Ecuador, living in the ­littoral. Remarks: M. clotho is distinguished from M. atropos by its less projecting labrum. In the latter, the peristome is almost entirely covered by the protruding labrum (Fig. 3.233).

A

B

C

D

Fig. 3.233: (A–C) Moira clotho. Test length 31 mm, Gulf of California, Mexico. ZMUC. (A) Aboral side: the peripetalous fasciole passes mostly immediately along the edges of the ambulacral invaginations. (B) Oral side. (C) Side view: the posterior end is vertically truncated. (D) Peristome, labrum, and adjacent plates. Scale bar is 6 mm (Mortensen 1951).

3.3.2 Order Spatangoida L. Agassiz, 1840 



The two American species differ from the three IndoPacific species by their subvertical posterior side. In those, the posterior end is concave, forming a more or less conspicuous beak. Moira lachesinella Mortensen, 1930 Main characteristics of the genus: length of the test reaches approximately 40 mm. Test: High, sloping toward the anterior, posterior side vertically concave, with conspicuous beak. Apical system: Subcentral, depressed, but with sloping sides, not abruptly sunken as in M. clotho and M. atropos. Labrum: Very short, moderately covering the large peristome. Depression above the subanal bow of the lateroanal fasciole very conspicuous; posterior beak strongly pointed.

A

B

 307

Distribution: East China Sea and Southern Japan northward to Sagami Bay, intertidal to a depth of approximately 50 m. Remarks: In the shape of the test, M. lachesinella strongly resembles M. lethe. Mortensen (1951) described the subanal depression in the former as being less sharply limited than in the latter, but this gradual difference is sometimes hardly recognizable. However, the globiferous pedicellariae show a distinct difference between both species: in M. lachesinella, there are freely projecting thorns on the basal thickening of the stalk, whereas in M.  lethe, these structures are not developed. In addition, the two species live in well separated geographic regions: M. lachesinella in the East China Sea and Southern Japan, M. lethe along the coasts of Southern Australia from Bowen, Queensland to Broome, Western Australia (Fig. 3.234).

C

D

E

F

G

Fig. 3.234: (A–D) Moira lachesinella. Test length 31 mm, Hakata Bay, South Japan. ZMUC ECH 482. (A) Aboral side: the spines are densely appressed to the test. (B) Oral side: the plastronal spines are spoon-shaped. (C) Side view. (D) Posterior side: below the periproct there is a deep depression, surrounded by the subanal branch of the fasciole. (E–G) Specimen without spines. Test length 37 mm, northeast of Taiwan. (E) Aboral side. (F) Oral side. (G) Posterior side.

308 

 3.3 Superorder Atelostomata von Zittel, 1879

Moira lethe Mortensen, 1930 Test: Very thin, very highly vaulted, posterior end concave, projecting heel below the periproct; in outline strongly pointed to the posterior; length up to 61 mm. Apical system: Anterior, depressed with two gonopores. Anterior ambulacrum: Deeply depressed, furrow at the margin shallow; pore pairs in uniserial row. Paired petals: Deeply sunken, opening slit-like. Oral side: Flattened, anterior furrow to the very large peristome only slightly depressed, short phyllodes; periproct high at the posterior side, sharply pointed. Fascioles: Peripetalous fasciole broad, leading mostly around the edge of the paired petals and the anterior ambulacrum; lateroanal fasciole narrow, forms a deep bend below the periproct, bounding the pit at the posterior beak. Spines: Short and densely packed; spoon-shaped on the sternum. Color: In life, M. lethe is light brown, the denuded test is white. Distribution: This species is confined to Australia from Bowen, Queensland, around the south coast, and at the west side northward to Broome, Western Australia. It lives buried from the intertidal to a depth of 10 m (Fig. 3.235). Literature: Rowe & Gates 1995; Miskelly 2002. Moira stygia (A. Agassiz, 1872) Main characteristics of the genus: Test: High with obliquely sloping anterior and steep, concave posterior side; posterior beak inconspicuously pointed. Apical system: Subcentral, depressed, but with sloping sides, not abruptly sunken as in M. clotho and M. atropos. Oral side: Peristome largely covered by the labrum, sternum hardly widening toward the posterior. Fascioles: Slight impression above the subanal bow of the lateroanal fasciole. Distribution: This species is recorded in the Indo-West Pacific from the Red Sea and East Africa to the Malayan Archipelago; littoral. Remarks: This species is well-distinguished from M. lachesinella and M lethe by its shallow subanal impression and by its longer labral plate (Fig. 3.236). Genus Moiropsis A. Agassiz, 1881 This genus is nearly related to Moira, but differs in the length of the anterior ambulacrum, which is shorter than the anterior paired petals. Mortensen (1951) treated Moiropsis as a subgenus of Moira. Moiropsis claudicans A. Agassiz, 1881 Test: High, anterior side obliquely sloping, posterior side concavely truncated; posterior interambulacrum keeled; in outline heart-shaped with distinct anterior notch.

Apical system: Posterior, ethmolytic with two gonopores; posterior genital plates distant from each other. Anterior ambulacrum: Deeply depressed near the apical system, shallower toward the ambitus; pore pairs and podia specialized for funnel-building, forming a single series; depression shorter than in anterior petals. Paired petals: Also with deep depressions, openings very narrow, but wider than in Moira. Oral side: Peristome large with projecting labrum, labral plate short and wide, extending to first adjacent ambulacral plates; periproct small, positioned high on the posterior face. Fascioles: Margino-peripetalous fasciole closely following the edges of the petals; lateroanal fasciole thin, strongly bent at the slightly protruding subanal area. Tubercles: Aborally fine, dense and uniform, oral side also densely tuberculated. Occurrence: This species is only recorded from the Kei Islands, Indonesia, at a depth of 220 m. A fossil species from the Miocene was found in Taiwan. Genus Ova Gray, 1825 This genus is distinguished from Brisaster and Schizaster by the number of their gonopores: Ova has usually two genital pores, whereas Brisaster has three, and Schizaster has four, usually. It has a well-developed lateroanal and a portion of marginal fasciole. Test: Oval with deep anterior notch, posterior end more or less pointed; in profile wedge-shaped with low anterior and high, steeply truncated posterior side. Apical system: Posterior, ethmolytic, with two gonopores. Anterior ambulacrum: Very deeply depressed adapically, becoming shallower toward the margin; pore pairs and podia specialized for funnel building. Paired ambulacra: Also distinctly sunken, posterior petals very short, anterior pair much longer and flexed outward, anterior column reduced. Oral side: Convex; peristome wider than long, more or less overarched by the labrum; periproct small, high on the subvertically truncated posterior side; labral plate short and wide, with lip; not extending to second adjacent ambulacral plate; sternal plates forming a broad oval, fully tuberculate. Fascioles: Complete marginal fasciole and peripetalous bow well-developed. Tuberculation: Aborally fine, dense and uniform; oral tuberculation larger. Spines: Aborally long, fine and slender; plastronal spines with spoon-shaped ends. Five Recent species are known: Ova canalifera (Lamarck 1816). Mediterranean. Ova edwardsi Cotteau, 1889. West Africa.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

 309

B

C

D

Fig. 3.235: (A–D) Moira lethe. Test length 31 mm; Mackay, Queensland, Australia. (A) Aboral side: the petals are deeply sunken, the peripetalous fasciole passes along the edges. (B) Posterior side: the projecting heel has a distinct pit. (C) Profile: the lower end of the posterior side projects and is pointed. (D) Specimen with spines. Spoon-shaped spines on the plastron. The spines are used for locomotion in fine, sandy or muddy sediments. The spines are used for locomotion in fine, sandy, or muddy sediments. MfN Berlin; scale bar is 5 mm.

Ova lacunosa (Linné, 1758). East Africa to North Australia and Japan. Ova orbignyana (A. Agassiz, 1880). Northwest Atlantic in deeper water. Ova portjacksonensis McNamara & Philip, 1980. Port Jackson, New South Wales, Australia. Literature: McNamara & Philip 1980; Shigei 1986; A.M. Clark & Rowe 1971; Rowe & Gates 1995; McNamara 1995 (Fig. 3.237).

Ova canaliferus (Lamarck, 1816) Test: Thin, in outline heart-shaped, in profile wedgeshaped, the low anterior side rising steeply toward the posterior, highest point behind the apical system, posterior interambulacrum keeled; maximum length approximately 74 mm. Anterior ambulacrum: Deeply sunken, pore pairs in irregular double row below the overhanging interambulacra.

310 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

D

C

E

F

G

H

Fig. 3.236: (A–D) Moira stygia. Test length 22 mm, Zanzibar, East Africa. ZMUC ECH 302. (A) Aboral side. (B) Oral side. (C) Side view. (D) Posterior side: the subanal impression is shallow. (E–H) Moiropsis claudicans. Kei Islands, Indonesia. No scale; from Agassiz (1881). (E) Aboral side. (F) Oral side. (G) Side view: the anterior is to the right. (H) Petaloid area.

3.3.2 Order Spatangoida L. Agassiz, 1840 



 311

B

A

C

D

E

F

Fig. 3.237: Ova canalifera. (A–D) Test length 53 mm; Hvar, Croatia. (A) Aboral side: the anterior ambulacrum is deeply depressed, the adjacent interambulacra are overhanging. (B) Oral side: the plastron is broad and fully tuberculated. (C) Profile: the posterior part of the test is much higher than the anterior, the end is truncated. (D) Oblique view into the anterior furrow: the row of pore pairs below the overhanging interambulacra is irregularly doubled. (E, F) Specimen with spines. Test length 62 mm, Naples, Italy. ZMH. (E) Aboral side: the spines are longer around the anterior. (F) Oral side.

312 

 3.3 Superorder Atelostomata von Zittel, 1879

Oral side: With flat plastron, the sides rising to the margin; peristome near the anterior furrow; very prominent lip; periproct high at the slightly concave posterior end. Fascioles: Peripetalous fasciole closely surrounding the ends of the petals, lateroanal branch narrow with deep bend below the periproct. Spines: Short, fine, and densely packed, size increasing toward anterior on aboral and oral sides, set with larger tubercles; spoon-shaped over the plastron. Color: In life, the spines are light brown, the fascioles marked by darker bands. The bare test is yellowish-gray. Distribution: The species is confined to the Mediterranean from Spain to the Dardanelles, Turkey. It is found from the littoral zone to a depth of approximately 100 m.

A

D

Ova edwardsi Cotteau, 1889 Main characteristics of the genus: length may reach 60 mm. Test: Strongly inflated aborally, posterior interambulacrum conspicuously keeled; posterior face concavely truncated. Anterior ambulacrum: With a single, fairly regular series of pore pairs per column, the adjacent interambulacra leaning over. Posterior petals: Broad, very short and diverging. Distribution: This species is restricted to West Africa from Cape Palmas to Angola; bathymetric range: 17 to 66 m. Remarks: Usually, the frontal ambulacrum in O. edwardsi is distinctly broader than in the other species, the adjacent interambulacra less overhanging the series of pore pairs (Fig. 3.238). The World Echinoidea Database speaks

C

B

E

G

F

Fig. 3.238: (A–D) Ova edwardsi. (A–C) Test length 60 mm; Angola, West Africa. ZMUC. (A) Aboral side: the sunken anterior ambulacrum is very broad. (B) Side view. (C) Anterior side. (D) Specimen with spines. Test length 31 mm, Gulf of Guinea, West Africa. ZMUC. Aboral side: long spines overarch the sunken ambulacra. (E–G) Ova lacunosus. Test length 49 mm; Sagami Bay, central Japan. ZMH. (E) Aboral side: the anterior furrow is broad. The pore pairs form one regular row on each side. (F) Side view. (G) Specimen with spines. Test length 59 mm, Hong Kong, China. ZMUC. Aboral side.

3.3.2 Order Spatangoida L. Agassiz, 1840 



from Schizaster canaliferus. The author cannot decide what is right. Ova lacunosa (Linné, 1758) This species is very similar to Ova canalifera. It differs in the arrangement of the pores in the deeply depressed anterior ambulacrum, which are developed as a single row of pore pairs on each side. Distribution: The species is recorded from Natal, East Africa to the northern coast of Australia and to Japan. It lives at a depth of 4 to 90 m. Remark: In the World Echinoidea Database O. lacunosa is written as Schizaster lacunosus, source A.M. Clark & Rowe (1971). Neither Clark & Rowe nor anything else mentioned the gonopores (two or four).

Ova orbignyana (A. Agassiz, 1881) Main characteristics of the genus: maximum length may reach 60 mm. Test: Strongly inflated, sloping strongly from the highest point behind the apical system toward the anterior margin, posterior side subvertically truncated, but not concave. Color: In the living echinoid, the fascioles are dark brown or purple. Distribution: From Massachusetts (Martha’s Vineyard), along the east coast of the USA and in the Eastern Gulf of Mexico, off Florida, and Louisiana; off the South American Coast from Venezuela to Barbados and Surinam. Bathymetric range: 26 to 500 m.

B

A

C

 313

D

E

Fig. 3.239: (A–C) Ova orbignyana. Test length 41 mm; off Martha’s Vineyard, MA, USA. ZMUC. (A) Aboral side. (B) Oral side: the sternum is oval. (C) Side view: the upper side slopes straightly from behind the apical system toward the anterior margin. (D, E) Ova portjacksonensis. Test length 37 mm, endemic to Port Jackson, New South Wales. (D) Aboral side. (E) Side view. Sketches after the photos in McNamara & Philip (1980).

314 

 3.3 Superorder Atelostomata von Zittel, 1879

Biology: The animals live buried in fine sediment, using their specialized anterior tube feet for funnel-building and food-gathering. Remarks: In O. orbignyana, the aboral side slopes more straightly from behind the apical system toward the anterior margin. In addition, the geographical range is well separated. However, the range of variation in the species is high (Fig. 3.239). Ova portjacksonensis McNamara & Philip, 1980 Main characteristics of the genus: test length 60 mm. Test: Short and broad; posterior petals widely spaced. Apical system: With two gonopores. Oral side: Labrum short, with ventral projections and medially constricted. Distribution: Port Jackson, New South Wales, Australia, at approximately 15 m. ­ ono­pores. Remarks: McNamara speaks definitely from two g Genus Protenaster Pomel, 1883 The genus is monospecific. Protenaster australis Gray, 1851 Test: Rather strong, in outline oval with broad, shallow anterior notch; in profile inflated with steep anterior and posterior sides, upper side more or less flattened; length up to 83 mm. Apical system: Slightly anterior, four gonopores, very long expanded madreporite Anterior ambulacrum: Broad and shallowly sunken; rows of pore pairs uniserial. Paired petals: Approximately equal length, anterior pair forming an angle of approximately 180°, distally slightly flexed; pore pairs near the apical system distinctly smaller. Oral side: Flattened; peristome kidney-shaped, slightly sunken, lip more or less projecting; phyllodes distinct; plastron broad; periproct vertically elongate, pointed, at half the height of the posterior side. Fascioles: Peripetalous fasciole surrounds the petals closely, indented in lateral and posterior interambulacra; lateroanal fasciole with an abruptly deep bend to the oral side below the periproct. Tubercles: On the aboral side small, larger ones bordering the anterior ambulacrum; spines small and uniform, larger at anterior side both aborally and orally.

Color: In life, light to darker brown, bare test whitish or very light pinkish. Habitat: The species lives in fine calcareous sands, buried to a depth of approximately 10 cm below the sea floor. It is found from the ebb zone to a depth of 10 m. Distribution: P. australis is confined to Australia from Sydney around the southern coast to Perth, Western Australia. Literature: McNamara & Bryce 1983; McNamara 1985; Rowe & Gates 1995; Miskelly 2002 (Fig. 3.240). Genus Schizaster L. Agassiz, 1836 Numerous species with two gonopores have been established in the genus Schizaster L. Agassiz, 1836. Unfortunately, the fossil type species turned out to have four gonopores, and consequently, the genus name Schizaster has to be synonymized with Paraster Pomel, 1869. The species with two gonopores now have the genus name Ova Gray, 1825. The genera Ova and Schizaster are still the subject of discussion. They are very similar, some authors felt them synonymous, and they show the amount of gonopores once with two or three and four. See the Remarks in Ova lacunosa. It is usually characterized by having four gonopores, the anterior pair may be smaller than the posterior pair. Seven extant species are included in the genus ­Schizaster: Schizaster compactus (Koehler, 1914). Indian Ocean, India, and West Australia. Schizaster gibberulus L. Agassiz & Desor, 1847. Red Sea and Indian Ocean. Schizaster doederleini Chesher, 1972. Caribbean. Schizaster floridiensis (Kier & Grant, 1965). Caribbean. Schizaster myorensis McNamara & Philip, 1980. Myora, Moreton Bay, Queensland, Australia. Schizaster obovatus Lindley, 2004. New Britain, Papua New Guinea. Schizaster rotundatus Döderlein, 1906. Galapagos Islands. Test: Ovoid with moderate frontal notch; aboral side high, posterior interambulacrum keeled, forming a slight hook above the periproct, posterior face vertically truncated. Apical system: Ethmolytic, four gonopores, the anterior pair usually being smaller than the posterior pair. Anterior ambulacrum: Distinctly depressed, deepest at fasciole, becoming shallow toward the ambitus; pore pairs and podia arranged in a single row at each column, specialized for funnel-building.

▸ Fig. 3.240: Protenaster australis. (A–C) Test length 61 mm; Sydney, Australia. (A) Aboral side: the petals are distinctly sunken, the anterior pair forming an angle of approximately 180°. (B) Oral side: the tubercles increase in size to the anterior and bear correspondingly larger spines. (C) Profile: the upper side is rather flattened. The fascioles are sharply curved. (D–F) Specimen with spines: length with spines 61 mm; Port Jackson, New South Wales, Australia. (D) Aboral side. (E) Oral side. (F) Side view. The spines on the anterior side are longer and more solid. Those on the plastron are spatulate.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

B

C

D

E

F

 315

316 

 3.3 Superorder Atelostomata von Zittel, 1879

Paired petals: Sunken, posterior pair half the length of anterior pair, more or less flexed. Oral side: Convex; peristome kidney-shaped, labrum with lip slightly projecting; periproct high on the posterior truncated face; labral plate short and wide, mushroomshaped, not reaching beyond the first adjacent ambulacral plates, broadly contacting the large and symmetrical sternal plates. Fascioles: Margino-peripetalous and lateroanal fasciole well-developed, subanal podia penicillate. Tuberculation: Aborally fine, dense and uniform, oral tuberculation larger, plastronal spines ending in spoonshaped tips. Schizaster compactus (Koehler, 1914) This species greatly resembles S. gibberulus. In that species, the pore pairs in the frontal ambulacrum are smaller and placed obliquely; in S. compactus, the pores of a pair are larger and positioned more horizontally. Test: Broadly oval with distinct anterior notch; aboral side inflated, interambulacral keel rounded, slightly overhanging the subvertical posterior face; length up to 40 mm. Apical system: With two, three, or four gonopores. Anterior ambulacrum: Sunken, very regular rows of pore pairs, the pairs positioned horizontally. Paired petals: Flexed and depressed. Oral side: Labral plate with conspicuous lip. Distribution: This species lives in the Indian Ocean in the Bay of Bengal and has also been recently recorded in Western Australia; depth 35 to 900 m (McNamara 1995). Remarks: McNamara (1995) reported on this species occurring off the coast of Northwest Australia, in the Dampier Archipelago. The population included specimens with two, three, and four gonopores (Fig. 3.241). Schizaster doederleini Chesher, 1972 Test: Strongly inflated, posterior part higher than anterior part; posterior interambulacrum keeled, forming a conspicuous hook overhanging the truncated posterior face; in outline nearly as broad as long, with distinct anterior notch, more or less pointed posteriorly; length may reach 75 mm. Apical system: Slightly posterior, anterior pair of gonopores much smaller than posterior pair. Anterior ambulacrum: Sunken and narrow, parallel-sided. Paired petals: Relatively short, posterior pair less than half the length of anterior pair. Oral side: Peristome not sunken; periproct small, high on posterior surface; labrum short, extending to the first adjoining ambulacral plates. Fascioles: Margino-peripetalous fasciole conspicuously widened below the petals; lateroanal fasciole forming a

straight line from below the anterior petals to the rear; well-developed subanal tube feet. Color: In live specimens, the test and spines are yellowish tan or light brown, the margino-peripetalous fasciole is darker brown. The podia in the anterior ambulacrum may be bright red. Distribution: Recorded from the Florida Keys, the Dry Tortugas and the Bahamas, and off Belize and Northern Colombia; 12 to 220 m. Biology: This species lives at a depth of approximately 20 to 100 mm deeply burrowed in mud fields. Remarks: S. doederleini can be distinguished from O. orbignyana by the shape of the sunken anterior ambulacrum: in O. orbignyana, the upper portion of the furrow is formed by a ridge of nontuberculated interambulacral plates, whereas in S. doederleini, the plates are densely set with tubercles up to the edge and without forming a distinct ridge (Fig. 3.242). Schizaster floridiensis (Kier & Grant, 1965) Main characteristics of the genus: recorded length 40 mm. S. floridiensis differs from S. doederleini by its more rounded outline of test, the larger petaloid area, the posterior petals being less distant from the posterior end, and by its larger periproct. Color: Test and spines light pink to faint yellow; fascioles and tube feet of petals are red, the podia in the anterior ambulacrum yellow-orange. Distribution: Recorded at the Florida Keys, the Bahamas, Dominica and the San Blas Islands, off Panama; depth 2 to 25 m. Biology: The species lives in mud or sand fields, burrowing to a depth of 250 mm. Schizaster gibberulus L. Agassiz & Desor, 1847 Test: Ovate with anterior notch; in profile anterior side curved to the margin, posterior side truncated or even concave; highest point behind the apex; length up to 40 mm. Apical system: Posterior to center (in adults), four gonopores, the anterior pair usually smaller than the posterior pair. Anterior ambulacrum: Distinctly sunken at the aboral side, becoming shallow toward the margin; row of pore pairs uniserial. Paired petals: Sunken, anterior pair much longer than posterior pair. Oral side: Flattened or slightly convex, peristome kidneyshaped, lip protruding; periproct high at the posterior end below overhanging interambulacrum. Fascioles: Peripetalous fasciole with indentations in  all interambulacra, distinct bend in anterior

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

 317

B

D

C

E

Fig. 3.241: Schizaster compactus. (A, B) Test length 35 mm; Madras, India. ZMUC. (A) Aboral side. (B) Side view. (C, D) Specimen with spines. Test length 37 mm, Madras, India. ZMUC. (C) Oral side: the lip is very conspicuous. (D) Side view. (E) This species may have two, three, or four gonopores. It is very unusual in that the anterior pore may either perforate genital plate G2 or G3. Scale bar is 1 mm; after McNamara (1995).

ambu­­lacrum; lateroanal fasciole with sharp bend below ­periproct. Spines: Short, uniform and densely packed, longer toward the oral side.

Color: In life, the sea urchin is dark purple, the bare test light purplish. Distribution: The species is known from the Red Sea and the Indian Ocean in shallow water (Fig. 3.243).

318 

 3.3 Superorder Atelostomata von Zittel, 1879

B

A

C

G

D

F

E

H

I

Fig. 3.242: (A, B) Schizaster doederleini. Test length 67 mm, Florida Keys, FL, USA. Photos courtesy of H. van Noordenburg. (A) Aboral side: the fasciole is strongly widened below the petals. (B) Side view. (C) Apical system. The anterior gonopores are conspicuously reduced. Scale bar is 2 mm. (D) The plastron is broad being bowed laterally. Scale bar is 6 mm. After Kier (1975). (E, F) Schizaster floridiensis. Length 40 mm, Florida Keys. Holotype. Photo courtesy of Doris J. Pawson. (E) Aboral side: the petaloid area is larger than in S. doederleini. (F) Profile. Sketch after photo in Kier & Grant (1965). (G–J) Schizaster gibberulus. Test length 19 mm; Ile de la Réunion, Indian Ocean. Rather young animal. (G) Aboral side: the well-developed petals are almost cruciform. The peripetalous fasciole is broad, closely surrounding the ends of the petals and indenting in the interambulacra. (H) Oral side: the anterior tubercles and their associated spines are the largest, serving for burrowing. (I) Profile: the lateroanal fasciole forms a sharp bend below the periproct.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

B

C

 319

D

E

Fig. 3.243: Schizaster rotundatus. (A–D) Holotype, length 26 mm. San Christóbal, Galapagos Islands. From photos in Döderlein (1906). (A) Aboral side. (B) Oral side. (C) Side view. (D) Posterior side. (E) Profile of paratype, test length 16.5 mm.

Schizaster obovatus Lindley, 2004 Test: Rounded with shallow anterior notch; narrow; not much inflated; the holotype, a naked test, is only 34 mm long. Apical system: Slightly posterior. Anterior ambulacrum: Shallow, with pore pairs arranged in single rows. Oral side: Convex, peristome oval and sunken; periproct nearly reaching aboral side; labrum thick, as long as broad, with several small tubercles; plastron pear-shaped. Fascioles: Peripetalous fasciole distinct; lateroanal fasciole narrower than peripetalous one. Occurrence: Lindley has found it in the vicinity of Cap Gazelle in New Britain, Papua New Guinea. Schizaster rotundatus Döderlein, 1906 Main characteristics of the genus: Test: Thin; broadly oval with anterior notch; upper side slightly arched, posterior side vertically. Labrum: Broad, projecting; narrow naked areas in posterior ambulacra. Fascioles: Margino-peripetalous fasciole conspicuously indented in both lateral interambulacra; lateroanal branch thin, but distinct. Occurrence: Two specimens of 26 and 16.5 mm are recorded from the Galapagos Islands. Genus Tripylaster Mortensen, 1907 This monospecific genus is closely related to Brisaster, with both having the same plate architecture, a lowly arched test, and an adapically shallow anterior ambulacrum. The apical disc in Tripylaster is positioned more subcentrally than posteriorly as in Brisaster. Consequently, the posterior petals are more divergent in Brisaster. This

distinguishes it from B. moseleyi, which is known from the same geographical range. Tripylaster philippii (Gray, 1851) Test: Lowly arched, posterior side vertically truncated; in outline subcircular to oval with deep anterior notch and truncated to concave posterior end; length may reach 78 mm. Apical system: Ethmolytic, three gonopores, subcentrally positioned. Anterior ambulacrum: Shallow adapically, deepening toward the anterior, notched ambitus, pore pairs uniserially arranged, podia with large, penicillate disc. Paired petals: Depressed, pore zones hardly flexed, posterior pair half the length of anterior pair. Oral side: Flat, peristome very anterior, facing forward into anterior furrow; labrum conspicuously projecting; periproct at the truncated, slightly concave posterior side; labral plate short and wide, not extending beyond first adjoining ambulacral plates; sternal plates long and symmetrical. Fascioles: Marginal fasciole complete, lateroanal branch running straight along the sides of the test, strongly bent below the anal area, surrounding three to four subanal podia to each side; peripetalous fasciole well indented laterally. Tuberculation: Aborally fine, dense and uniform, slightly larger tubercles along anterior ambulacrum, oral tuberculation larger and less dense, plastronal spines with spoonlike tips. Distribution: This species is recorded from Southern South America, at the east coast northward up to the Rio de la Plata and at the west coast up to Calbuco, Chile. Bathymetric range: from the littoral to a depth of 340 m (Fig. 3.244).

320 

A

 3.3 Superorder Atelostomata von Zittel, 1879

B

C

D

E

F

Fig. 3.244: Tripylaster philippii. (A–C) Test length 49 mm, Patagonian Shelf, Argentina. ZMH. (A) Aboral side: the petals are only very faintly flexed. (B) Oral side. (C) Side view: the aboral side is gently arched. (D–F) Specimen with spines. Test length 25 mm, Falkland Islands. ZMUC. (D) Aboral side: in young specimens, the posterior petals may be relatively shorter than in larger individuals. (E) Side view: the long plastronal spines end in spoon-like tips. (F) Test length 57 mm, Patagonia, Argentina. ZMUC. Aboral side.

3.3.2 Order Spatangoida L. Agassiz, 1840 



Paleopneustina incertae sedis A Genus Schizocosmus Markov, 1990 The phylogenetic place is temporary, it is a combination of the schizasterids and the paleopneustid echinoids. The genus was placed in Pericomus (H.L. Clark 1925), in Tripylus with subgenus Parapneustes (Mortensen 1951), in Parapneustes (de Ridder et al. 1992), and in a new genus Schizocosmus (Mironov 2001). It is monospecific with its development, nonpelagic, and brood-protecting. The genus is characterized by having a more or less complete marginal fasciole with lateroanal branch. The peripetalous branch, which mostly coalesces with the former behind the anterior petals, may continue in some species as a separate branch toward the anterior side. The variation in the course of the fascioles is high (Fig. 3.245). Schizocosmus abatoides (H.L. Clark, 1925) Test: Broadly ovoid, frontal notch very faint; slightly to moderately elevated; posterior end truncated; length reaches usually 40 mm.

B

A

 321

Apical system: Subcentral to anterior, ethmolytic with three gonopores. Anterior ambulacrum: Narrow with uniserial pore pairs; podia not strongly developed. Paired petals: Of subequal length, parallel-sided, deeply sunken in females, less so in males. Oral side: Peristome kidney-shaped, facing forward into anterior depression, labrum projecting; labral plate short, extending only to first or third adjacent ambulacral plates; sternal plates long; extending to the seventh or eighth adjoining ambulacral plates; plate pattern variable. Fascioles: Marginal fasciole crossing the anterior ambulacrum ambital or just below the ambitus; well-developed, but thin, complete lateroanal branch; peripetalous branch distinct, sometimes continuing onto the anterolateral interambulacra, joining the marginal fasciole or not. Color: Light brown. Distribution: This species is known from the South Sandwich, South Shetland, and South Orkney islands, off Coats Land, and in the Ross Sea. Bathymetric range 64 to 1,120 m.

C

D

E

F

G

Fig. 3.245: Schizocosmus abatoides. (A) Male, test length 42 mm. The petals are slightly depressed. (B) Female, length 39 mm. Weddell Sea. AWI. In the female, the petals are transformed into deeply sunken marsupia. (C) Specimen with spines. Test length 28 mm, South Shetland Islands. AWI. (C) Side view: the peripetalous fasciole continues anteriorly, not coalescing with the marginal fasciole. (D) Offspring, found in the female. There are three well-defined stages of development: (1) eggs, (2) “short-spined” juveniles, and (3) “long-spined” juveniles that are almost ready to leave the brood pouch. (E–G) Plate pattern: (E) Aboral side of female. (F) Oral plate pattern. (G) Aboral side of male. The course of the fascioles is very variable: the well-developed peripetalous branch may coalesce with the marginal fasciole behind the anterior petals or in front of them, or it may pass parallel and well separated toward the anterior, but never crossing the anterior ambulacrum. Scale bar is 10 mm. (E) and (F) modified after Madon-Senez (1998). (G) Modified after De Ridder et al. (1992).

322 

 3.3 Superorder Atelostomata von Zittel, 1879

Remarks: There is a great deal of variation in the course of the fasciolar bands.

Palaeopneustina incertae sedis B In this clade, the marginal fasciole, which runs around the test, and the peripetalous fasciole, which crosses the aboral side like a bow, are coalesced anteriorly. The posterior part of the marginal fasciole passing below the periproct forms the lateroanal branch, which may be lost in some genera. The remaining fasciole appears to be a single one, but actually it is composed of the peripetalous bow and the anterior branch of the marginal fasciole. In The Echinoid Directory, A.B. Smith separated several genera into still unnamed groups, including the Antarctic brood-protecting genera Amphipneustes, Brachysternaster, Genicopatagus, and Parapneustes. However, using RNA analysis, Féral et al. (1994) stated that Abatus and the four above named genera are monophyletic. These brooding species are concentrated around Antarctica in the Cidaridae and the Paleopneustina. The genera are closely related, despite their morphological differences, and they share a similar evolutionary scenario. Almost all Antarctic or Subantarctic irregulars are brood-protecting, the young do not run through a pelagic larval stage with following metamorphosis, but develop from large yolk-rich eggs directly into small juvenile. The females have evolved deep brood pouches (marsupia) on the upper side, where the young grow, sheltered below the covering spines. Their development is nonpelagic. In contrast to the cycle of Abatus cordatus from the Kerguelen Islands, the members of high-Antarctic waters are likely to reproduce throughout the year. In the pouches of a single female, several stages of juveniles are found together. In some species, e.g., Brachysternaster chesheri, three to four well-defined stages are recognizable: round and irregularly formed eggs, followed by short-spined and long-spined juveniles. In other species, the development seems to be more or less continuous. That means they are spawning either several times a year or continually. Perhaps the cycle of development may last even longer than at the Kerguelen Islands because the water temperature is lower (Pearse & McClintock 1990). The fact that most of the echinoids live in deep water, as well as the conditions in the Antarctic winter, has made systematic research of these urchins rather difficult. Literature: Mortensen 1910; Koehler 1926; Bernasconi 1969; 1973; Pearse & McClintock 1990; Lockhart et al. 1994; Madon-Senez 1998; 2002; David et al. 2000 CDRom; David

et al. 2005 with a comprehensive bibliography; Pearse et al. 2008 (comprehensive bibliography) (Fig. 3.246). Five genera are included in the group: Amphipneustes Koehler, 1900. Antarctic and Subantarctic. Nine species, brood-protecting. Brachysternaster Larrain, 1985. Antarctic and Subantarctic. One species, brood-protecting. Genicopatagus A. Agassiz, 1879. Antarctic. Two species, brood-protecting. Heterobrissus Manzoni & Mazzetti, 1878. Indo-West Pacific. Five species, not brood-protecting. Parapneustes Koehler, 1912. Antarctic and Subantarctic. Two species, brood-protecting. Genus Amphipneustes Koehler, 1900 All species of the genus show nonpelagic development, they are brood-protecting their young in the sunken ambulacral pouches. The genus, exclusively distributed in Antarctic waters, in the islands south of the Atlantic and around the continent itself, is usually characterized by the total loss of fascioles. In a few species, a marginal fasciole is developed in juveniles. Test: More or less oval or ovoid, no anterior notch, variably high, reaching a medium to large size. Apical system: Subcentral, ethmolytic; three gonopores, larger in females than in males. Paired petals: Deeply depressed in females, forming marsupia; faintly sunken in males. Oral side: Peristome kidney-shaped; periproct marginal or inframarginal; labrum prominent, partly covering the peristome. Fascioles: Usually lacking in adults. The genus consists of nine species. Amphipneustes bifidus Mortensen, 1950. Antarctic coastal region. Amphipneustes brevisternalis Koehler, 1926. Eastern Antarctic Coast. Amphipneustes davidi Madon-Senez, 2002. Princess Ragnhild Coast, Antarctica. Amphipneustes koehleri Mortensen, 1905. South Georgia Islands, Subantarctic. Amphipneustes lorioli Koehler, 1901. Circum-Antarctic. Amphipneustes marsupialis Koehler, 1926. South Orkney Islands to Terre Adélie, Antarctica. Amphipneustes mironovi Markov, 1991. South Georgia and South Shetland islands, Antarctica. Amphipneustes rostratus Koehler, 1926. Antarctic Peninsula to King George V Coast. Amphipneustes similis Mortensen, 1936. Circum-Antarctic. The architectural variability of the species is conspicuously high, so they are often difficult to distinguish.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

D

B

 323

C

E

Fig. 3.246: (A–C) Development from the yolk-rich egg to the short-spined embryo. (A) The round eggs are filled with yolk (Brachysternaster). (B) The buds of the spines and podia have already appeared, when the embryonic epidermis opens and the echinid embryo comes out. It is approximately half-way through its development. On the yellow test, the primary (= terminal) podia are visible near the edge of the membrane. (Amphipneustes). (C) The upper short-spined juvenile shows the oral side, the future mouth in the center, surrounded by the developing spines and podia. The lower specimen is seen in profile and shows the protruding five primary podia around the naked upper side. (Brachysternaster). (AWI). (D, E) Long-spined juveniles. Many spines, podia, and pedicellariae have appeared. (D) Aboral side: in the center lies the apical system with the madreporite, which has only one pore at this stage. It is surrounded by the five primary (= terminal) podia. To the posterior, the periproct is still connected to the apical system. Only later in ontogeny will it shift to the posterior or oral side. (Amphipneustes). (E) Oral side: the peristome in the center is still closed. It is surrounded by the (darker) podia, whose finger-like digits for food-gathering have already begun to develop. This juvenile is almost ready to leave the mother and to live independently. (Brachysternaster) (AWI). All scale bars are 2 mm.

324 

 3.3 Superorder Atelostomata von Zittel, 1879

Amphipneustes bifidus Mortensen, 1950 Main characteristics of the genus: length may reach 73 mm. Test: Regularly ovoid, no frontal notch; aboral side regularly domed, with a vertically truncated posterior end, oral side slightly concave. Paired petals: Short, with approximately 12 pore pairs in each column. Oral side: Labrum protruding over the peristome, extending between the second and fifth adjoining ambulacral plates; sternal plates may reach the ninth to twelfth ambulacral plates. Spines: Spines of oral and ambital test terminating in two glassy peaks (bifide). Distribution: A. bifidus is obviously restricted to the Antarctic coastal region, along Coats Land, Princess Martha Coast, Enderby Land, and Mac-Robertson Coast; depth range from 190 to 623 m.

Distribution: This species is known only from the Subantarctic South Georgia Islands; 75 to 204 m.

Amphipneustes brevisternalis Koehler, 1926 Main characteristics of the genus: length usually 60 mm, maximum 70 mm. Test: Ovoid, aboral side hemispherical, posterior end with marginal truncation. Oral side: Periproct on the marginal truncation, fully visible from below; labrum of moderate size, reaching fourth adjacent plates; sternal plates short, as broad as long, plates extend to tenth adjoining ambulacral plates, for those are low. Distribution: This species is known from the Eastern Antarctic Coast, from Enderby Land over Adélie Coast to the Ross Sea at depths of 190 to 832 m (Fig. 3.247).

Amphipneustes marsupialis Koehler, 1926 Main characteristics of the genus: length usually 40 mm, may reach 60 mm. Test: Usually low, in outline slightly angular, anteriorly rounded, posteriorly attenuated; brood pouches in females large, extending almost to the margin. Oral side: Labrum long and narrow, extending to the second or third adjacent ambulacral plates; sternal plates elongated and narrow, reaching to the sixth adjoining ambulacral plates. Fasciole: Distinct marginal fasciole in juveniles with a length of up to 3 mm. Distribution: This species is found off South Orkney Islands, in the Weddell Sea, and along the coast of eastern Antarctica to Terre Adélie; depth 219 to 898 m (Fig. 3.249).

Amphipneustes davidi Madon-Senez, 2002 Main characteristics of the genus: length of holotype 57 mm. Test: Low, anterior side rounded without frontal notch, attenuated toward the posterior. Paired petals: Short, brood pouches in females end well above the margin. Oral side: Periproct low on the obliquely truncated posterior side, visible from above; labrum extends to third or fourth adjoining ambulacral plates; sternal plates very long and very broad with almost concave sides, forming a conspicuous triangle. Distribution: This species is only known from Princess Ragnhild Coast; depth range 150 to 216 m. Amphipneustes koehleri Mortensen, 1905 Main characteristics of the genus: maximum length 40 mm. Test: In outline broadly oval, anterior side straight and vertically truncated. Oral side: Periproct at vertical posterior end; labrum short, extending to the first or second ambulacral plates; sternal plates long and broad, laterally bowed.

Amphipneustes lorioli Koehler, 1901 Main characteristics of the genus: maximum length 60 mm. Test: Regularly ovoid, posterior end narrower and not truncated, aboral side high and subconical. Oral side: Periproct almost inframarginal, fully visible from below; labrum broad and stout, extending to the third and fourth adjacent ambulacral plates; sternal plates short, posteriorly slightly indented by flexed ambulacra. Color: In living specimens, the test and spines are violet to dark purple. Distribution: This species is found around the entire Antarctic continent, from South Shetland, Weddell Sea, MacRobertson Coast to the Ross and Bellingshausen Seas; 70 to 2,012 m (Fig. 3.248).

Amphipneustes mironovi Markov, 1991 Test very thin and fragile, known only from fragments. Faint anterior notch; periproct supramarginal at the posterior side; labrum very large and slightly elevated, extending to the second or third adjacent ambulacral plates. Distribution: Reported from off the South Georgia and South Shetland Islands, at depths of 3,060 and 4,704 m (Fig. 3.250). Amphipneustes rostratus Koehler, 1926 Main characteristics of the genus: length may reach 60 mm. Test: Aborally high and conical; posterior interambulacrum keeled; in outline anteriorly rounded, strongly attenuated toward the posterior truncated end; brood pouches in females relatively short. Oral side: Flattened, peristome deeply depressed; periproct on the truncated posterior face, visible from below; labrum large, extending to the second adjacent

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

 325

B

D

C

F

E

G H

Fig. 3.247: (A–C) Amphipneustes bifidus, male. Test length 42 mm; Weddell Sea, Antarctica. AWI. (A) Aboral side. (B) Oral side. (C) The spines end distally in two glassy (hyaline) peaks. The shaft is finely ridged. Scale bar is 1 mm. (D, E) Amphipneustes brevisternalis, female. (D) Aboral side. (E) Oral plate pattern. Scale bar is 10 mm. Modified after Madon-Senez (1998). (F, G) Amphipneustes davidi, female. (F) Aboral side. (G) Oral plate pattern. Scale bar is 10 mm. Modified after Madon-Senez (2002). (H) Amphipneustes koehleri. Oral plate pattern. Scale bar is 10 mm. Modified after Madon-Senez (1998).

ambulacral plates; sternal plates very long, reaching the seventh and eighth ambulacral plates. Fasciole: Marginal fasciole passing below the periproct and often well-developed in juveniles up to a length of 29 mm; lost in adult specimens.

Color: Light brown to yellow-brown. Distribution: A. rostratus is found off the northern tip of the Antarctic Peninsula, and along the coast of Antarctica from Coats Land to King George V Land; depth 164 to 1,353 m (Fig. 3.251).

326 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

C

D

G

E

F

H

Fig. 3.248: Amphipneustes lorioli. (A–C) Female, test length 59 mm; Weddell Sea, Antarctica. AWI. (A) Aboral side: the deep marsupia are overarched by flattened paddle-shaped spines. Below this dense cover numerous fine spines are developed, in which the most advanced, “long-spined” juveniles cling with the thorns of their own spines. On the ground of the pouches the smaller “short-spined” juveniles and the eggs are densely packed. (B) Oral side: the periproct lies inframarginal. (C) Side view: the test is high and conical. (D) Long-spined juveniles, almost ready to leave the mother. Oral side (left) with the growing phyllodes; (right) aboral side. Scale bar is 5 mm. (E, F) Male specimen, test length 63 mm, Weddell Sea. AWI. (E) Aboral side: the petals are narrow and only slightly depressed. (F) Oral side: the lip greatly protrudes over the peristome. (G) Oral plate pattern. Scale bar is 10 mm. Modified after Madon-Senez (1998).

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

D

 327

B

C

E

Fig. 3.249: Amphipneustes marsupialis. (A–C) Female, test length 51 mm, Weddell Sea, Antarctica. AWI. (A) Aboral side: the marsupia are broad and long. (B) Oral side. (C) Side view. (D) Male specimen, length 47 mm, Weddell Sea, Antarctica. AWI. Aboral side: the petals are very long. (E) Long-spined juvenile. ZMH. It still has a fasciole, which becomes lost in larger specimens. Scale bar is 2 mm.

328 

 3.3 Superorder Atelostomata von Zittel, 1879

B

A

C

D

E

Fig. 3.250: Amphipneustes rostratus. (A–C) Male. Test length 40 mm; Weddell Sea. AWI. (A) Aboral side. In males, the petals are slightly sunken. (B) Oral side: the plastron is narrow. The whole oral side was made free of spines. (C) Side view: the test is highly vaulted. (D, E) Specimen with spines, female. Test length 41 mm; Weddell Sea. AWI. (D) Aboral side: the spines adjacent to the petals are long, densely covering the deeply sunken marsupia. (E) Oral side.

3.3.2 Order Spatangoida L. Agassiz, 1840 



B

A

D

E

 329

C

F

H G

Fig. 3.251: (A–D) Amphipneustes rostratus, female. Test length 48 mm, Weddell Sea, Antarctica. AWI. Aboral side: the marsupia are short. (B) Aboral side of juvenile: the periproct is positioned in the center, the podia in the anterior ambulacrum III (above) are already welldeveloped. In the fasciole, tiny tubercles are densely spaced. Scale bar is 10 mm. (C) Offspring in three well defined stages: eggs, shortspined, and long-spined embryos. Scale bar is 2 mm. AWI. (D) Oral plate pattern. Scale bar is 10 mm; modified after Madon-Senez (1998). (E–H) Amphipneustes similis, female. Test length 58 mm, Weddell Sea, Antarctica. AWI. (A) Aboral side. (B) Oral side. (C) Side view. The large black pedicellariae are typical. (D) Oral plate pattern. Scale bar is 10 mm. Modified after Madon-Senez (1998).

Amphipneustes similis Mortensen, 1936 Main characteristics of the genus: length may reach up to 70 mm. Test: Regularly ovoid with rounded anterior side, aboral side hemispherical; marsupia in females very deep. Oral side: Peristome sunken, periproct on slightly truncated posterior face, visible from below; labrum broad, extending to third adjacent ambulacral plates; sternal plates short and stout. Spination: Not very dense. Distribution: This species is likely to be restricted to the Antarctic Peninsula and the Weddell Sea from South

Shetland and South Orkney islands to Coats Land; depth range 20 to 898 m. Genus Brachysternaster Larrain, 1985 The genus is monospecific and restricted to Antarctic and Subantarctic waters. The development is nonpelagic and brood-protecting (Fig. 3.252). Brachysternaster chesheri Larrain, 1985 Test: Rather low, vertex at the apical system, no posterior truncation; in outline elongated ovoid; anterior side

330 

 3.3 Superorder Atelostomata von Zittel, 1879

convex, narrowing toward the posterior; length may reach 77 mm. Apical system: Ethmolytic with three gonopores, positioned subcentrally. Anterior ambulacrum: Flush with specialized podia. Paired petals: Sunken in females, forming deep marsupia, only slightly sunken in males. Oral side: Somewhat concave; peristome reniform, hardly visible below the protruding labrum; periproct inframarginal, visible from below; labral plate broad, pointed toward the posterior, extending at least to the third adjacent ambulacral plates; separated from the sternal plates by one or two pairs of ambulacral plates that join at the midline; sternal plates very short and broadly triangular. Fascioles: No trace. Tubercles: Sparse; spines relatively long. Color: Test and spines usually purple, more rarely yellowish. Distribution: This species is known only in the Weddell Sea, at the northwest side off the Antarctic Peninsula, South Shetland and South Orkney islands, at the east side along Coats Land and Princess Ragnhild Coast. Bathymetric range 115 to 1,353 m; mostly recorded from depths of 200 to 500 m (Fig. 3.253). Genus Genicopatagus A. Agassiz, 1879 The genus contains two species, living exclusively in Antarctic deep waters. Delopatagus is made a synonym for Genicopatagus by Mironov (1993). The development is nonpelagic and brood-protecting. Genicopatagus affinis A. Agassiz, 1879 Test: Very thin and fragile; in outline broadly oval or subcircular, no anterior notch; aboral side raised, vertex posterior, posterior slope steeper than anterior; length may reach 77.3 mm. Apical system: Eccentric to the posterior, ethmolytic with three gonopores, much larger in females than in males; a specimen of 41 mm length has not yet developed gonopores, so it is likely the species reaches a larger size. Anterior ambulacra: Narrow; flush, adapically faintly sunken; pore pairs distinct. Paired ambulacra: Also narrow, subpetaloid with double pores, deeply sunken in females forming marsupia, but not depressed in males. Oral side: Flat, peristome subcentral, sunken; overarched by labrum; phyllodes well-developed; periproct supramarginal, high on the steeply sloping posterior side; labrum long and stout, extending to third to fifth adjoining ambulacral plates; sternal plates reaching the fifth to ninth adjacent ambulacral plates. Tuberculation: Fine and uniform, not dense. Color: Test and spines are variably violet to yellowishgreen.

A

B

C

D

Fig. 3.252: (A) Brachysternaster chesheri. Pedicellariae at the apical system. There are tridentate and bidentate pedicellariae of various sizes the globiferous (arrow) having a very long stem. Scale bar is 10 mm. (B, C) Juveniles found in the marsupia of a female, test length 51 mm. (AWI). There are three well-defined stages: round eggs (above), embryos with spines just evolving (in the middle) and long-spined juveniles (below), much larger and almost ready to leave the mother. Scale bar is 2 mm. (C) Offspring from the female, test length 46 mm. Eggs and short-spined embryos. Scale bar is 10 mm. (D) Oral plate pattern: the ambulacral plates touch each other. Scale bar is 20 mm.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

 331

B

C

D

E

F

Fig. 3.253: Brachysternaster chesheri. (A–C) Female. Length 46 mm; Weddell Sea. AWI. (A) Aboral side: the offspring in the still shallow marsupia was densely packed. (B) Oral side: the ambulacral plates, joining at the midline, form a large naked area. (C) Side view: the labrum is very prominent, the plastron being concave. (D) Female specimen with spines; length 63 mm; Weddell Sea. AWI. Aboral side: the juveniles are kept in the large marsupia. (E) Female with spines; length 55 mm; South Shetland Islands. ZMH. (E) Aboral side: the primary spines are long and coarse. (F) Male specimen; length 61 mm; Weddell Sea. AWI. Aboral side: the petals are hardly depressed.

332 

 3.3 Superorder Atelostomata von Zittel, 1879

Distribution: This species is recorded from the South Shetlands Islands and off Coats Land in the Weddell Sea, in the South Atlantic, and in the southern Indian Ocean at depths of 621 to 5,651 m (Fig. 3.254). Genus Delopatagus Koehler, 1907 Mironov (1993) already suggested that Delopatagus and Genicopatagus might be synonymous. The author

A

considered this possibility after having observed a wide range of variation in the 12 specimens of Genicopatagus collected by the Polarstern 1998 and 2000. However, more material must be examined, particularly from Delopatagus brucei, from which only fragmentary specimens are available. Further molecular data are necessary. In any case, the two species are closely related, and both are close to Amphipneustes. The differences between Delopatagus and

B

C

D

E

F

Fig. 3.254: Genicopatagus affinis. (A–C) Male, test length 48 mm, Scotia Sea. AWI. (A) Aboral side: the paired ambulacra are subpetaloid, the anterior ambulacrum being flush with the test. (B) Oral side: the peristome is subcentral, surrounded by well-developed phyllodes. (C) Side view: the posterior slope is much steeper than the anterior. (D, E) Specimen without spines, male. Length 61 mm, Weddell Sea, Antarctic. ZMH. (D) Aboral side: the paired ambulacra are subpetaloid. (E) Oral side. (F) Female specimen, length 59 mm, Weddell Sea, Antarctica. (F) Aboral side: the marsupia are still short and relatively shallow.

3.3.2 Order Spatangoida L. Agassiz, 1840 



Genicopatagus are gradual, particularly the petals are difficult to define. Koehler called them “slightly petaloid” in D. brucei, in Genicopatagus affinis they are said to be “subpetaloid”. Delopatagus brucei Koehler, 1907 Test: Fragile, in outline subcircular, no anterior notch; aboral side raised, posterior slope much steeper than the anterior; holotype measures 41 mm in length. Apical system: Positioned conspicuously to the posterior; ethmolytic, three gonopores. Anterior ambulacrum: Flush, pores rudimentary.

 333

Paired petals: Weakly petaloid, forming deep marsupia in females, flush in males, posterior pair much shorter than anterior pair. Oral side: Flattened; peristome slightly anterior, somewhat sunken below projecting labrum; periproct high on the steep posterior face, clearly visible from above. Fascioles: No trace. Color: Test and spines in living specimens are grayyellowish. Distribution: This species is seldom collected, known from off Fireland, South Shetland Islands, and Coats Land; bathymetric range 1,996 to 4,462 m (Fig. 3.255).

A

B

C

D

E

F

G

Fig. 3.255: (A–C) Genicopatagus affinis. (A, B) From Agassiz (1904), no scale. (A) Aboral side. (B) Oral side. (C) Oral plate pattern. (D–G) Delopatagus brucei, male. Test length 41 mm, south of the South Orkney Islands. From Koehler (1907). (D) Aboral side: the apical disc is strongly eccentric toward the posterior. (E) Oral side. (F) Side view: the posterior side is very steep, toward the anterior the test slopes gently. (G) Posterior side: the periproct is positioned high on the aboral side.

334 

 3.3 Superorder Atelostomata von Zittel, 1879

Genus Heterobrissus Manzoni & Mazzetti, 1877 This genus is characterized by a very coarse aboral tuberculation, the total lack of fascioles and by the ambulacra on the oral side being fully tuberculated beyond the phyllodes. It is a tropical echinoid and not brood-protecting. Test: Regularly oval to ovoid, no anterior notch; aboral side more or less highly domed or subconical, rounded margin. Apical system: Subcentral, ethmolytic with four gonopores. Anterior ambulacrum: Narrow, flush, with very small pore pairs. Paired petals: Subpetaloid; parallel series of enlarged pore pairs expand almost to the margin. Oral side: Flattened, peristome slightly anterior; deeply sunken, facing forward into a groove, overarched by the labrum; phyllodes strongly developed; periproct circular and flush, positioned inframarginally on oblique posterior end; labral plate short and broad, extending to the second or third adjacent ambulacral plates; sternum symmetrical. Tubercles: Aboral side more or less sparsely set with coarse, heterogeneous tubercles, crenulate and perforate; more densely packed on oral side, ambulacra fully tuberculated beyond phyllodes. Spines: Aborally solid, long, curved at the base and sharply pointed distally. Literature: Döderlein 1901; Koehler 1914; Baker & Rowe 1990; van Noordenburg 2008. Five species are included in the genus: Heterobrissus erinaceus Baker & Rowe, 1990. East Aus­tralia. Heterobrissus gigas Baker & Rowe, 1990. Southeast Australia. Heterobrissus hemingi (Anderson, 1902). Laccadive Sea, Indian Ocean. Heterobrissus hystrix (A. Agassiz, 1880). Caribbean. Heterobrissus niasicus (Döderlein, 1901). East Africa, Malayan Archipelago to Philippines and Southeast Australia. Heterobrissus erinaceus Baker & Rowe, 1990 Main characteristics of the genus: test length of holotype 140.4 mm; width 74.1 mm. Test: Steeply arched; ambitus abruptly curved; upper side conical. Apical system: Posterior; four gonopores, ethmolytic. Anterior ambulacrum: Frontal pores slit-like. Paired petals: Narrowly divergent, anterior pair almost reaching ambitus. Oral side: Peristome partly obscured by projecting labrum; circular periproct at the truncated end; labrum pointed, longer than wide; labrum and sternum fully tuberculated.

Tuberculation: Small primary tubercles, crowded and smaller at ambitus; spines sharply pointed. Color: Test muddy brown, spines mauve to lavender. Distribution: Restricted to the East of Australia, from southern New South Wales to southern Queensland at a depth of 183 to 421 m. Heterobrissus gigas Baker & Rowe, 1990 Main characteristics of the genus: test length of holotype is 146 mm, it may reach 165 mm. It is very much like Heterobrissus niasicus, but larger and with distinctly different pedicellariae. Test: Very large, ambitus rounded. Oral side: Periproct vertically oval; labrum curved, wider than long. Tuberculation: Coarse. Distribution: Southeastern coast of Australia from Cape Hawke, New South Wales, to Cape Everard, Victoria. Bathymetric range 128 to 503 m. Heterobrissus hemingi (Anderson, 1902) Main characteristics of the genus: maximum length reaches more than 115 mm. This species differs from H. niasicus by its test being higher and more conical, the oral side being flat, not depressed; by its narrower petals, and shorter spines being more densely distributed. Distribution: This species was collected in the Laccadive Sea in enormous quantities (“filling six buckets”); depth 390 to 510 m (Fig. 3.256). Heterobrissus hystrix (A. Agassiz, 1880) Main characteristics of genus: length may reach 142 mm. Test: Low conical, in outline elliptical, attenuated toward the posterior, anterior side convex. Paired petals: Extend approximately three-fourths of the distance toward the margin, pore zones set with small secondary tubercles, widely open distally. Oral side: Flattened, deeply sunken toward the peristome, phyllodes depressed, appearing conspicuously naked, for the ambulacra beyond are densely tuberculated; periproct inframarginal at the obliquely truncated posterior end. Primary tubercles: Aborally scarce within a mass of miliaries, oral tuberculation more densely set. Spines: Coarse, up to 40 mm, sharply pointed. Color: In living specimens, the color of the test and spines are very variable, from chocolate brown with yellow-gray spines, to greenish-purple, Indian-red or dark-purplish. The denuded test is whitish with a pink tint.

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

D

B

 335

C

E F

Fig. 3.256: (A–C) Heterobrissus erinacea. Test length 140.4 mm, holotype, east Australia. After photos in Baker & Rowe (1990). (A) Aboral side: the test is almost round. (B) Oral side: the large peristome is anterior. (C) Side view. The test is rather low. (D–F) Heterobrissus hemingi. Test length 99 mm, Laccadive Sea. From Koehler (1914). (D, E) Aboral and oral side: the oral ambulacra are fully tuberculated beyond the distinct phyllodes. (F) Side view: the test is high and conical.

Distribution: Recorded only from the West Indies at depths of 38 to 500 m (Fig. 3.257). Heterobrissus niasicus (Döderlein, 1901) Test: Conically raised, in outline regularly oval; length reaches more than 140 mm. Apical system: Subcentral, four gonopores; ethmolytic. Anterior ambulacrum: Flush, minuscule pore pairs, no specialized tube feet. Paired ambulacra: Also flush; nonpetaloid, rows of adapical pore pairs rudimentary. Oral side: Flattened, peristome deeply sunken, overhung by the prominent labrum; phyllodes strongly developed and depressed, periproct large, on the short, vertically

truncated posterior end; plastron and adjacent ambulacra fully tuberculate. Fascioles: None. Spination: Very coarse spines scattered over the aboral surface, finer and more densely packed on the oral side. Distribution: This species is reported from deep water in the Indian Ocean from the East African coast to the Malayan Archipelago, the Philippines, and southeast Australia (Fig. 3.258). Genus Parapneustes Koehler, 1912 The genus is nonpelagic and brood-protecting. It contains two species, which are very similar. Both live in the Antarctic and Subantarctic.

336 

 3.3 Superorder Atelostomata von Zittel, 1879

A

B

C

D

E

F

G

Fig. 3.257: (A–C) Heterobrissus hystrix. Test length 131 mm; Virgin Island, West Indies. ZMUC. (A) Aboral side: the petals are parallel-sided. (B) Oral side. (C) Side view: the test is lowly conical. (D, E) Specimen with spines; test length 142 mm; Virgin Island, West Indies. ZMUC. (D) Aboral side. The spines are coarse and sharply pointed. (E) Oral side. (F) Apical system. Scale bar is 4 mm. (G) Labrum and adjacent plates: the ambulacral plates beyond the phyllodes are fully tuberculated like the labral and sternal plates. Scale bar is 10 mm. After Mortensen (1950).

3.3.2 Order Spatangoida L. Agassiz, 1840 



A

D

B

 337

C

E

F

Fig. 3.258: (A–C) Heterobrissus niasicus. (A, B) Test length 115 mm; southeast of Sumatra, Indonesia. ZMH. (A) Aboral side: the spines are very coarse and bowed near the base. (B) Oral side: the spines are much smaller and finer than on the aboral side. (C) Test length 94 mm; Philippines. Photo courtesy of H. van Noordenburg. Aboral side: Only a few, large primary tubercles are scattered over the upper surface. (D–F) Specimen without spines; length of test 134 mm; Mozambique, East Africa. (D) Aboral side: the large primary tubercles are sparsely distributed; the series of pore pairs do not reach the apical system. (E) Oral side: the peristome is deeply depressed, covered by the prominent lip. The strongly developed phyllodes lie in sunken, rather naked grooves, whereas the lateral ambulacra are tuberculated like the plastron. (F) Close-up of peristomial area: the phyllodes are strongly developed. The lip expands far across the peristome.

338 

 3.3 Superorder Atelostomata von Zittel, 1879

Parapneustes cordatus Koehler, 1912 Test: Elongated; outline somewhat angular, anterior side straight, distinctly narrowing toward the pointed posterior end; length 50 mm. Paired petals: Long, almost reaching the ambitus. Oral side: Labral plate extending to the second or third adjoining ambulacral plates; sternal plates long and narrow. Fascioles: Marginal and peripetalous fascioles thin, usually not wider than three miliaries, lateroanal branch sometimes reduced and hardly visible. Color: In life, the test is reddish-brown, the spines rather yellowish-green. Distribution: This species is recorded from the western part of Antarctica from the Ross Sea, the Bellingshausen Sea, and off South Shetland Islands. Depth range 160 to 592 m (Fig. 3.259).

Parapneustes reductus Koehler, 1912 Main characteristics of the genus: length may reach approximately 40 mm. Test: Ovoid, no frontal notch; posterior side rounded and narrower than anterior side. Oral side: Labral plate extending at least to second adjacent ambulacral plates; sternal plates long and narrow, slightly bowed laterally. Fascioles: Not more than three miliaries in width and sometimes interrupted; anterior branch passing lowly around the ambitus, peripetalous branch also rather low. Distribution: This species is only reported from Fireland and the South Shetland and South Orkney islands. Bathymetric range 50 to 761 m.

A

B

C

D

E

Fig. 3.259: (A–C) Parapneustes cordatus, male. Test length 50 mm; Palmer Archipelago, Antarctica. ZMUC. (A) Aboral side. The ambulacra are moderately depressed. (B) Oral side. (C) Side view: there is a complete marginal fasciole and a distinct peripetalous branch. (D, E) Parapneustes reductus, male. (D) Aboral side. (E) Oral plate pattern. Scale bar is 10 mm. Modified after Madon-Senez (1998), as Tripylus reductus.

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Acknowledgments Many people kindly supported me. Without their help, I would not have been able to write these books on echinoids. Thanks to Claus Nielsen and Tom Schiötte, Zoological Museum of Copenhagen, for making available to me specimens of the echinoid collection of Theodor Mortensen. I reveled in examining echinoids and photographing all I needed. Thanks to Carsten Lüter, Museum für Naturkunde, Berlin, for allowing me to take pictures of specimens housed in the collection, just before they were moved into the rooms of the newly built, state-of-the-art, wet collection. Thanks to Andreas Schmidt-Rhaesa and Peter Stiewe for making it possible to work in the echinoid collection of the Zoological Institute and Museum of the University of Hamburg. I felt at home in that basement. Thanks to Rich Mooi, California Academy of Sciences, San Francisco, for answering the many questions on the classification of echinoids at once. Many thanks to Andreas Schmidt-Rhaesa, in his role as chief editor of the Handbook of Zoology, for his help in bringing the text and figures into the right format. And for doing hard work when I was ill. I know it only today. Thanks for answering questions, for pictures, for specimens, for literature and for good advice to the following: –– Owen F. Anderson, National Institute of Water & Atmospheric Research Ltd., Wellington, New Zealand –– Andrew Cabrinovic and Harry Taylor, The National History Museum, London, UK –– Willi Clarenbach, Haan, Germany

DOI 10.1515/9783110368536-005

–– Simon E. Coppard, Hamilton College, New York, USA –– Bruno David, Biogéoscienes, Université de Bourgogne, Dijon, France –– Heidi Friedhoff, Norderstedt, Germany –– Takashi Ito, Shizuoka, Japan –– Roland and Renate Kraft, Groß-Grönau, Germany –– Andreas Kroh, Naturhistorisches Museum, Vienna, Austria –– Harileos A. Lessios, Smithsonian Tropical Research Institute, Balboa, Panama –– Ashley Miskelly, Sydney, Australia –– Christian Neumann, Museum für Naturkunde, Berlin, Germany –– Henk van Noordenburg, Leusden, The Netherlands –– Doris J. and David L. Pawson, National Museum of Natural History, Washington, DC, USA –– Bernhard Ruthensteiner, Zoologische Staatssammlungen, Munich, Germany –– Andrew B. Smith, The Natural History Museum, London, UK –– Bas van der Steld, Zoetermeer, The Netherlands –– Hans-Volker Thiel, Bonn, Germany And last but not least, thanks to my husband Peter, who accompanied me in this work for many years, encouraging me to go on and helping me with numerous “small” technical things that turned out to be so important. I am grateful to all of you, Heinke Schultz

Index abatoides, Schizocosmus 321 Abatus 4, 282, 283, 285, 322 Abatus agassizii 283, 284 Abatus beatriceae 283, 284, 285 Abatus bidens 283, 284, 285 Abatus cavernosus 147, 282, 283, 285, 286 Abatus cordatus 283, 285, 287, 322 Abatus curvidens 284, 285 Abatus elongatus 287, 288 Abatus ingens 284, 285, 288 Abatus koehleri 284, 287, 288 Abatus philippii 288, 289 Abatus shackletoni 284, 289, 290 Abatus (Pseudabatus) nimrodi 284, 289, 290 aberrans, Encope 102, 103, 105 abnormalis, Koehleraster 7, 8 Acanthotrema 194 Aceste 4, 282, 289, 291 Aceste bellidifera 289, 293 Aceste ovata 289, 291, 293 Aceste weberi 289, 291 aculeata, Argopatagus 260, 261 aculeata, Phrissocystis 260 acuta, Fibulariella 76, 77, 78 Aeropsis 4, 261 Aeropsidae 4, 261 Aeropsis fulva 262, 265 Aeropsis rostrata 262, 263, 265 affinis, Genicopatagus 330, 332, 333 affinis, Hypselaster 298, 300 affinis, Peronella 83 africanus (Plagiobrissus), Plagiobrissus 183, 185 Agassizia 4, 278, 279 Agassizia excentrica 279, 280 Agassizia scrobicutata 279, 280 agassizii, Abatus 283, 284 agassizii (Allobrissus), Brissus 167, 169, 170 akabanus, Pericosmus 273, 274 alcocki, Pourtalesia 131, 135 alexandri, Echinolampas 20, 21, 23 Allobrissus 167, 170 alta, Brissopsis 155, 157 altus, Nacospatangus 245, 246, 247 altus, Spatangus 253, 255, 256 Ammotrophus 3, 34, 36 Ammotrophus arachnoides 36, 39 Ammotrophus cyclius 37, 39 Ammotrophus platyterus 39, 40 Amphipneustes 5, 282, 322, 323, 332 Amphipneustes bifidus 322, 324, 325 Amphipneustes brevisternalis 322, 324, 325 Amphipneustes davidi 322, 324, 325 Amphipneustes koehleri 322, 324, 325 Amphipneustes lorioli 322, 324, 326 Amphipneustes marsupialis 322, 324, 327 Amphipneustes mironovi 322, 324 DOI 10.1515/9783110368536-006

Amphipneustes rostratus 322, 324, 325, 328, 329 Amphipneustes similis 322, 329 amphora, Echinosigra (Echinogutta) 130 amplificatus, Clypeaster 43, 45 Anabrissus  4, 151, 152 Anabrissus damesi 152, 153 analis, Peronella 83 Anametalia 4, 151, 152, 15, 170 Anametalia grandis 152, 153 Anametalia regularis 152, 154, 155 Anametalia sternaloides 152, 154, 155 angularis, Plethotaenia 253, 254 angulipora, Fibulariella 77, 78 angustus, Metalia 174, 176 annandalei, Clypeaster 43, 44, 45 Anochanus 3, 29, 30 Anochanus sinensis 30 antarctica, Echinosigra (Echinogutta) 130 antarcticus, Brisaster 291, 293 antepodeanus, Urechinus 145, 147 Antrechinus 4, 139 Antrechinus drygalskii 139 Antrechinus mortenseni 139, 142 Antrechinus nordenskjoldi 139, 142 aoteanus, Plexechinus 122, 123, 124 Apatopygids 11 Apatopygidae 3, 9 Apatopygus 3, 10 Apatopygus occidentalis 10, 11 Apatopygus recens 10, 11, 30 Aphanopora 3, 29, 30 Aphanopora echinobrissoides 30, 31 apicatus, Echinocyamus 68, 69 Arachnoides 3, 39 Arachnoides placenta 37, 39, 40 Arachnoides tenuis 37, 39, 40 arachnoides, Ammotrophus 36, 39 Arachnoids 34, 36 Arachnoididae 3, 34 Araeolampas 4, 212, 215 Araeolampas atlantica 216, 217 Araeolampas fulva 216, 217 Araeolampas glauca 216, 217 Araeolampas hastata 216, 218 Araeolampas rostrata 216, 218 Argopatagus 4, 260 Argopatagus aculeata 260, 261 Argopatagus multispinus 260, 261 Argopatagus planus 260, 261 Argopatagus vitreus 260, 261, 262 Astriclypeid 96 Astriclypeidae 3, 67, 93, 95 Astriclypeus 3, 36, 95 Astriclypeus manni 95 Atelostomata 3, 7, 118, 145 atlantica, Araeolampas 216, 217

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 Index

atlantica, Brissopsis 157, 159, 162 atropos Moira 222, 302, 303, 304, 305, 307 auritus Sculpsitechinus 96 auritus, Echinodicus 35, 96, 97 aurorae, Pourtalesia 131, 135 australasiae, Breynia 218, 219, 220 australasiae, Clypeaster  35, 43, 44, 46, 63 australe cordatum, Echinocardium 222 australis, Mortonia  67, 73, 74 australis, Protenaster 314 barbatus, Diploporaster 297, 298 beatriceae, Abatus 283, 284, 285 bellidifera, Aceste 289, 293 belyaevi, Pilematechinus 141, 145 bengalensis, Brissopsis 157, 159, 161 beryl, Spatangus 253, 255, 260 bidens, Abatus 283, 284, 285 bidens, Pericosmus 273, 274 bifidus, Amphipneustes 322, 324, 325 bisperforatus, Echinodiscus 96, 97 boninensis, Laganum 79 borealis, Encope micropora 105, 107 brachypetalus, Hypselaster 298, 300 brachypetalus, Linopneustes 205, 208 Brachysternaster 5, 282, 322, 323, 329 Brachysternaster chesheri 322, 329, 330, 331 brevisternalis, Amphipneustes 322, 324, 325 Breynia 4, 212, 218, 219 Breynia australasiae 218, 219, 220 Breynia desorii 218, 219, 220 Breynia elegans 218, 219, 221 Breynia neanika 218, 219, 221 Breynia vredenburgi 218, 219 Brisaster 4, 282, 291, 308 Brisaster antarcticus 291, 293 Brisaster capensis 291, 294 Brisaster fragilis 291, 294 Brisaster kerguelensis 291, 293, 294 Brisaster latifrons 291, 293, 294 Brisaster moseleyi 291, 293, 296, 319 Brisaster owstoni 291, 293, 296 Brisaster tasmanicus 291, 293 Brisaster townsendi 291, 293, 297 Brissalius 4, 151, 155 Brissalius vannoordenburgi 155, 157 Brissid 151 Brissidae 4, 150, 151, 152, 170 Brissidea 150 Brissidina 4, 150 Brissopsis 4, 150, 151, 155, 157, 179 Brissopsis alta 155, 157 Brissopsis atlantica 157, 159, 162 Brissopsis bengalensis 157, 159, 161 Brissopsis columbaris 157, 159, 161 Brissopsis elongata 157, 159, 161 Brissopsis evanescens 157, 159, 161 Brissopsis jarlii 157, 159 Brissopsis luzonica 157, 159, 161, 167 Brissopsis lyrifera 157, 162, 163

Brissopsis lyrifera capensis 157, 162, 163 Brissopsis mediterranea 157, 159, 16, 164 Brissopsis micropetala 157, 162, 165 Brissopsis obliqua 157, 162, 165 Brissopsis oldhami 157, 162, 165 Brissopsis pacifica 157, 162, 166 Brissopsis parallela 157, 165, 168 Brissopsis persica 179 Brissopsis similis 157, 16, 168 Brissopsis zealandiae 157, 162, 165 Brissus 1, 4, 118, 118, 151, 152, 167 Brissus (Allobrissus) agassizii 167, 169, 170 Brissus gigas 167 Brissus latecarinatus 167, 169, 170 Brissus obesus 167, 170 Brissus unicolor 1, 167, 170, 171 brucei, Delopatagus 333 brykovi, Scaphechinus 115 cadenati, Meoma 171, 172, 173 californica, Encope micropora 105, 107 californicus, Spatangus 253, 255, 256 Calymne 3, 119, 120 Calymne relicta 119, 121 Calymnidae 3, 119 camarota, Lovenia 228, 231 canalifera, Ova 308, 309, 311, 312 capense, Echinocardium 222, 223 capensis lyrifera, Brissopsis 157, 162, 163 capensis, Brisaster 291, 294 capensis, Spatangus 253, 255, 256 Cardiasteridae 127 caribaearum, Cassidulus 10, 12, 14, 15, 17, 18 carinata, Helgocystis 131, 135 carinata, Maretia 242, 244 Carnarechinidae 3, 119, 122 Carnarechinus 3, 122 Carnarechinus clypeatus 122 Cassidulidae 3, 11 Cassiduloid 12, 13, 14, 15 Cassiduloida 3, 7, 9, 11, 34 Cassidulus 3, 10, 12 Cassidulus caribaearum 10, 12, 14, 15, 17, 18 Cassidulus delectus 12 Cassidulus infidus 12, 14 Cassidulus malayanus 12, 13 Cassidulus mitis 12, 14, 15 Cassis tuberosa (Gastropod) 63 cavernosus, Abatus 147, 282, 283, 285, 286 centrale, Laganum 79 Ceratophysa 3, 128 Ceratophysa ceratopyga 128, 129 ceratopyga, Ceratophysa 128, 129 challengeri, Spatagocystis 139, 141 chesheri, Brachysternaster 322, 329, 330, 331 chesheri, Clypeaster 43, 44, 46 chinensis, Faorina 269, 272 chuni, Echinolampas 20, 21, 23, 29 Cidaridae 283, 322 Cidaroidea 3

Index 

cinctus, Peripatagus 266, 268 cinctus, Plexechinus 122, 124, 125, 126 Cionobrissus 4, 152, 170 Cionobrissus revinctus 170, 172 claudicans, Moiropsis 308, 310 clotho, Moira 303, 306, 307 Clypeaster 3, 40, 63 Clypeaster amplificatus 43, 45 Clypeaster annandalei 43, 44, 45 Clypeaster australasiae 35, 43, 44, 46, 63 Clypeaster chesheri 43, 44, 46 Clypeaster cyclopilus 43, 44, 47 Clypeaster durandi 43, 44, 47 Clypeaster elongatus 43, 44 Clypeaster euclastus 43, 44, 47 Clypeaster europacificus 43, 45, 47, 49, 63 Clypeaster eurychorius 35, 43, 47, 49 Clypeaster eurypetalus 43, 49, 51 Clypeaster fervens 43, 49, 51, 54 Clypeaster humilis 43, 44, 49, 51 Clypeaster isolatus 43, 49 Clypeaster japonicus 43, 49, 51, 56 Clypeaster kieri 43, 49, 52 Clypeaster lamprus 43, 51, 52 Clypeaster latissimus 43, 51, 52 Clypeaster leptostracon 43, 54 Clypeaster luetkeni 43, 52, 54 Clypeaster lytopetalus 43, 54 Clypeaster miniaceus 43, 54, 55 Clypeaster nummus 43, 54 Clypeaster ochrus 43, 54, 55 Clypeaster oshimensis 43, 54, 56 Clypeaster pallidus 43, 56 Clypeaster pateriformis 43, 56 Clypeaster prostratus 34, 43, 57 Clypeaster rangianus  43, 57, 58 Clypeaster rarispinus 43, 58, 60 Clypeaster ravenelii 43, 58, 61 Clypeaster reticulatus 43, 49, 58, 61 Clypeaster rosaceus 35, 43, 58, 60, 62, 63 Clypeaster rotundus 43, 61, 62, 63 Clypeaster speciosus 43, 47, 62, 63, 64 Clypeaster subdepressus 35, 43, 54, 57, 60, 63, 64 Clypeaster telurus 43, 63, 65 Clypeaster tumidus 43, 63, 65 Clypeaster virescens 43, 63, 66 Clypeasteridae 3, 34, 40 Clypeasterids 81 Clypeasterina 3, 34, 37 Clypeasteroid 34, 36, 37, 145 Clypeasteroida 2, 3, 7, 9, 34 clypeatus, Carnarechinus 122 clypeatus, Cystechinus 122 cocosi, Encope micropora 107 coeleus, Idiobryssus 170, 172 columbaris, Brissopsis 157, 159, 161 compactus, Schizaster 283, 314, 316, 317 Conolampas 3, 18, 19 Conolampas diomedeae 18, 19 Conolampas malayana 18, 20

Conolampas murrayana 18, 19 Conolampas sigsbei 18, 19, 21 convergens, Echinocyamus 68, 69 cordatum, Echinocardium 219, 222, 223, 224, 226, 228 cordatus, Abatus 283, 285, 287, 322 cordatus, Parapneustes 338 cordatus, Pericosmus 273, 275 cordiformis, Lovenia 228, 230, 231 cordiformis, Maretia 242, 244, 245 Corystus 3, 120, 122 Corystus relictus 118, 120, 121, 123 Corystusidae 3, 119, 120, 122 costae (Rhabdobrissus), Plagiobrissus 184, 185, 186, 187 crassa, Echinolampas 20, 23 cribellum, Fibularia 73, 75 crispus, Echinocyamus 68, 69 cristatus; Paleopneustes 266, 268 cuneata, Echinocrepis 128, 129, 130 cuneus, Rhynobrissus 187, 189 curvidens, Abatus 284, 285 Cyclaster 4, 263 Cyclaster recens 263, 265 Cyclaster regalis 263, 265 cyclius, Ammotrophus 37, 39 cyclopilus, Clypeaster 43, 44, 47 cyclostomus, Echinoneus 7, 8 Cystechinus 4, 139, 141, 143 Cystechinus clypeatus 122 Cystechinus giganteus 141, 143 Cystechinus loveni 141, 143 Cystechinus wyvilli 141, 143 Cystocrepis setigera 128, 129 Cystocrepis 3, 128 damesi, Anabrissus 152, 153 davidi, Amphipneustes 322, 324, 325 debilis, Pourtalesia 131, 135 decagonale, Laganum 79, 80 deciesdigitata, Rotula 67, 91, 93 delectus, Cassidulus 12 Delopatagus 332 Delopatagus brucei 333 Dendraster 3, 36, 96, 98 Dendraster excentricus 98, 99 Dendraster laevis 98, 99 Dendraster terminalis 98, 99 Dendraster vizcainoensis 98 Dendrasteridae 3, 67, 93, 96 denudatus, Elipneustes 202, 204 depressa, Echinolampas 20, 24 depressum, Jacksonaster 79, 80 depressum, Laganum 79, 80 depressus, Nacospatangus 245, 246 desorii, Breynia 218, 219, 220 Diadematecea 1 dicrana, Metalia 174, 176, 180, 181 diomedeae, Conolampas 18, 19 diomedeae, Plesiozonus 269, 270, 271 diomedeae, Spatangus 253, 255 Diploporaster 4, 282, 297

 351

352 

 Index

Diploporaster barbatus 297, 298 Diploporaster savigny 297, 298 djakonovi, Marginoproctus 117 doederleini, Lovenia 228, 229 doederleini, Schizaster 314, 316, 318 dolosus, Hypselaster 298, 300, 301, 302 drygalskii, Antrechinus 139 dubium, Echinocardium 226 durandi, Clypeaster 43, 44, 47 dyscritus, Granobrissoides 236 Echinarachniidae 3, 67, 93, 98 Echinarachnius 3, 98 Echinarachnius parma 37, 99, 100 echinobrissoides, Aphanopora 30, 31 Echinocardium 4, 127, 147, 157, 212, 219, 222 Echinocardium capense 222, 223 Echinocardium cordatum australe 222 Echinocardium cordatum 219, 222, 223, 224, 226, 228 Echinocardium dubium 226 Echinocardium fenauxi 222, 223, 224 Echinocardium flavescens 219, 222, 223, 224, 226, 227 Echinocardium keiense 224 Echinocardium laevigaster 222, 224 Echinocardium lymani 222, 224, 226 Echinocardium mediterraneum 222, 223, 226, 227, 229 Echinocardium meteorense 222, 226, 229 Echinocardium mortenseni 222, 227, 229 Echinocardium pennatifidum 222, 223, 228, 229 Echinocrepis 3, 127, 128 Echinocrepis cuneata 128, 129, 130 Echinocrepis rostrata 129, 130 Echinocyamidae 3, 67, 68 Echinocyamus 3, 68, 72, 73 Echinocyamus apicatus 68, 69 Echinocyamus convergens 68, 69 Echinocyamus crispus 68, 69 Echinocyamus elegans 69 Echinocyamus grandiporus 69 Echinocyamus grandis 71 Echinocyamus incertus 71 Echinocyamus insularis 71 Echinocyamus macrostomus 71 Echinocyamus megapetalus 67, 68, 71 Echinocyamus planissimus 71, 72 Echinocyamus platytatus 67, 71, 72 Echinocyamus provectus 71, 72 Echinocyamus pusillus 35, 68, 71, 72, 73, 116 Echinocyamus scaber 71 Echinocyamus sollers 71, 72 Echinodiscus 3, 95, 96 Echinodiscus auritus 35, 96, 97 Echinodiscus bisperforatus 96, 97 Echinodiscus tenuissimus 96, 97 Echinodiscus truncatus 95, 96, 97 Echinogutta 127, 130 Echinolampadidae 3, 11, 18 Echinolampas 3, 10, 18, 19, 23 Echinolampas alexandri 20, 21, 23 Echinolampas chuni 20, 21, 23, 29

Echinolampas crassa 20, 23 Echinolampas depressa 20, 24 Echinolampas keiensis 20, 24, 25 Echinolampas koreana 20 Echinolampas ovata 12, 20, 21, 23, 25, 27 Echinolampas rangii 20, 25, 27 Echinolampas sternopetala 20, 24, 25, 27, 29 Echinolampas sumatrana 20, 21, 23, 29 Echinoneid 8 Echinoneidae 3, 7 Echinoneoida 3, 7 Echinoneus 3, 7, 8, 9 Echinoneus cyclostomus 7, 8 Echinosigra 3, 128, 130, 133, 135 Echinosigra amphora fabrefacta 133 Echinosigra amphora 133 Echinosigra (Echinogutta) amphora 130 Echinosigra (Echinogutta) antarctica 130 Echinosigra (Echinogutta) fabrefacta 130, 133 Echinosigra (Echinogutta) valvaedentata 130 Echinosigra (Echinogutta) 130 Echinosigra (Echinosigra) phiale 130, 133 Echinosigra (Echinosigra) porrecta 130 Echinosigra (Echinosigra) vityazi 130 Echinosigra (Echinosigra) 130 Echinosigra paradoxa 130 Echinosigra phiale 130 ecuadorensis, Encope micropora 107 edwardsi, Ova 308, 312 elegans, Breynia 218, 219, 221 elegans, Echinocyamus 69 elevata persica, Metalia 179 elevata, Hemimaretia 239 Elipneustes 4, 202, 205 Elipneustes denudatus 202, 204 Elipneustes rubens 202 elongata, Brissopsis 157, 159, 161 elongata, Lovenia 223, 228, 229, 230, 232 elongatus, Abatus 287, 288 elongatus, Clypeaster 43, 44 emarginata, Encope 102, 103 Encope 3, 100, 101, 102, 105, 106 Encope aberrans 102, 103, 105 Encope emarginata 102, 103 Encope grandis 67, 102, 103, 104 Encope laevis 102, 103, 104 Encope michelini 35, 102, 103, 104, 105 Encope micropora borealis 105, 107 Encope micropora californica 105, 107 Encope micropora cocosi 107 Encope micropora ecuadorensis 107 Encope micropora fragilis 107 Encope micropora insularis 107 Encope micropora irregularis 107 Encope micropora micropora 107 Encope micropora tetrapora 107 Encope micropora 105, 106, 107 Encope perspectiva 107 Encope wetmorei 107 enodatus, Prenaster 279, 281

Index 

epigonus, Oligopodia 15, 16 erinaceus, Heterobrissus 334, 335 euclastus, Clypeaster 43, 44, 47 Euechinoidea 3 Eupatagidae 4, 150, 198 Eupatagus 4, 198, 201, 202, 242 Eupatagus flindersi 198, 200 Eupatagus lymani 198, 200 Eupatagus micropetalus 198, 199, 201 Eupatagus obscurus 198, 201 Eupatagus rubellus 198, 201 Eupatagus valenciennesi 198, 202, 203 Eurhodia 3, 12, 15 Eurhodia relicta 15 europacificus, Clypeaster 43, 45, 47, 49, 63 eurychorius, Clypeaster 35, 43, 47, 49 Eurypatagidae 4, 150, 202 Eurypatagus 4, 202, 204 Eurypatagus grandiporus 204, 205 Eurypatagus ovalis 204, 205, 206 Eurypatagus parvituberculatus 204, 205, 207 eurypetalus, Clypeaster 43, 49, 51 evanescens, Brissopsis 157, 159, 161 excavatus, Tripylus 280, 281 excentrica, Agassizia 279, 280 excentricus, Dendraster 98, 99 excentricus, Linopneustes  205, 208 expergitus, Holanthus 148, 149 fabrefacta, Echinosigra (Echinogutta) 130, 133 Faorina 4, 269, 272 Faorina chinensis 269, 272 Fellaster 3, 34, 39 Fellaster zelandiae 37, 40, 42 fenauxi, Echinocardium 222, 223, 224 fervens, Clypeaster 43, 49 Fibularia 3, 67, 68, 73 Fibularia japonica 73, 76 Fibularia nutriens 73, 76 Fibularia ovulum 75, 77 Fibularia plateia 76, 77 Fibularia volva 76 Fibularia, cribellum 73, 75 Fibulariella acuta 76, 77, 78 Fibulariella angulipora 77, 78 Fibulariella  3, 67, 68, 73, 76 Fibulariella oblonga 78 Fibulariidae 3, 67, 73 fisheri, Scrippsechinus 198, 199 flavescens, Echinocardium 219, 222, 223, 224, 226, 227 flindersi, Eupatagus 198, 200 floridiensis, Schizaster 314, 318 fragilis, Brisaster 291, 294 fragilis, Encope micropora 107 fragilis, Homolampas 242, 243 fragilis, Hypselaster 298, 300, 301 fragilis, Linopneustes 205, 208, 210 frangibilis, Meoma 171, 172, 173 fudsiyama, Laganum 79, 81 fulva, Araeolampas 216, 217, 262, 265

Genicopatagus 5, 282, 322, 332 Genicopatagus affinis 322, 330, 332, 333 gibberulus, Schizaster 314, 316, 318 gibbosus, Holanthus 148, 149 giganteus, Cystechinus 141, 143 gigas, Brissus 167 gigas, Heterobrissus 334 glauca, Araeolampas 216, 217 Gonimaretia 245 Gonimaretia laevis 245 gracilis, Nacospatagus 245, 246 grandiporus, Echinocyamus 69 grandiporus, Eurypatagus 204, 205 grandis (Plagiobrissus), Plagiobrissus 184, 185 grandis (ventricosa) Meoma 171, 174, 175 grandis, Anametalia 152, 153 grandis, Echinocyamus 71 grandis, Encope 67, 102, 103, 104 Granobrissoides 4, 236 Granobrissoides dyscritus 236 Granobrissoides hirsutus 236, 238 Granopatagus 255 grantii, Melitta 103, 108, 109 gregalis, Lovenia 228, 231, 235 griegii, Sarsiaster 149 grisea, Lovenia 228 griseus, Scaphechinus 115 Gymnopatagus 4, 236, 238 Gymnopatagus magnus 239, 241 Gymnopatagus valdiviae 239, 241 Hagenowia rostrata 127 hastata, Araeolampas 216, 218 hawaiiensis, Lovenia 228, 230, 232, 235 Helgocystis 4, 127, 128, 131 Helgocystis carinata 131, 135 Heliophora 3, 90 Heliophora orbiculus 90, 93 Hemiaster 147 Hemiasterid 147 Hemiasteridae 4, 147 hemiasteroides, Rhynobrissus 187, 189 Hemimaretia 4, 236, 239 Hemimaretia elevata 239 hemingi, Heterobrissus 334, 335 heptneri, Pourtalesia 131, 135 Heterobrissus 5, 322, 334 Heterobrissus erinaceus 334, 335 Heterobrissus gigas 334 Heterobrissus hemingi 334, 335 Heterobrissus hystrix 334, 336 Heterobrissus niasicus 334, 335, 337 hickmani, Holanthus 148, 150 hilgardi, Palaeobrissus 194, 195 hinemoae, Peronella 83, 84 hirsuta, Pseudolovenia 236 hirsutus, Granobrissoides 236, 238 hirsutus, Lissospatangus 238 hirsutus, Plesiozonus 269, 271 hirsutus, Plexechinus 122, 124, 125, 126

 353

354 

 Index

hispida, Pourtalesia 131, 136 Holanthus 4, 147 Holanthus expergitus 148, 149 Holanthus gibbosus 148, 149 Holanthus hickmani 148, 150 Holasteroid 118 Holasteroida 3, 7, 18 Homolampas 4, 215, 236, 239 Homolampas fragilis 242, 243 Homolampas lovenioides 242, 243 humilis, Clypeaster 43, 44, 49, 51 humilis, Phryssocystis 260, 262 Hypselaster affinis 298, 300 Hypselaster brachypetalus 298, 300 Hypselaster dolosus 298, 300, 301, 302 Hypselaster fragilis 298, 300, 301 Hypselaster jukesii 300, 301, 302 Hypselaster kempi 300, 302, 303 Hypselaster 4, 282, 298 Hypselaster limicolus 300, 302, 303 Hypselaster maximus 300, 302 Hypselaster rotundus 300, 302 hystrix, Heterobrissus 334, 336 Idiobryssus 4, 152, 170 Idiobryssus coelus 170, 172 imitans, Solenocystis 137, 141 incerta sedis A, Paleopneustina 266, 321 incerta sedis B, Paleopneustina 266, 322 incertus, Echinocyamus 71 incus, Spatagobrissus 250, 252 inermis, Spatangus 253, 256 infidus, Cassidulus 12, 14 ingens, Abatus 284, 285, 288 insularis, Echinocyamus 71 insularis, Encope micropora 107 interruptus, Lonchophorus 248 interruptus, Nacospatangus 245, 246, 248 Irregularia 1, 7 irregularis, Encope micropora 107 isolatus, Clypeaster 43, 49 isometra, Mellita 95, 108, 109, 110 Isopatagus 4, 263 Isopatagus obovatus 263, 267 Jacksonaster depressum 79, 80 japonica, Fibularia 73, 76 japonica, Peronella 1, 79, 83, 85 japonicus, Clypeaster 43, 49, 51, 56 jarlii, Brissopsis 157, 159 jeffreysi, Pourtalesia 127, 131, 133, 137 jensenae, Rictocystis 133 josephinae, Palaeotropus 194, 195, 196 joubini, Laganum 81 jukesii, Hypselaster 300, 301, 302 jullieni (Rhabdobrissus), Plagiobrissus 184, 185, 186, 187 kanakoffi, Mellita 108, 109, 110, 111 keiense, Echinocardium 224 keiensis, Echinolampas 20, 24, 25

keiensis, Pericosmus 273, 275, 276 keiensis, Peronella 83, 85, 86 kempi, Hypselaster 300, 302, 303 kerguelensis, Brisaster 291, 293, 294 Kermabrissoides 4, 192, 194 Kermabrissoides siculum 194 kermadecensis, Metalia 174, 178 kieri, Clypeaster 43, 49, 52 kieri, Porterpygus 11 Koehleraster 3, 7, 9 Koehleraster abnormalis 7, 8 koehleri, Abatus 284, 287, 288 koehleri, Amphipneustes 322, 324, 325 koreana, Echinolampas 20 Kupeia 4, 148 Kupeia toi 147, 148, 150 lachesinella, Moira 303, 307 lacunosa, Ova 309, 312, 313 laevigaster, Echinocardium 222, 224 laevis, Dendraster 98, 99 laevis, Encope 102, 103, 104 laevis, Gonimaretia 245 laevis, Nacospatangus 246, 247, 248 Laganidae 3, 67, 78 Laganids 78, 79 Laganiformes 3, 37, 67 Laganum 3, 78 Laganum boninensis 79 Laganum centrale 79 Laganum decagonale 79, 80 Laganum depressum 79, 80 Laganum fudsiyama 79, 81, 82 Laganum joubini 81 Laganum laganum 35, 67, 81, 82 Laganum putnami 82 Laganum retinens 82 Laganum versatile 82 laganum, Laganum 35, 67, 81, 82 laguncula, Pourtalesia 131, 133, 137 lamprus, Clypeaster 43, 51, 52 Lanthonia 108 lata, Lovenia 228, 232, 235 latecarinatus, Brissus 167, 169, 170 latifrons, Brisaster 291, 293, 294 latissima, Metalia 174, 176, 178 latissimus, Clypeaster 43, 51, 52 Leodia  3,100, 101, 107 Leodia sexiesperforata 101, 102, 107, 108 leptostracon, Clypeaster 43, 54 lesueuri, Peronella 83, 84, 86 lethe, Moira 303, 307, 308, 309 limicolus, Hypselaster 300, 302, 303 Linopneustes 4, 202, 205, 250 Linopneustes brachypetalus 205, 208 Linopneustes excentricus 205, 208 Linopneustes fragilis 205, 208, 210 Linopneustes longispinus 205, 208, 210 Linopneustes murrayi 205, 208, 210, 211 Linopneustes spectabilis 205, 210, 213

Index 

Lissospatangus hirsutus 238 Lonchophorus interruptus 248 longifissa, Mellita 108, 109, 110, 111, 112 longispinus, Linopneustes 205, 208, 210 lorioli, Amphipneustes 322, 324, 326 loveni, Cystechinus 141, 143 loveni, Paleotrema 195, 196 loveni, Tropholampas 30, 33 Lovenia 4, 212, 219, 228, 242 Lovenia camarota 228, 231 Lovenia cordiformis 228, 230, 231 Lovenia doederleini 228, 229 Lovenia elongata 223, 228, 229, 230, 232 Lovenia gregalis 228, 231, 235 Lovenia grisea 228 Lovenia hawaiiensis 228, 230, 232, 235 Lovenia lata 228, 232, 235 Lovenia subcarinata 228, 230, 235 Lovenia triforis 218, 228, 236 Loveniidae 4, 150, 212 lovenioides, Homolampas 242, 243 luetkeni, Clypeaster 43, 52, 54 luetkeni, Spatangus 253, 255, 257, 258 luzonica, Brissopsis 157, 159, 163, 167 lymani, Echinocardium 222, 224, 226 lymani, Eupatagus 198, 200 lyrifera, Brissopsis 157, 161, 163 lytopetalus, Clypeaster 43, 54 macronesius, Pericosmus 273, 276, 278 Macropneustidae 4, 150, 260 macroproctes, Peronella 84, 86, 87 macrostomus, Echinocyamus 71 magnus, Gymnopatagus 239, 241 mai, Sinaechinocyamus 116, 117 malayana, Conolampas 18, 20 malayanus, Cassidulus 12, 13 manni, Astriclypeus 95 Maretia 4, 202, 212, 236, 242, 244, 245 Maretia carinata 242, 244 Maretia cordiformis 242, 244, 245 Maretia planulata 242, 244, 246 Maretia tuberculata 242, 245 Maretiidae 4, 150, 151, 236 Marginoproctus 3, 94, 117 Marginoproctus djakonovi 117 marsupialis, Amphipneustes 322, 324, 327 mathesoni, Spatangus 253, 257, 258 mauretianus, Pericosmus 273, 276 maximus, Hypselaster 300, 302 mediterranea, Brissopsis 157, 159, 162, 164 mediterraneum, Echinocardium 222, 223, 226, 227, 229 megapetalus, Echinocyamus 67, 68, 71 melanostomus, Pericosmus 273, 276 Mellita 3, 101, 108, 109 Melitta grantii 103, 108, 109 Mellita isometra 95, 108, 109, 110 Mellita kanakoffi 108, 109, 110, 111 Mellita longifissa 108, 109, 110, 111, 112 Mellita notabilis 108, 110, 111, 113

Mellita quinquiesperforata 108, 111, 113 Mellita tenuis 93, 94, 101, 108, 112, 114 Mellitella 3, 101, 113 Mellitella stokesii 101, 113, 114 Mellitidae 3, 67, 94, 95, 100 Meoma 4, 152, 171 Meoma (ventricosa) grandis 171, 174, 175 Meoma (ventricosa) ventricosa 171, 172, 174, 175 Meoma cadenati 171, 172, 173 Meoma frangibilis 171, 172, 173 Meoma ventricosa  173 mergulensis, Peronella 86 Metalia 4, 152, 155, 174, 176, 179, 180 Metalia angustus 174, 176 Metalia dicrana 174, 176, 180, 181 Metalia kermadecensis 174, 178 Metalia latissima 174, 176, 178 Metalia nobilis 174, 178 Metalia persica 174, 179 Metalia robillardi 174, 176, 180, 181 Metalia spatagus 151, 174, 176, 179, 180, 181 Metalia sternalis 174, 176, 180, 182 Metalia townsendi 174, 180, 183 meteorense, Echinocardium 222, 226, 229 michelini, Encope 35, 102, 103, 104, 105 Micrasteridae 4, 261, 263 Micrasterina 4, 261 micrasteroides, Neopneustes 180, 183 micropetala, Brissopsis 157, 162, 165 Micropetalon 3, 8, 9 Micropetalon purpureum 8, 9 micropetalus, Eupatagus 198, 199, 201 micropora Encope 10, 106, 107 micropora, Encope micropora 107 miniaceus, Clypeaster 43, 54, 55 minuta, Peronella 86 mirabilis , Palaeostoma  148, 150, 151 mirabilis, Scaphechinus 115, 116 mirabilis, Spatagobrissus 250 miranda, Pourtalesia 131, 133, 138 mironovi, Amphipneustes 322, 324 mitis, Cassidulus 12, 14, 15 Moira 4, 127, 157, 282, 302, 304, 308 Moira atropos 222, 302, 303, 304, 305, 307 Moira clotho 303, 306, 307 Moira lachesinella 303, 307 Moira lethe 303, 307, 308, 309 Moira stygia 303, 308, 310 Moiropsis 4, 28, 308 Moiropsis claudicans 308, 310 mortenseni, Antrechinus 139, 142 mortenseni, Echinocardium 222, 227, 229 mortenseni, Taimanawa 191, 192, 193 Mortonia 3, 68, 72, 73 Mortonia australis 67, 73, 74 Mortonia polyporus 73, 74 moseleyi, Brisaster 291, 293, 296, 319 multispina, Phrissocystis 260 multispinus, Argopatagus 260, 261 multispinus, Spatangus 254, 257, 258

 355

356 

 Index

multituberculata, Paramaretia 212 murrayana, Conolampas 18, 19 murrayi, Linopneustes 205, 208, 210, 211 myorensis, Schizaster 314 Nacospatangus 4, 236, 242, 245 Nacosatangus gracilis 245, 246 Nacospatangus altus 245, 246, 247 Nacospatangus depressus 245, 246 Nacospatangus interruptus 245, 246, 248 Nacospatangus laevis 246, 247, 248 Nacospatangus oblongus 246, 248, 249 Nacospatangus tylotus 246, 248, 249 Nannolampas 3, 30 Nannolampas tenera 30, 31 naresianus, Urechinus 14, 147 neanika, Breynia 218, 219, 221 Neognathostomata 3, 7, 9, 10 Neolampadidae 3, 11, 29 Neolampas 3, 10, 30, 31 Neolampas rostellata 30, 32 Neopneustes 4, 152, 180 Neopneustes micrasteroides 180, 183 niasicus, Heterobrissus 334, 335, 337 nimrodi (Pseudabatus), Abatus 284, 289, 290 nobilis, Metalia 174, 178 nordenskjoldi, Antrechinus 139, 142 notabilis, Mellita 108, 110, 111, 113 nummus, Clypeaster 43, 54 nutriens, Fibularia 73, 76 obesus, Brissus 167, 170, 171 obliqua, Brissopsis 157, 162, 165 oblonga, Fibulariella 78 oblonga, Peronella 86 oblongus, Nacospatangus 246, 248, 249 oblongus, Pericosmus 273, 276 obovatus, Isopatagus 263, 267 obovatus, Schizaster 314, 319 obscurus, Eupatagus 198, 201 occidentalis, Apatopygus 10, 11 ochrus, Clypeaster 43, 54, 55 oldhami, Brissopsis 157, 162, 165 Oligopodia 3, 12, 15, 34 Oligopodia epigonus 15, 16 Oligopygoida 3 orbicularis, Peronella 87, 88 orbiculus, Heliophora 90, 93 orbignyana, Ova 309, 313, 314, 316 oshimensis, Clypeaster  43, 54, 56 Ova 4, 282, 291, 308, 314 Ova canalifera 308, 309, 311, 312 Ova edwardsi 308, 312 Ova lacunosa 309, 312, 313 Ova orbignyana 309, 313, 314, 316 Ova portjacksonensis 309, 313, 314 ovalis, Eurypatagus 204, 205, 206 ovata, Aceste 289, 291, 293 ovata, Echinolampas 12, 20, 21, 23, 25, 27 ovatum, Paleotrema 195, 196

oviformis, Pycnolampas 250, 251 ovulum, Fibularia 75, 77 owstoni, Brisaster 291, 293, 296 pacifica, Brissopsis 157, 162, 166 pacificus, Plagiobrissus 184, 185, 186, 187 pacificus, Rhyncholampas 12, 17, 18 Palaeobrissus 4, 193, 194 Palaeobrissus hilgardi 194, 195 Palaeostoma 4, 148 Palaeostoma mirabilis 148, 150, 151 Palaeostomatid 147 Palaeostomidae 4, 148 Palaeotrema 4 Palaeotropidae 4, 150, 192 Palaeotropus josephinae 194, 195, 196 Palaeotropus 4, 193, 194 Palaeotropus uniporum 195 Paleopneustes 4, 266 Paleopneustes cristatus 266, 268 Paleopneustid 269, 283, 321 Paleopneustidae 4, 266, 269 Paleopneustina 4, 5, 266, 267, 322 Paleopneustina incertae sedis A 266, 321 Paleopneustina incertae sedis B 266, 322 Paleotrema 4, 193, 194, 195 Paleotrema loveni 195, 196 Paleotrema ovatum 195, 196 pallidus, Clypeaster 43, 56 pallidus, Spatangus 254, 257, 258 Paraamphiope raimondii 96 paradoxa, Echinosigra 130 parallela, Brissopsis 157, 165, 168 Paramaretia 4, 202, 212 Paramaretia multituberculata 212 Paramaretia peloria 212, 214 Parapneustes 5, 321, 322, 335 Parapneustes cordatus 338 Parapneustes reductus 338 Paraster 314 parma, Echinarachnius 37, 99, 100 parvituberculatus, Eurypatagus 204, 205, 207 parvus, Plexechinus 122, 123, 124 pateriformis, Clypeaster 43, 56 paucituberculatus, Spatangus 254, 255, 257, 258 pellucida, Peronella 88, 89 peloria, Paramaretia 212, 214 pennatifidum, Echinocardium 222, 223, 228, 229 Pericosmid 269 Pericosmidae 4, 266, 269 Pericosmus 4, 269, 272, 273, 321 Pericosmus akabanus 273, 274 Pericosmus bidens 273, 274 Pericosmus cordatus 273, 275 Pericosmus keiensis 273, 275, 276 Pericosmus macronesius 273, 276, 278 Pericosmus mauretianus 273, 276 Pericosmus melanostomus 273, 276 Pericosmus oblongus 273, 276 Pericosmus porphyrocardius 273, 276, 278

Index 

Pericosmus tenuis 273, 276, 278 Peripatagus 4, 266 Peripatagus cinctus 266, 268 Peronella 1, 3, 78, 83, 86 Peronella affinis 83 Peronella analis 83 Peronella hinemoae 83, 84 Peronella japonica 1, 79, 83, 85 Peronella keiensis 83, 85, 86 Peronella lesueuri 83, 84, 86 Peronella macroproctes 84, 86, 87 Peronella mergulensis 86 Peronella minuta 86 Peronella oblonga 86 Peronella orbicularis 87, 88 Peronella pellucida 88, 89 Peronella peronii 88, 89 Peronella rubra 86, 88, 90 Peronella rullandi 88 Peronella strigata 88 Peronella tuberculata 88, 90 peronii, Peronella 88, 89 persica, Brissopsis 179 persica, Metalia 174, 179 perspectiva, Encope 107 phiale, Echinosigra (Echinosigra) 130, 133 phiale, Echinosigra 130 philippii, Abatus 288, 289 philippii, Tripylaster 319, 320 Phrissocystis 260 Phrissocystis aculeata 260 Phrissocystis humilis 260, 262 Phrissocystis multispina 260 Pilematechinus 4, 139, 141, 144 Pilematechinus belyaevi 141, 145 Pilematechinus rathbuni 141, 144, 145 Pilematechinus vesica 141, 144, 145 placenta, Arachnoides 37, 39, 40 placopetalus, Rhynobrissus 189 Plagiobrissus 4, 152, 174, 183 Plagiobrissus (Plagiobrissus) africanus 183, 185 Plagiobrissus (Plagiobrissus) grandis 184, 185 Plagiobrissus (Rhabdobrissus) costae 184, 185, 186, 187 Plagiobrissus (Rhabdobrissus) jullieni 184, 185, 186, 187 Plagiobrissus (Rhabdobrissus) 184, 185 Plagiobrissus pacificus 184, 185, 186, 187 planissimus, Echinocyamus 71, 72 planulata, Maretia 242, 244, 246 planus, Argopatagus 260, 261 planus, Plexechinus 122, 123, 124 planus, Sinaechinocyamus 117 plateia, Fibularia 76, 77 Platybrissus roemeri 212, 215 Platybrissus 4, 202, 212 platytatus, Echinocyamus 67 platyterus, Ammotrophus 39, 40 Plesiozonus 4, 266, 269, 271 Plesiozonus diomedeae 269, 270, 271 Plesiozonus hirsutus 269, 271 Plesiozonus tenuis 269, 271

Plethotaenia 4, 252 Plethotaenia angularis 253, 254 Plethotaenia spatangoides 253, 254 Plexechinidae 3, 119, 122 Plexechinus 3, 122 Plexechinus aoteanus 122, 123, 124 Plexechinus cinctus 122, 124, 125, 126 Plexechinus hirsutus 122, 124, 125, 126 Plexechinus parvus 122, 123, 124 Plexechinus planus 122, 123, 124 Plexechinus spectabilis 122, 124, 125 Plexechinus sulcatus 122, 124, 126 polyporus, Mortonia 73, 74 porphyrocardius, Pericosmus 273, 276, 278 porrecta, Echinosigra (Echinosigra) 130 Porterpygus 3, 11 Porterpygus kieri 11 portjacksonensis, Ova 309, 313, 314 Pourtalesia 4, 125, 127, 128, 131, 133, 135 Pourtalesia alcocki 131, 135 Pourtalesia aurorae 131, 135 Pourtalesia debilis 131, 135 Pourtalesia heptneri 131, 135 Pourtalesia hispida 131, 136 Pourtalesia jeffreysi 127, 131, 133, 137 Pourtalesia laguncula 131, 133, 137 Pourtalesia miranda 131, 133, 138 Pourtalesia tanneri 131, 133, 138 Pourtalesia wandeli 138 Pourtalesiid 119 Pourtalesiidae 3, 125, 128 Prenaster 4, 278, 279 Prenaster enodatus 279, 281 Prenasterid 281 Prenasteridae 4, 266, 278 prostatus, Clypeaster 34, 43, 57 Protenaster 5, 282, 314 Protenaster australis 314 provectus, Echinocyamus 71, 72 Pseudabatus 284, 290 Pseudolovenia  4, 212, 236 Pseudolovenia hirsuta 236 Pseudomaretia 245 platytatus, Echinocyamus 71, 72 purpureum, Micropetalon 8, 9 purpureus, Spatangus 68, 254, 255, 257, 259 pusillus, Echinocyamus 35, 68, 71, 72, 73, 116 putnami, Laganum 82 Pycnolampas 4, 236, 250 Pycnolampas oviformis 250, 251 pyramidalis, Rhynobrissus 187, 189, 190 quinquiesperforata, Mellita 108, 111, 113 raimondii, Paraamphiope 96 rangianus, Clypeaster 43, 57, 58 rangii, Echinolampas 20, 25, 27 rarispinus, Clypeaster 43, 58, 60 raschi, Spatangus 254, 255, 259, 260 rathbuni, Pilematechinus 141, 144, 145

 357

358 

 Index

ravenelii, Clypeaster 43, 58, 61 recens, Apatopygus 10, 11, 30 recens, Cyclaster 263, 265 recens, Studeria 32 reductus, Parapneustes 338 reductus, Tripylus 338 regalis, Cyclaster 263, 265 regularis, Anametalia 152, 154, 155 relicta, Calymne 119, 121 relicta, Eurhodia 15 relictus, Corystus 118, 120, 121, 123 relictus, Stereopneustes 121 relictus, Taimanawa 192, 193 reticulatus, Clypeaster 43, 49, 53, 54, 58, 61 reticulatus, Urechinus 145 retinens, Laganum 82 revinctus, Cionobrissus 170, 172 Rhabdobrissus 184, 185 Rhyncholampas 3, 17 Rhyncholampas pacificus 12, 17, 18 Rhynobrissus 4, 152, 187, 191 Rhynobrissus cuneus 187, 189 Rhynobrissus hemiasteroides 187, 189 Rhynobrissus placopetalus 189 Rhynobrissus pyramidalis 187, 189, 190 Rhynobrissus tumulus 187, 189, 191 Rictocystis 4, 133 Rictocystis jensenae 133 robillardi, Metalia 174, 176, 180, 181 roemeri, Platybrissus 212, 215 rosaceus, Clypeaster 35, 43, 58, 60, 62, 63 rostellata, Neolampas 30, 32 rostrata, Aeropsis 262, 263, 265 rostrata, Araeolampas 216, 218 rostrata, Echinocrepis 129, 130 rostrata, Hagenowia 127 rostratus, Amphipneustes 322, 324, 325, 328, 329 Rotula 3, 88, 91 Rotula deciesdigitata 67, 91, 93 Rotulidae 3, 67, 88 rotundatus, Schizaster 314, 319 rotundus Hypselaster 300, 302 rotundus, Clypeaster 43, 61, 62, 63 rubellus, Eupatagus 198, 201 rubens, Elipneustes 202 rubra, Peronella 86, 88, 90 rullandi, Peronella 88 Sarsiaster 4, 148, 149, 151 Sarsiaster griegii 149, 151 savigny, Diploporaster 297, 298 scaber, Echinocyamus 71 Scaphechinus brykovi 115 Scaphechinus griseus 115 Scaphechinus mirabilis 115, 116 Scaphechinus 3, 113, 115, 116 Scaphechinus tenuis 115 Schizaster 5, 282, 291, 308, 314 Schizaster compactus 283, 314, 316, 317 Schizaster doederleini 314, 316, 318

Schizaster floridiensis 314, 318 Schizaster gibberulus 314, 316, 318 Schizaster myorensis 314 Schizaster obovatus 314, 319 Schizaster rotundatus 314, 319 Schizasterid 281, 282, 283, 302, 321 Schizasteridae 4, 266, 278, 281, 282 Schizocosmus 5, 321 Schizocosmus abatoides 321 Scrippsechinus 4, 193, 194, 198 Scrippsechinus fisheri 198, 199 scrobiculata, Agassizia 279, 280 Sculpsitechinus 96 Sculpsitechinus auritus 96 Sculpsitechinus tenuissimus 96 Scutellidae 3, 67, 94, 113 Scutelliform 91, 93, 95 Scutelliformes 3, 37, 67, 91 Scutellina 3, 67 setigera, Cystocrepis 128, 129 sexiesperforata, Leodia 101, 102, 108 shackletoni, Abatus 284, 289, 290 sibogae, Sternopatagus 120, 122 siculum, Kermabrissoides 194 sigsbei, Conolampas 18, 19, 21 similis, Amphipneustes 322, 329 similis, Brissopsis 157, 167, 168 Sinaechinocyamus 3, 94, 113, 115, 116, 117 Sinaechinocyamus mai 116, 117 Sinaechinocyamus planus 117 sinensis, Anochanus 30 Solenocystis 4, 128, 137 Solenocystis imitans 137, 141 sollers, Echinocyamus 71, 72 Spatagobrissus 4, 236, 250 Spatagobrissus incus 250, 252 Spatagobrissus mirabilis 250 Spatagocystis 4, 128, 139 Spatagocystis challengeri 139, 141 spatagus, Metalia 174, 176, 179, 180, 181 Spatangidae 4, 150, 252 Spatangoid 118, 302 Spatangoida 2, 4, 7, 118, 145 Spatangoidea 4, 150, 198 spatangoides, Plethotaenia 253, 254 Spatangus 4, 252, 253, 255 Spatangus altus 253, 255, 256 Spatangus beryl 253, 255, 260 Spatangus californicus 253, 255, 256 Spatangus capensis 253, 255, 256 Spatangus diomedeae 253, 255 Spatangus inermis 253, 256 Spatangus luetkeni 253, 255, 257, 258 Spatangus mathesoni 253, 257, 258 Spatangus multispinus 254, 257, 258 Spatangus pallidus 254, 257, 258 Spatangus paucituberculatus 254, 255, 257, 258 Spatangus purpureus 68, 254, 255, 257, 259 Spatangus raschi 254, 255, 259, 260 Spatangus thor 254

Index 

spatagus, Metalia 151 speciosus, Clypeaster 43, 47, 62, 63, 64 spectabilis, Linopneustes 205, 210, 213 spectabilis, Plexechinus 122, 124, 125 Stereopneustes 120 Stereopneustes relictus 121 sternalis, Metalia 174, 176, 180, 182 sternaloides, Anametalia 152, 154, 155 Sternopatagus 3, 119, 120 Sternopatagus sibogae 120, 122 sternopetala, Echinolampas 20, 24, 25, 27, 29 stokesii, Mellitella 101, 113, 114 strigata, Peronella 88 Studeria 3, 30, 32 Studeria recens 32 stygia, Moira 303, 308, 310 subcarinata, Lovenia 228, 230, 235 subdepressus, Clypeaster 35, 43, 54, 57, 60, 63, 64 sulcatus, Plexechinus 122, 124, 126 sumatrana, Echinolampas 20, 21, 23, 29 Taimanawa 4, 152, 189, 192, 193 Taimanawa mortenseni 191, 192, 193 Taimanawa relictus 192, 193 Taiwanaster 115 Taiwanasteridae 3, 67, 94, 113, 115 tanneri, Pourtalesia 131, 133, 138 tasmanicus, Brisaster 291, 293 telurus, Clypeaster 43, 63, 65 tenera, Nannolampas 30, 31 tenuis, Arachnoides 37, 39, 40 tenuis, Mellita 93, 94, 101, 108, 112, 114 tenuis, Pericosmus 273, 276, 278 tenuis, Plesiozonus 269, 271 tenuis, Scaphechinus 115 tenuissimus, Echinodiscus 96, 97 tenuissimus, Sculpsitechinus 96 terminalis, Dendraster 9, 99 tetrapora, Encope micropora 107 thor, Spatangus 254 toi, Kupeia 148, 150 townsendi, Brisaster 291, 293, 297 townsendi, Metalia 174, 180, 183 triforis, Lovenia 218, 228, 236 Tripylaster 5, 282, 291, 319

Tripylaster philippii 319, 320 Tripylus 4, 279, 282, 321 Tripylus excavatus 280, 281 Tripylus reductus 338 Tropholampas 3, 30, 33 Tropholampas loveni 30, 33 truncatus, Echinodiscus 95, 96, 97 tuberculata, Maretia 242, 245 tuberculata, Peronella 88, 90 tumidus, Clypeaster 43, 63, 65 tumulus, Rhynobrissoides 187, 189, 191 tylotus, Nacospatangus 246, 248, 249 unicolor, Brissus 1, 167, 170 uniporum, Palaeotropus 195 Urechinidae 4, 139 Urechinina 119 Urechinus 4, 139, 145 Urechinus antepodeanus 145, 147 Urechinus naresianus 145, 147 Urechinus reticulatus  145 valdiviae, Gymnopatagus 239, 241 valenciennesi, Eupatagus  198, 202, 203 valvaedentata, Echinosigra (Echinogutta) 130 vannoordenburgi, Brissalius 155, 157 ventricosa (ventricosa) Meoma 171, 172, 174, 175 ventricosa, Meoma 173 versatile, Laganum 82 vesica, Pilematechinus 141, 144, 145 virescens, Clypeaster 43, 63, 66 vitreus, Argopatagus 260, 261, 262 vityazi, Echinosigra (Echinosigra) 130 vizcainoensis, Dendraster 98 volva, Fibularia 76 vredenburgi, Breynia 218, 219 wandeli, Pourtalesia 138 weberi, Aceste 289, 291 wetmorei, Encope 107 wyvilli, Cystechinus 141, 143 zealandiae, Brissopsis 157, 162, 165 zelandiae, Fellaster 37, 40, 42

 359