West African Brachyuran Crabs - Smithsonian Institution Libraries
West African Brachyuran Crabs - Smithsonian Institution Libraries
West African Brachyuran Crabs - Smithsonian Institution Libraries
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<strong>West</strong> <strong>African</strong> <strong>Brachyuran</strong> <strong>Crabs</strong><br />
(Crustacea: Decapoda)<br />
RAYMOND B. MANNING<br />
and<br />
L. B. HOLTHUIS<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 306<br />
;
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S. Dillon Ripley<br />
Secretary<br />
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S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y • N U M B E R 3 0 6<br />
<strong>West</strong> <strong>African</strong> <strong>Brachyuran</strong> <strong>Crabs</strong><br />
(Crustacea: Decapoda)<br />
Raymond B. Manning<br />
and L. B. Holthuis<br />
SMITHSONIAN INSTITUTION PRESS<br />
City of Washington<br />
1981
ABSTRACT<br />
Manning, Raymond B., and L. B. Holthuis. <strong>West</strong> <strong>African</strong> <strong>Brachyuran</strong> <strong>Crabs</strong><br />
(Crustacea: Decapoda). <strong>Smithsonian</strong> Contributions to Zoology, number 306, 379<br />
pages, 88 figures, 1981.—The <strong>West</strong> <strong>African</strong> marine brachyuran crab fauna,<br />
comprising 218 named species in 120 genera and 26 familes, is surveyed.<br />
Sixteen new genera and 24 new species are recognized. Synonymies are<br />
updated for the tropical species, and all 300 + Eastern Atlantic species are<br />
listed. Original references and synonymies are provided for all 146 Eastern<br />
Atlantic genera. Synonymies have been compiled for all 36 currently recognized<br />
families of marine crabs. Twenty-nine families are represented in the<br />
Eastern Atlantic fauna. One family, Hexapodidae Miers, 1886, and one<br />
subfamily, Camptandriinae Stimpson, 1858 (Ocypodidae) are revised at the<br />
generic level. The genera Liocarcinus Stimpson, 1871 (Portunidae), Machaerus<br />
Leach, 1818 (Goneplacidae), and Lambdophallus Alcock, 1900, Paeduma Rathbun,<br />
1897, Parahexapus Balss, 1922, Pseudohexapus Monod, 1956, and Thaumastoplax<br />
Miers, 1881 (all Hexapodidae), are defined and recognized. It is<br />
suggested that the family Geryonidae Colosi, 1923, shows closest affinities<br />
with the family Portunidae Rafinesque, 1815.<br />
OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded<br />
in the <strong>Institution</strong>'s annual report, <strong>Smithsonian</strong> Year. SERIES COVER DESIGN: The coral Montastrea<br />
cavemosa (Linnaeus).<br />
Library of Congress Cataloging in Publication Data<br />
Manning, Raymond B. 1934-<br />
<strong>West</strong> Africa brachyuran crabs (Crustacea: Decapoda)<br />
(<strong>Smithsonian</strong> contributions to zoology ; no. 306)<br />
Bibliography: p.<br />
Supt. of Docs, no.: SI 1.27:306<br />
1. <strong>Crabs</strong>—Africa, <strong>West</strong>—Classification, 2. Crustacea—Classification. 3. Crustacea—Africa,<br />
<strong>West</strong>—Classification. I. Holthuis, L. B., joint author. II. Title. III. Series: <strong>Smithsonian</strong><br />
<strong>Institution</strong>. <strong>Smithsonian</strong> contributions to zoology ; no. 306.<br />
QL1.S54 no. 306 (QL444.M33] 591s [595.3'842] 80-607105
Contents<br />
Page<br />
Introduction 1<br />
Collections Studied 3<br />
Format Considerations 4<br />
Repositories and Abbreviations 5<br />
Acknowledgments 6<br />
Family RANINIDAE de Haan, 1839 6<br />
Subfamily NOTOPODINAE Serene and Umali, 1972 7<br />
Genus Ranilia H. Milne Edwards, 1837 7<br />
* Ranilia constricta (A. Milne Edwards, 1880) 7<br />
Subfamily RANININAE de Haan, 1839 9<br />
Genus Cyrtorhina Monod, 1956 9<br />
Cyrtorhina granulosa Monod, 1956 9<br />
Genus Raninoides H. Milne Edwards, 1837 10<br />
* Raninoides bouvieri Capart, 1951 10<br />
Family HOMOLODROMIIDAE Alcock, 1899 11<br />
Family DROMIIDAE de Haan, 1833 11<br />
Genus Dromia Weber, 1795 11<br />
Dromia bollorei Forest, 1974 11<br />
Dromia marmorea Forest, 1974 11<br />
* Dromia monodi Forest and Guinot, 1966 15<br />
Dromia nodosa A. Milne Edwards and Bouvier, 1898 19<br />
Genus Sternodromia Forest, 1974 19<br />
* Sternodromia spinirostris (Miers, 1881) 19<br />
Family DYNOMENIDAE Ortmann, 1892 23<br />
Genus Dynomene Desmarest, 1823 23<br />
* Dynomene filholi Bouvier, 1894 23<br />
Family LATREILLIIDAE Stimpson, 1858 24<br />
Genus Latreillia Roux, 1830 24<br />
Latreillia elegans Roux, 1830 25<br />
Family HOMOLIDAE de Haan, 1839 25<br />
Genus Homola Leach, 1815 25<br />
* Homola barbata (Fabricius, 1793) 25<br />
Genus Paromola Wood-Mason and Alcock, 1891 27<br />
Paromola cuvieri (Risso, 1816) 27<br />
Family CYCLODORIPPIDAE Ortmann, 1892 28<br />
Family CYMONOMIDAE Bouvier, 1898 28<br />
Family DORIPPIDAE MacLeay, 1838 28<br />
Key to Subfamilies and Genera of Dorippidae 29<br />
Subfamily DORIPPINAE MacLeay, 1838 30<br />
Genus Medorippe, new genus 31<br />
iii
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
* Medorippe lanata (Linnaeus, 1767), new combination 31<br />
Genus Phyllodorippe, new genus 35<br />
* Phyllodorippe armata (Miers, 1881), new combination 36<br />
Subfamily ETHUSINAE Guinot, 1977 38<br />
Genus Ethusa Roux, 1830 38<br />
* Ethusa rosacea A. Milne Edwards and Bouvier, 1897 38<br />
* Ethusa rugulosa A. Milne Edwards and Bouvier, 1897 39<br />
* Ethusa vossi, new species 39<br />
Genus Ethusina Smith, 1884 42<br />
Key to Eastern Atlantic Species of Ethusina 43<br />
Ethusina alba (Filhol, 1884) 43<br />
* Ethusina beninia, new species 46<br />
Family CALAPPIDAE de Haan, 1833 49<br />
Subfamily CALAPPINAE de Haan, 1833 49<br />
Genus Acanthocarpus Stimpson, 1871 49<br />
* Acanthocarpus brevispinis Monod, 1946 50<br />
Genus Calappa Weber, 1795 51<br />
Calappa gallus (Herbst, 1803) 51<br />
Calappa granulata (Linnaeus, 1758) 51<br />
* Calappa pelii Herklots, 1851 52<br />
* Calappa rubroguttata Herklots, 1851 54<br />
Genus Cycloes de Haan, 1837 56<br />
Cycloes cristata (Brulle, 1837) 56<br />
Subfamily MATUTINAE de Haan, 1835 56<br />
Genus Matuta Weber, 1795 56<br />
Matuta michaelseni Balss, 1921 56<br />
Family LEUCOSIIDAE Samouelle, 1819 57<br />
Subfamily EBALIINAE Stimpson, 1871 59<br />
Genus Atlantotlos Doflein, 1904 59<br />
* Atlantotlos rhombifer Doflein, 1904 59<br />
Genus Ebalia Leach, 1817 60<br />
* Ebalia qffinis Miers, 1881 60<br />
Ebalia cranchii Leach, 1817 61<br />
Ebalia nux A. Milne Edwards, 1883 61<br />
* Ebalia tuberculata Miers, 1881 61<br />
Ebalia tuberosa (Pennant, 1777) 63<br />
Genus Merocryptus A. Milne Edwards, 1873 64<br />
Merocryptus obsoletus A. Milne Edwards and Bouvier, 1898 ... 64<br />
Subfamily ILIINAE Stimpson, 1871 64<br />
Genus Ilia Leach, 1817 64<br />
Ilia nucleus (Linnaeus, 1758) 64<br />
* Ilia spinosa Miers, 1881 64<br />
Subfamily LEUCOSIINAE Samouelle, 1819 65<br />
Genus Philyra Leach, 1817 65<br />
Philyra cristata Miers, 1881 66<br />
* Philyra laevidorsalis Miers, 1881 66<br />
Page
NUMBER 306<br />
Genus Pseudomyra Capart, 1951 66<br />
* Pseudomyra mbizi Capart, 1951 66<br />
Family BELLIIDAE Dana, 1852 67<br />
Family ATELECYCLIDAE Ortmann, 1893 68<br />
Genus Atelecyclus Leach, 1814 68<br />
Atelecydus rotundatus (Olivi, 1792) 68<br />
Atelecyclus undecimdentatus (Herbst, 1783) 69<br />
Family THIIDAE Dana, 1852 69<br />
Genus Thia Leach, 1815 69<br />
Thia scutellata (Fabricius, 1793) 69<br />
Family CANCRIDAE Latreille, 1803 69<br />
Family PIRIMELIDAE Alcock, 1899 69<br />
Genus Pirimela Leach, 1816 70<br />
Pirimela denticulata (Montagu, 1808) 70<br />
Genus Sirpus Gordon, 1953 70<br />
Sirpus monodi Gordon, 1953 70<br />
* Sirpus gordonae, new species 70<br />
Family CORYSTIDAE Samouelle, 1819 72<br />
Genus Nautilocorystes H. Milne Edwards, 1837 72<br />
Nautilocorystes ocellatus (Gray, 1831) 72<br />
Family BYTHOGRAEIDAE Williams, 1980 72<br />
Family PORTUNIDAE Rafinesque, 1815 72<br />
Subfamily CARCININAE MacLeay, 1838 75<br />
Genus Carcinus Leach, 1814 75<br />
Carcinus maenas (Linnaeus, 1758) 75<br />
Genus Xaiva MacLeay, 1838 75<br />
Xaiva biguttata (Risso, 1816) 75<br />
Xaiva mcleayi (Barnard, 1947) 76<br />
Subfamily POLYBIINAE Ortmann, 1893 76<br />
Genus Bathynectes Stimpson, 1871 76<br />
Bathynectes maravigna (Prestandrea, 1839), new combination .. 76<br />
* Bathynectes piperitus, new species 77<br />
Genus Liocarcinus Stimpson, 1871 83<br />
Liocarcinus arcuatus (Leach, 1814), new combination 84<br />
Liocarcinus corrugatus (Pennant, 1777), new combination 84<br />
Genus Macropipus Prestandrea, 1833 85<br />
Macropipus australis Guinot, 1961 85<br />
* Macropipus rugosus (Doflein, 1904) 86<br />
Subfamily PORTUNINAE Rafinesque, 1815 87<br />
Genus Callinectes Stimpson, 1860 87<br />
Key to <strong>West</strong> <strong>African</strong> Species of Callinectes 88<br />
* Callinectes amnicola (De Rochebrune, 1883), new combination 88<br />
* Callinectes marginatus (A. Milne Edwards, 1861) 92<br />
* Callinectes pallidus (De Rochebrune, 1883), new combination 95<br />
Genus Cronius Stimpson, 1860 98<br />
* Cronius ruber (Lamarck, 1818) 98<br />
Page
vj SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Genus Portunus Weber, 1795 100<br />
Portunus hastatus (Linnaeus, 1767) 101<br />
* Portunus inaequalis (Miers, 1881) 102<br />
* Portunus validus Herklots, 1851 103<br />
Portunus vocans (A. Milne Edwards, 1878) 107<br />
Genus Thalamita Latreille, 1829 107<br />
Thalamita poissonii (Audouin, 1826) 107<br />
Family GERYONIDAE Colosi, 1923 108<br />
Genus Geryon Kr0yer, 1837 109<br />
Geryon affinis A. Milne Edwards and Bouvier, 1894 110<br />
* Geryon maritae, new species 112<br />
Family PLATYXANTHIDAE Guinot, 1977 118<br />
Family XANTHIDAE MacLeay, 1838 118<br />
Genus Cataleptodius Guinot, 1968 120<br />
* Cataleptodius floridanus (Gibbes, 1850) 120<br />
Genus Coralliope Guinot, 1967 121<br />
Coralliope parvula (A. Milne Edwards, 1869) 121<br />
Genus Cycloxanthops Rathbun, 1897 122<br />
Cycloxanthops occidentalis (A. Milne Edwards, 1867) 122<br />
Genus Domecia Eydoux and Souleyet, 1842 122<br />
* Domecia acanthophora africana Guinot, 1964 122<br />
Genus Epixanthus Heller, 1861 123<br />
* Epixanthus hellerii A. Milne Edwards, 1867 123<br />
Genus Eriphia Latreille, 1817 124<br />
Eriphia verrucosa (Forskal, 1775) 124<br />
Genus Euryozius Miers, 1886 124<br />
Euryozius bouvieri (A. Milne Edwards, 1869) 125<br />
* Euryozius pagalu, new species 126<br />
Genus Eurypanopeus A. Milne Edwards, 1878 130<br />
* Eurypanopeus blanchardi (A. Milne Edwards, 1881) 130<br />
Genus Globopilumnus Balss, 1933 133<br />
Globopilumnus africanus (A. Milne Edwards, 1867) 133<br />
* Globopilumnus stridulans Monod, 1956 134<br />
Genus Glyptoxanthus A. Milne Edwards, 1879 135<br />
* Glyptoxanthus angolensis (De Brito Capello, 1866) 135<br />
Glyptoxanthus cavernosus (A. Milne Edwards, 1878) 135<br />
Glyptoxanthus corrosus (A. Milne Edwards, 1869) 135<br />
Genus Heteropanope Stimpson, 1858 136<br />
Heteropanope acanthocarpus Crosnier, 1967 136<br />
Heteropanope tuberculidens Monod, 1956 136<br />
Genus Leopoldius Serene, 1971 136<br />
* Leopoldius pisifer (MacLeay, 1838), new combination 136<br />
Genus Menippe de Haan, 1833 137<br />
Menippe nodifrons Stimpson, 1859 137<br />
Genus Microcassiope Guinot, 1967 138<br />
* Microcassiope minor (Dana, 1852) 138<br />
Page
NUMBER 306 Vll<br />
Genus Monodaeus Guinot, 1967 140<br />
Monodaeus couchii (Couch, 1851) 141<br />
Monodaeus rectifrons (Crosnier, 1967) 141<br />
* Monodaeus rouxi (Gapart, 1951) 142<br />
Genus Nanocassiope Guinot, 1967 142<br />
* Nanocassiope melanodactyla (A. Milne Edwards, 1867) 143<br />
Genus Nanopilumnus Takeda, 1974 145<br />
* Nanopilumnus boletifer (Monod, 1956) 145<br />
Genus Panopeus H. Milne Edwards, 1834 145<br />
* Panopeus africanus A. Milne Edwards, 1867 146<br />
Genus Paractaea Guinot, 1969 148<br />
* Paractaea margaritaria (A. Milne Edwards, 1867) 148<br />
Paractaea monodi Guinot, 1969 149<br />
* Paractaea rufopunctata africana Guinot, 1976 150<br />
Genus Paraxanthias Odhner, 1925 151<br />
Paraxanthias eriphioides (A. Milne Edwards, 1867) 151<br />
Genus Pilumnopeus A. Milne Edwards, 1863 151<br />
* Pilumnopeus africanus (De Man, 1902) 151<br />
Pilumnopeus caparti (Monod, 1956) 151<br />
Genus Pilumnus Leach, 1815 152<br />
Pilumnus hirtellus (Linnaeus, 1761) 152<br />
Pilumnus inermis A. Milne Edwards and Bouvier, 1894 152<br />
* Pilumnus perrieri A. Milne Edwards and Bouvier, 1898 153<br />
Pilumnus spinifer H. Milne Edwards, 1834 154<br />
* Pilumnus stebbingi Capart, 1951 154<br />
Genus Platychelonion Crosnier and Guinot, 1969 155<br />
Platychelonion planissimum Crosnier and Guinot, 1969 155<br />
Genus Platypodiella Guinot, 1967 155<br />
Platypodiella picta (A. Milne Edwards, 1869) 155<br />
Genus Pseudomedaeus Guinot, 1968 155<br />
* Pseudomedaeus africanus (Monod, 1956) 155<br />
Genus Xantho Leach, 1814 156<br />
Xantho incisus (Leach, 1814) 156<br />
Xantho pilipes A. Milne Edwards, 1867 156<br />
Xantho sexdentatus (Miers, 1881) 157<br />
Genus Xanthodius Stimpson, 1859 157<br />
* Xanthodius denticulatus (White, 1848) 157<br />
Xanthodius inaequalis faba (Dana, 1852) 158<br />
* Xanthodius inaequalis inaequalis (Olivier, 1791) 158<br />
Family GONEPLACIDAE MacLeay, 1838 159<br />
Subfamily CARCINOPLACINAE H. Milne Edwards, 1852 160<br />
Genus Carcinoplax H. Milne Edwards, 1852 160<br />
* Carcinoplax barnardi Capart, 1951 160<br />
Subfamily EURYPLACINAE Stimpson, 1871 161<br />
Genus Machaerus Leach, 1818 161<br />
* Machaerus atlanticus (Miers, 1881), new combination 162<br />
Page
vlll<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
* Machaerus oxyacantha (Monod, 1956), new combination 163<br />
Subfamily GONEPLACINAE MacLeay, 1838 163<br />
Genus Goneplax Leach, 1814 163<br />
Goneplax rhomboides (Linnaeus, 1758) 164<br />
Subfamily RHIZOPINAE Stimpson, 1858 165<br />
Genus Acidops Stimpson, 1871 165<br />
Acidops cessacii (A. Milne Edwards, 1878) 165<br />
Subfamily TYPHLOCARCINOPINAE Rathbun, 1909 165<br />
Genus Typhlocarcinodes Alcock, 1900 165<br />
* Typhlocarcinodes integrifrons (Miers, 1881) 165<br />
Family HEXAPODIDAE Miers, 1886 166<br />
Key to Genera of Hexapodidae 168<br />
Genus Hexapinus, new genus 169<br />
Hexapinus buchanani (Monod, 1956), new combination 171<br />
Genus Hexaplax Doflein, 1904 171<br />
Genus Hexapus de Haan, 1833 171<br />
Genus Lambdophallus Alcock, 1900 173<br />
Genus Paeduma Rathbun, 1897 173<br />
Genus Parahexapus Balss, 1922 175<br />
Parahexapus africanus Balss, 1922 175<br />
Genus Pseudohexapus Monod, 1956 176<br />
Pseudohexapus platydactylus Monod, 1956 176<br />
Genus Spiroplax, new genus 176<br />
Genus Stevea, new genus 177<br />
Genus Thaumastoplax Miers, 1881 177<br />
Thaumastoplax anomalipes Miers, 1881 179<br />
Genus Tritoplax, new genus 180<br />
Family PINNOTHERIDAE de Haan, 1833 181<br />
Subfamily ASTHENOGNATHINAE Stimpson, 1858 181<br />
Genus Asthenognathus Stimpson, 1858 181<br />
Asthenognathus atlanticus Monod, 1933 181<br />
Subfamily PINNOTHERINAE de Haan, 1833 182<br />
Genus Pinnotheres Bosc, 1802 182<br />
Pinnotheres conicola, new species 182<br />
Pinnotheres leloeuffi Crosnier, 1969 185<br />
Pinnotheres mccainae Schmitt, 1973 185<br />
Pinnotheres pinnotheres (Linnaeus, 1758) 187<br />
Pinnotheres pisum (Linnaeus, 1767) 187<br />
Pinnotheres tellinae, new species 187<br />
Pinnotheres sp. A 190<br />
Pinnotheres sp. B 190<br />
Pinnotheres sp. D 191<br />
Pinnotheres sp 191<br />
Family RETROPLUMIDAE Gill, 1894 191<br />
Family MICTYRIDAE Dana, 1851 191<br />
Family PALICIDAE Rathbun, 1898 191<br />
Page
NUMBER 306<br />
Genus Palicus Philippi, 1838 191<br />
* Palicus caromi (P. Roux, 1830) 191<br />
Family OCYPODIDAE Rafinesque, 1815 192<br />
Subfamily CAMPTANDRIINAE Stimpson, 1858 193<br />
Key to Genera of Camptandriinae 193<br />
Genus Calabarium, new genus 195<br />
Calabarium crinodytes, new species 196<br />
Genus Camptandrium Stimpson, 1858 199<br />
Genus Cleistostoma de Haan, 1833 200<br />
Genus Deiratonotus, new genus 201<br />
Genus Ecphantor, new genus 202<br />
Ecphantor modestus, new species 203<br />
Genus Ilyogynnis, new genus 206<br />
Genus Leipocten Kemp, 1915 207<br />
Genus Paracleistostoma De Man, 1895 208<br />
Genus Paratylodiplax Serene, 1974 209<br />
Genus Serenella, new genus 211<br />
Genus Telmatothrix, new genus 212<br />
Telmatothrix powelli, new species 213<br />
Genus Tylodiplax De Man, 1895 217<br />
Subfamily OCYPODINAE Rafinesque, 1815 217<br />
Genus Ocypode Weber, 1795 217<br />
Ocypode africana De Man, 1881 218<br />
* Ocypode cursor (Linnaeus, 1758) 219<br />
Genus Uca Leach, 1814 220<br />
* Uca tangeri (Eydoux, 1835) 221<br />
Family GRAPSIDAE MacLeay, 1838 225<br />
Subfamily GRAPSINAE MacLeay, 1838 226<br />
Genus Geograpsus Stimpson, 1858 226<br />
* Geograpsus lividus (H. Milne Edwards, 1837) 226<br />
Genus Goniopsis de Haan, 1833 227<br />
* Goniopsis pelii (Herklots, 1851) 227<br />
Genus Grapsus Lamarck, 1801 232<br />
* Grapsus grapsus (Linnaeus, 1758) 232<br />
Genus Pachygrapsus Randall, 1840 233<br />
* Pachygrapsus gracilis (De Saussure, 1858) 233<br />
* Pachygrapsus transversus (Gibbes, 1850) 234<br />
Genus Planes Bowdich, 1825 235<br />
Planes cyaneus Dana, 1851 235<br />
Planes minutus (Linnaeus, 1758) 236<br />
Subfamily PLAGUSIINAE Dana, 1851 237<br />
Genus Pennon Gistel, 1848 237<br />
* Pennon gibbesi (H. Milne Edwards, 1853) 238<br />
Genus Plagusia Latreille, 1804 238<br />
* Plagusia depressa (Fabricius, 1775) 239<br />
Subfamily SESARMINAE Dana, 1851 239<br />
Page
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Genus Cyclograpsus H. Milne Edwards, 1837 239<br />
* Cyclograpsus integer H. Milne Edwards, 1837 239<br />
Genus Metagrapsus H. Milne Edwards, 1853 240<br />
* Metagrapsus curvatus (H. Milne Edwards, 1837) 240<br />
Genus Sesarma Say, 1817 .' 241<br />
Subgenus Chiromantes Gistel, 1848 242<br />
Sesarma (Chiromantes) angolense De Brito Capello, 1864 243<br />
* Sesarma (Chiromantes) buettikoferi De Man, 1883 243<br />
Sesarma (Chiromantes) elegans Herklots, 1851 244<br />
Subgenus Perisesarma De Man, 1895 245<br />
Sesarma (Perisesarma) alberti Rathbun, 1921 245<br />
Sesarma (Perisesarma) huzardi (Desmarest, 1825) 245<br />
Sesarma (Perisesarma) kamermani De Man, 1883 247<br />
Subfamily VARUNINAE H. Milne Edwards, 1853 247<br />
Genus Brachynotus de Haan, 1833 247<br />
Brachynotus atlanticus Forest, 1957 247<br />
Genus Euchirograpsus H. Milne Edwards, 1853 247<br />
Euchirograpsus liguricus H. Milne Edwards, 1853 247<br />
Family GECARCINIDAE MacLeay, 1838 248<br />
Genus Cardisoma Latreille, 1828 249<br />
Cardisoma armatum Herklots, 1851 249<br />
Genus Gecarcinus Leach, 1814 250<br />
Gecarcinus weileri (Sendler, 1912) 250<br />
Family HAPALOCARCINIDAE Caiman, 1900 250<br />
Genus Neotroglocarcinus Fize and Serene, 1957 250<br />
Neotroglocarcinus balssi (Monod, 1956) 251<br />
Family HYMENOSOMATIDAE MacLeay, 1838 251<br />
Genus Elamena H. Milne Edwards, 1837 252<br />
Elamena (Trigonoplax) gordonae Monod, 1956 252<br />
Genus Hymenosoma Desmarest, 1825 252<br />
Hymenosoma orbiculare Desmarest, 1825 252<br />
Family MAJIDAE Samouelle, 1819 252<br />
Subfamily EPIALTINAE MacLeay, 1838 255<br />
Genus Acanthonyx Latreille, 1828 255<br />
Key to Eastern Atlantic Species of Acanthonyx 255<br />
Acanthonyx brevijrons A. Milne Edwards, 1869 256<br />
Acanthonyx depressifrons, new species 258<br />
Acanthonyx lunulatus (Risso, 1816) 260<br />
* Acanthonyx minor, new species 261<br />
Subfamily INACHINAE MacLeay, 1838 263<br />
Key to Eastern Atlantic Genera of Inachinae 265<br />
Genus Achaeus Leach, 1817 266<br />
Key to Eastern Atlantic Species of Achaeus 268<br />
* Achaeus buderes, new species 269<br />
Achaeus cranchii Leach, 1817 271<br />
* Achaeus foresti Monod, 1956 271<br />
Page
NUMBER 306 XI<br />
Achaeus monodi (Capart, 1951) 272<br />
Achaeus trifalcatus Forest and Guinot, 1966 272<br />
* Achaeus turbator, new species 272<br />
Genus Calypsachaeus, new genus 275<br />
* Calypsachaeus calypso (Forest and Guinot, 1966), new combination<br />
275<br />
Genus Capartiella, new genus 277<br />
* Capartiella longipes (Capart, 1951), new combination 278<br />
Genus Dorhynchus Thomson, 1873 279<br />
Dorhynchus thomsoni Thomson, 1873 281<br />
Genus Ergasticus Studer, 1883 281<br />
Ergasticus clouei Studer, 1883 281<br />
Genus Inachus Weber, 1795 282<br />
Key to Tropical <strong>West</strong> <strong>African</strong> Species of Inachus 282<br />
Inachus aguiarii De Brito Capello, 1876 283<br />
* Inachus angolensis Capart, 1951 283<br />
* Inachus biceps, new species 285<br />
Inachus dorsettensis (Pennant, 1777) 287<br />
* Inachus grallator, new species 287<br />
Inachus guentheri (Miers, 1879) 291<br />
Inachus leptochirus Leach, 1817 291<br />
* Inachus nanus, new species 291<br />
Inachus phalangium (Fabricius, 1775) 293<br />
Inachus thoracicus Roux, 1830 294<br />
Genus Macropodia Leach, 1814 294<br />
Key to Tropical <strong>West</strong> <strong>African</strong> Species of Macropodia 295<br />
Macropodia doracis, new species 295<br />
* Macropodia gilsoni (Capart, 1951) 297<br />
* Macropodia hesperiae, new species 298<br />
Macropodia intermedia Bouvier, 1940 300<br />
Macropodia longicornis (A. Milne Edwards and Bouvier, 1899) 300<br />
Macropodia longipes (A. Milne Edwards and Bouvier, 1899) ... 300<br />
* Macropodia macrocheles (A. Milne Edwards and Bouvier, 1898) 301<br />
* Macropodia spinulosa (Miers, 1881) 301<br />
* Macropodia straeleni Capart, 1951 303<br />
Genus Stenorhynchus Lamarck, 1818 304<br />
* Stenorhynchus lanceolatus (Brulle, 1837) 304<br />
Subfamily MAJINAE Samouelle, 1819 307<br />
Genus Maja Lamarck, 1801 307<br />
Maja crispata Risso, 1827 307<br />
Maja goltziana d'Oliveira, 1888 307<br />
Maja squinado (Herbst, 1788) 307<br />
Subfamily PISINAE Dana, 1851 307<br />
Genus Apiomithrax Rathbun, 1897 307<br />
Apiomithrax bocagei (Osorio, 1887) 308<br />
* Apiomithrax violaceus (A. Milne Edwards, 1867) 309<br />
Page
xjj SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Genus Eurynome Leach, 1814 311<br />
* Eurynome aspera (Pennant, 1777) 311<br />
* Eurynome parvirostris Forest and Guinot, 1966 312<br />
Genus Herbstia H. Milne Edwards, 1834 312<br />
Key to Species of Adult Herbstia from <strong>West</strong> Africa 312<br />
* Herbstia condyliata (Fabricius, 1787) 312<br />
* Herbstia nitida, new species 315<br />
Herbstia rubra A. Milne Edwards, 1869 317<br />
Genus Micropisa Stimpson, 1858 318<br />
Micropisa ovata Stimpson, 1858 318<br />
Genus Pisa Leach, 1814 318<br />
* Pisa armata (Latreille, 1803) 318<br />
* Pisa calva Forest and Guinot, 1966 319<br />
* Pisa carinimana Miers, 1879 320<br />
Pisa nodipes (Leach, 1815) 321<br />
Pisa tetraodon (Pennant, 1777) 321<br />
Family MIMILAMBRIDAE Williams, 1979 322<br />
Family PARTHENOPIDAE MacLeay, 1838 322<br />
Subfamily AETHRINAE Dana, 1851 322<br />
Genus Heterocrypta Stimpson, 1871 322<br />
* Heterocrypta maltzami Miers, 1881 322<br />
Genus Sakaila, new genus 324<br />
* Sakaila africana, new species 325<br />
Subfamily PARTHENOPINAE MacLeay, 1838 327<br />
Genus Daldorfia Rathbun, 1904 327<br />
Daldorfia bouvieri (A. Milne Edwards, 1869) 327<br />
Genus Parthenope Weber, 1795 327<br />
* Parthenope expansa (Miers, 1879) 328<br />
* Parthenope massena (Roux, 1830) 330<br />
Parthenope miersii (A. Milne Edwards and Bouvier, 1898) 331<br />
* Parthenope notialis, new species 331<br />
Genus Solenolambrus Stimpson, 1871 336<br />
* Solenolambrus noordendei (Capart, 1951) 336<br />
Appendix I: Station Data 337<br />
Appendix II: Gazetteer 342<br />
Addendum 348<br />
Genus Lillyanella, new genus 348<br />
Lillyanella plumipes, new species 349<br />
Addendum to Key 352<br />
Literature Cited 353<br />
Page
<strong>West</strong> <strong>African</strong> <strong>Brachyuran</strong> <strong>Crabs</strong><br />
(Crustacea: Decapoda)<br />
Introduction<br />
Raymond B. Manning<br />
and L. B. Holthuis<br />
This report is based primarily on collections<br />
made in the Gulf of Guinea in 1964 and 1965<br />
during two cruises of the University of Miami<br />
Research Vessel John Elliott Pilhbury. Material<br />
from the Pillsbury collections has been supplemented<br />
by other brachyuran crabs from <strong>West</strong><br />
Africa in the collections of the Rijksmuseum van<br />
Natuurlijke Historie, Leiden, and the National<br />
Museum of Natural History, <strong>Smithsonian</strong> <strong>Institution</strong>,<br />
Washington, D.C.<br />
Our original intent was to prepare a list of the<br />
Brachyura obtained during the two Pillsbury<br />
cruises, a job that we anticipated would be quite<br />
simple, as the <strong>West</strong> <strong>African</strong> <strong>Brachyuran</strong> fauna<br />
has been investigated in detail in the last 30 years.<br />
In 1951, A. Capart published a beautifully illustrated<br />
volume on the <strong>West</strong> <strong>African</strong> crabs collected<br />
by the Belgian Oceanographic Expedition, 1948-<br />
1949, aboard the Noordende III. Capart's study was<br />
overshadowed by the publication in 1956 by Th.<br />
Monod of a review of <strong>West</strong> <strong>African</strong> crabs.<br />
Monod's work, a monumental compilation of<br />
information on <strong>West</strong> <strong>African</strong> Brachyura, was illustrated<br />
with almost 900 figures. It remains the<br />
basic tool in the study of <strong>West</strong> <strong>African</strong> crabs.<br />
Raymond B. Manning, Department of Invertebrate Zoology, National<br />
Museum of Natural History, <strong>Smithsonian</strong> <strong>Institution</strong>, Washington,<br />
D.C. 20560. L. B. Holthuis, Rijksmuseum van Natuurlijke Historie,<br />
Postbus 9517, 2300 RA Leiden, The Netherlands.<br />
These reports were soon followed by others<br />
based on smaller collections, those by M. Rossignol<br />
(1957 and 1962) on decapods from the Congo<br />
region and by D. Guinot and A. Ribeiro (1962)<br />
on collections from the Cape Verde Islands and<br />
Angola, as well as several other smaller reports.<br />
Material collected by the Calypso in 1956 from<br />
<strong>West</strong> <strong>African</strong> localities between Spanish Sahara<br />
and Gabon on the mainland, as well as from the<br />
offshore islands of the Gulf of Guinea, Principe,<br />
Sao Tome, and Annobon, formed the basis of an<br />
important report by J. Forest and D. Guinot in<br />
1966.<br />
Realizing that the <strong>West</strong> <strong>African</strong> tropical brachyuran<br />
fauna was one of the best known in the<br />
world, we anticipated few difficulties in preparing<br />
a report on the materials collected by the Pillsbury.<br />
As our study progressed, however, we continually<br />
encountered problems that demonstrated that<br />
our original optimism was largely unfounded.<br />
Whenever comparative study material was<br />
available in our collections, we compared specimens<br />
from <strong>West</strong> <strong>African</strong> localities with those from<br />
other areas. Several of the common species in the<br />
Gulf of Guinea have in the past been identified<br />
with European-Mediterranean species. Thus<br />
Ethusa mascarone, Inachus dorsettensis, Macropodia rostrata,<br />
and Parthenope macrochelos, among others, all<br />
have been included in the tropical fauna by<br />
Monod (1956), as well as other authors. In the
case of these four species, direct comparison of<br />
material from northern and southern areas has<br />
demonstrated that the southern forms are distinct;<br />
southern counterparts of three of these are<br />
described as new, and a name is resurrected from<br />
synonymy for the fourth species.<br />
Similarly, direct comparison of material from<br />
both sides of the Atlantic yielded surprising results.<br />
In one case, that of one species of Ranilia,<br />
such a comparison showed that populations on<br />
each side of the Atlantic were conspecific whereas<br />
they had previously been considered to be distinct.<br />
Other comparisons, based on species in<br />
Eurypanopeus, Goniopsis, Callinectes, and Bathynectes,<br />
resulted in our recognition of distinct species on<br />
each side of the Atlantic.<br />
Two family-group taxa, the family Hexapodidae<br />
and the subfamily Camptandriinae of the<br />
family Ocypodidae, had to be surveyed at the<br />
generic level in order to determine the status of<br />
the <strong>West</strong> <strong>African</strong> species. These revisions are incorporated<br />
herein. In addition, a key to the<br />
subfamilies and genera of family Dorippidae is<br />
presented.<br />
We also encountered unanticipated problems<br />
in various aspects of nomenclature: Citations of<br />
brachyuran family and subfamily names vary<br />
widely in the existing literature and often are<br />
incorrect; information on type-species and original<br />
citations of genera is widely scattered and has<br />
never been compiled for the Eastern Atlantic<br />
species; and numerous older names in the literature<br />
have never been properly identified.<br />
Finally, there is no single reference work containing<br />
at least a list of all of the Eastern Atlantic<br />
Brachyura. The Scandinavian species have been<br />
dealt with by M. Christiansen (1969) in a wellprepared<br />
handbook; the British species have been<br />
studied by R. W. Ingle (1980) of the British<br />
Museum (Natural History); the Spanish and Portuguese<br />
species were the subject of the most upto-date<br />
review of European-Mediterranean crabs<br />
by the late R. Zariquiey Alvarez (1968); the Black<br />
Sea species were studied by Bacescu (1967); the<br />
French species were studied by Bouvier (1940) in<br />
a work in which the nomenclature is now com-<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
pletely out of date; the Adriatic species were<br />
reported by O. Pesta in 1918, but, again, all the<br />
names are not current; and the eastern Mediterranean<br />
species were listed by L. B. Holthuis and<br />
E. Gottlieb in 1958. The latter fauna contains<br />
several species that have entered and colonized<br />
the region from the Red Sea via the Suez Canal.<br />
About 300 species of Brachyura are known from<br />
the Eastern Atlantic in literature that is widely<br />
scattered.<br />
Thus our simple list of species taken off <strong>West</strong><br />
Africa by the Pillsbury changed into a guide to the<br />
Eastern Atlantic crab fauna, with major emphasis<br />
on that of tropical <strong>West</strong> Africa. We have listed<br />
all Eastern Atlantic species of Brachyura known<br />
to us, and we have included all references to <strong>West</strong><br />
<strong>African</strong> crabs published since 1956 that have<br />
come to our attention. This work, then, is basically<br />
an update of Monod's monumental work,<br />
based primarily on the collections made by the<br />
Pillsbury.<br />
In our accounts, the term "<strong>West</strong> <strong>African</strong>" or<br />
"tropical" is used to indicate tropical <strong>West</strong> Africa:<br />
the Cape Verde Islands and the <strong>African</strong> mainland<br />
from Mauritania southward to Angola, including<br />
the offshore islands of the Gulf of Guinea.<br />
"Extralimital" is generally used to describe more<br />
northern forms, including those that live in the<br />
Mediterranean. We have not attempted to deal<br />
with the poorly known fauna of South-<strong>West</strong><br />
Africa.<br />
In the systematic account herein, we include<br />
original citations for all 36 families of marine<br />
Brachyura, as well as references to synonyms at<br />
the family-group level. Twenty-nine families are<br />
represented in the Eastern Atlantic fauna and 26<br />
of these have representatives in the tropical fauna.<br />
The seven families of crabs not occurring in the<br />
Eastern Atlantic are the Belliidae, the Bythograeidae,<br />
the Mimilambridae, the Cyclodorippidae,<br />
the Mictyridae, the Platyranthidae, and the<br />
Retroplumidae. The three families of crabs occurring<br />
in the Eastern Atlantic but not represented<br />
in the tropical fauna are the Cancridae,<br />
the Cymonomidae, and the Homolodromiidae.<br />
Original citations are provided for all 146 genera
NUMBER 306<br />
now known from the Eastern Atlantic, along with<br />
an indication of their type-species and gender;<br />
there are representatives of 120 genera in the<br />
tropical fauna. The Eastern Atlantic fauna comprises<br />
about 300 species, 218 of which have been<br />
recorded from localities between Mauritania and<br />
Angola.<br />
Sixteen new genera, six of which are extralimital,<br />
and 24 new species from <strong>West</strong> Africa are<br />
recognized.<br />
Most of the 300+ Eastern Atlantic species are<br />
indigenous to the Eastern Atlantic. Fifteen species<br />
have been introduced into the Eastern Mediterranean<br />
through the Suez Canal and apparently<br />
have become established there: Atergatis rosens,<br />
Charybdis helleri, C. longicollis, Eucrate crenata, Heteropanope<br />
laevis, Hyastenus hilgendorfi, Ixa monodi,<br />
Leucosia signata, Myra fugax, Notopus dorsipes, Pilumnopeus<br />
vauquelini, Pilumnus hirsutus, Portunus pelagicus,<br />
Sphaerozius nitidus, and Thalamita poissonii. Five<br />
other species have been introduced from other<br />
areas: Callinectes sapidus, Eriocheir sinensis, Neopanope<br />
sayi, Pilumnoides perlatus, and Rhithropanopeus<br />
harrisii.<br />
Several other species have been introduced into<br />
the Eastern Atlantic but apparently have not<br />
become established there. Catta (1876) reported<br />
the following species taken from a vessel in Marseilles<br />
harbor: Pachygrapsus advena, new species (=<br />
P. transversus), Nautilograpsus minutus (= Planes minutus),<br />
Plagusia squamosa (= P. depressd), and P.<br />
tomentosa (= P. chabrus (Linnaeus, 1758)); the first<br />
three are included in the <strong>West</strong> <strong>African</strong> fauna. De<br />
Man (1913) found Menippe convexa Rathbun,<br />
1893, Leptodius voeltzkowi Lenz, 1905, Pilumnus longicomis<br />
Hilgendorf, 1878, Pilumnus malardi De<br />
Man, 1913 (= Parapilumnus malardi), and Pilumnus<br />
truncatospinosus De Man, 1913 (= Parapilumnus truncatospinosus)<br />
in the harbor of St. Vaast-la-Hougue,<br />
Normandy, France, where they were obtained<br />
from barnacles attached to a ship coming from<br />
Madagascar.<br />
The following species have been erroneously<br />
recorded from <strong>West</strong> Africa: Chlorodiella longimana,<br />
Hepatus princeps, Libinia erinacea, Metopograpsus mes-<br />
sor, Notolopas brasiliensis, Ocypode ceratophthalmus, O.<br />
quadrata, Pilumnoides hassleri, Platychirograpsus spectabilis,<br />
Pseudograpsus elongatus, Rochinia gracilipes,<br />
Sesarma roberti, and Uca burgersi.<br />
In addition, there are several species described<br />
or recorded from Eastern Atlantic localities based<br />
on incorrectly labeled material, such as Portunus<br />
sanguinolentus (Herbst, 1783), reported from the<br />
Adriatic by Pesta (1918:458). These species are<br />
not included in our lists of extralimital species<br />
under each family.<br />
COLLECTIONS STUDIED.—Most of the specimens<br />
reported here were collected during two cruises<br />
of the research vessel John Elliot Pillsbury of the<br />
Rosenstiel School of Marine and Atmospheric<br />
Science, University of Miami, in 1964 and 1965.<br />
A narrative of the cruise was given by Voss (1966)<br />
and Bayer (1966), who summarized dredging and<br />
trawling records. Cruise tracks of the Pillsbury off<br />
<strong>West</strong> Africa are shown in Figure 87. Species taken<br />
by the Pillsbury are marked with an asterisk (*) in<br />
the discussion sections under each family heading<br />
and in the headings of the species accounts. The<br />
Pillsbury collections have been deposited in the<br />
Rijksmuseum van Natuurlijke Historie, Leiden<br />
(L) and the National Museum of Natural History,<br />
<strong>Smithsonian</strong> <strong>Institution</strong>, Washington, D.C. (W).<br />
The Pillsbury collections are quite important.<br />
They are slightly smaller than those taken by the<br />
Calypso, containing the representatives of 98<br />
rather than 108 species (Forest and Guinot, 1966:<br />
27). Although this represents less than half of the<br />
known <strong>West</strong> <strong>African</strong> crab fauna, 16 of the 24 new<br />
species recognized here are based on specimens<br />
taken by the Pillsbury. A smaller collection from<br />
off Gabon, taken during an exploratory cruise of<br />
the U. S. Fish and Wildlife Service vessel Geronimo<br />
also is reported on here. All of the material from<br />
the Geronimo collections has been deposited at<br />
Washington. The bulk of the brachyuran collection<br />
made off southern Angola and northern<br />
South-<strong>West</strong> Africa by the U. S. Fish and Wildlife<br />
Service vessel Undaunted and reported upon by<br />
Crosnier (1970) has been deposited at Leiden and<br />
is reported here.<br />
In the species accounts, station data for the
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
collections made by the Pillsbury, Geronimo, andcitations,<br />
not an exhaustive list of references.<br />
Undaunted are abbreviated to station number, Where appropriate, we have indicated which<br />
depth in meters, and observed bottom type. Com- family-group names are on the Official List of<br />
plete station data and a list of brachyuran species Family Group Names in Zoology established by the<br />
taken at each station are given in Appendix I. International Commission on Zoological Nomen-<br />
Smaller collections from the northwestern coast clature (ICZN) by stating "name 00 on Official<br />
of Africa by the Rijksmuseum van Natuurlijke List."<br />
Historie aboard the research vessel Onversaagd also In each family account we have included a<br />
are reported under "Material Examined" sections paragraph entitled "Eastern Atlantic Genera," in<br />
of the appropriate species discussion. Many of the which we state how many genera of that family<br />
species collected during the Onversaagd cruises do occur in the area. Extralimital or nontropical<br />
not occur in the tropical fauna and will be studied<br />
and possibly reported upon separately. In addition,<br />
the collections at Washington include series<br />
of decapods from Liberia donated by G. C.<br />
Miller, then with the U. S. Fish and Wildlife<br />
Service; material from Ghana donated by G. W.<br />
Bane, then with Cornell University; and several<br />
collections from Gulf of Guinea, donated by A.<br />
Crosnier, then with the Centre ORSTOM,<br />
Pointe-Noire.<br />
genera are then listed along with their original<br />
citations, an indication of their type-species and<br />
genders, and a statement that they are on the<br />
Official List, as appropriate. We have not tried to<br />
cite synonyms of extralimital genera.<br />
That paragraph is followed by one entitled<br />
"Eastern Atlantic Species," in which the numbers<br />
of species occurring in that region are given. This<br />
is followed by a comparison of the names of<br />
tropical species, as cited by Monod (1956), com-<br />
In both Washington and Leiden are a few lots pared to the current names; in several families,<br />
from the Travailleur and Talisman collections, ac- notably the Xanthidae and the Majidae, there<br />
quired on exchange from the Museum national have been numerous name changes since 1956. In<br />
d'Histoire naturelle, Paris. The <strong>Smithsonian</strong> re- those lists, the names used by Monod are given<br />
ceived representatives of 80 species from those in the same order in which he cited them, to<br />
collections in 1899 through E.-L. Bouvier. make it easier to work from our paper back to<br />
FORMAT CONSIDERATIONS.—We have adopted a Monod's. Also in those lists, species taken by the<br />
fairly rigid format in order to ensure that our Pillsbury are marked with an asterisk (*). Those<br />
accounts are comparable at the family, genus, lists are then followed by a list, in alphabetical<br />
and species levels.<br />
order by genus and species, of extralimital species,<br />
At the family level, we have first given the along with a brief comment on their range and<br />
original citation for each family of marine brach- one or more pertinent references. These two inyuran<br />
crabs; the freshwater families have been troductory paragraphs may be followed by a<br />
excluded. In addition to the original citation we<br />
have included original citations to all synonyms<br />
that we could find at the family-group level,<br />
without indicating necessarily which of these are<br />
now in current use as family-group categories.<br />
Thus in the Portunidae, for example, we have<br />
cited 20 different family-group taxa that have<br />
been recognized in the literature; only a few of<br />
these are now in current use. The list of synonyms<br />
for each family does not include erroneous spellings<br />
or subsequent uses of the family names; each<br />
of the family synonymies is a guide to original<br />
third entitled "Remarks" or by a key to genera.<br />
The families usually are arranged in the order<br />
in which they were treated in Monod. Within<br />
families, subfamilies, if recognized, are arranged<br />
alphabetically except in the Xanthidae where<br />
taxa are arranged alphabetically by genus and by<br />
species within each genus, genera are arranged<br />
alphabetically within subfamilies, and species are<br />
arranged alphabetically within genera.<br />
For genera represented in the tropical fauna<br />
we have included not only the original citation<br />
but references to all synonyms known to us. In
NUMBER 306<br />
the cases of some genera, such as Dorippe sensu abbreviated; full data and a list of species taken<br />
lato, Sesarma, and Uca, all of which are in need of at each station are in Appendix 1. Geographic<br />
revision, we have given a complete list of original localities in the sections on material and distri-<br />
citations of genus-group names in these groups as bution are corrected to approved spellings by the<br />
a guide to their future revision. We do not mean U. S. Board on Geographic Names as published<br />
to imply that the names so cited are to be consid- in their gazetteers. A list of localities, by country,<br />
ered as synonyms.<br />
with coordinates, is given in Appendix 2. Locali-<br />
Tropical species are treated in two ways. If ties in the literature accompanied by coordinates<br />
material from the tropical region was studied, we are not repeated in Appendix 2. Alternate spell-<br />
give an expanded account, with a synonymy, a ings often are given in the text. The names of<br />
list of synonyms, material examined (with a sep- most countries (e.g., Senegal, Guinea) have been<br />
arate paragraph for Pillsbury, Geronimo, and Un- anglicized. In the sections of ecology we have<br />
daunted collections), citations to a description, to summarized available information on habitat,<br />
a good illustration, and to an illustration of the depth distribution, and occurrences of ovigerous<br />
gonopod (used interchangeably with male pleo- females; much of this information has been taken<br />
pod and first male pleopod), an expanded section from the recent literature, that is since 1956.<br />
on biology, and an expanded section on distri- In the text, author and date citations accombution.<br />
If no tropical material is available, we panying a name (e.g., Dehaanius MacLeay, 1838)<br />
have given an abbreviated account, with refer- are considered to be part of the name, not a<br />
ences to Capart (1951) and Monod (1956), and bibliographic citation. References to Monod<br />
subsequent references, with parenthetic remarks without a date always refer to the monograph of<br />
on origin of the material reported in the litera- <strong>West</strong> <strong>African</strong> crabs published by Monod in 1956.<br />
ture, followed by a brief statement on distri- Although we have tried to make our lists of<br />
bution.<br />
family citations, generic names and synonyms,<br />
In our synonymies we have tried to include and species synonyms as complete as possible, we<br />
references to Capart (1951) and Monod (1956), realize that these lists are by no means exhaustive.<br />
all subsequent references based on <strong>West</strong> <strong>African</strong> We would appreciate having any omissions or<br />
specimens, and a few minor references overlooked errors brought to our attention.<br />
by Monod. We have not duplicated the extensive In some accounts in the literature based on<br />
synonymies given by Monod, except in the case expeditions sponsored by the French in the latter<br />
of new species, where complete synonymies are half of the 19th century (A. Milne Edwards and<br />
given and in cases where the name has been Bouvier, 1894, 1899, 1900), coordinates were<br />
changed since 1956. Original citations for all based on the Paris observatory, not Greenwich,<br />
species then reported were given by Monod in his which is 2° 20' <strong>West</strong> of Paris. Later (Bouvier,<br />
synonymies. The separate list of synonyms may 1922), these coordinates were based on Green-<br />
seem redundant, but even though the nomenclawich. In our text we have corrected the earlier<br />
ture of the Eastern Atlantic brachyurans is rela- observations from Paris to Greenwich; to avoid<br />
tively stable, current studies are demonstrating errors, we often have given both, so that a coor-<br />
the existence of previously unrecognized species dinate may be cited as: 05°10'N, 05°05'W of<br />
there, and many older names are available in the Paris (= 02°45'W of Greenwich).<br />
literature dealing with the European-Mediterra- REPOSITORIES AND ABBREVIATIONS.—The bulk<br />
nean fauna. In this region older names must be of the materials reported herein have been depos-<br />
searched for carefully before new species are ited in two institutions: The Rijksmuseum van<br />
named.<br />
Natuurlijke Historie, Leiden, The Netherlands,<br />
In the sections on material based on the Pills- abbreviated to "L" in the material sections and<br />
bury, Geronimo, and Undaunted collections, data are often referred to as "Leiden" in the text. Registry
numbers (Crust. D. 000) are for the Decapod<br />
collection at Leiden. The National Museum of<br />
Natural History, <strong>Smithsonian</strong> <strong>Institution</strong>, Washington,<br />
D. C, abbreviated to "W" in the material<br />
and referred to as "Washington" or "USNM" in<br />
the text. Catalog numbers for material at Washington<br />
are given under the acronym USNM,<br />
referring to catalogs established under the former<br />
U. S. National Museum. For new species deposited<br />
in these collections, registry or catalog numbers<br />
are given only for holotypes.<br />
A few specimens from other museums also have<br />
been studied. These are cited as follows:<br />
BM British Museum (Natural History), London<br />
MP Museum national d'Histoire naturelle, Paris<br />
(Paris Museum)<br />
ZMB Zoologisches Museum, Berlin<br />
A few abbreviations have been used throughout<br />
the text:<br />
cb<br />
cl<br />
cm<br />
fm<br />
juv<br />
m<br />
mm<br />
ov<br />
carapace breadth<br />
carapace length<br />
centimeter<br />
fathom<br />
juvenile<br />
meter<br />
millimeter<br />
ovigerous<br />
ACKNOWLEDGMENTS.—We are indebted to Gilbert<br />
L. Voss, Rosenstiel School of Marine and<br />
Atmospheric Science, University of Miami, for<br />
inviting us to participate in the Pillsbury cruises<br />
and for making the material available for study.<br />
Participation in the cruises was supported by the<br />
National Geographic Society under grants to the<br />
University of Miami. Part of this study was supported<br />
by the National Geographic Society under<br />
grant NGS 1042 to one of us (RBM).<br />
R. W. Ingle, British Museum (Natural History),<br />
London, and J. Forest, Museum national<br />
d'Histoire naturelle, Paris, provided working<br />
space on several occasions; Ingle also read and<br />
commented on part of the manuscript. H. Zibrowius,<br />
Station Marine d'Endoume, Marseille,<br />
and H.-E. Gruner, Zoologisches Museum, Berlin,<br />
sent material to us. C. B. Powell, University of<br />
Port Harcourt, Nigeria, collected and donated<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
specimens to the Rijksmuseum van Natuurlijke<br />
Historie, Leiden and the National Museum of<br />
Natural History, <strong>Smithsonian</strong> <strong>Institution</strong>, Washington,<br />
D. C.<br />
C. Froglia, Laboratorio di Tecnologia della<br />
Pesca, Ancona, brought an obscure paper by N.<br />
Prestandrea and the account of Portunus Maravigna<br />
to our attention. P. Le Loeuff, Antenne OR-<br />
STOM, Centre de Bretagne, Brest, provided us<br />
with a copy of a paper by C. A. Dias and J. F.<br />
Seita Machado that we were unable to obtain<br />
elsewhere. Angelo A. DiMauro, University of<br />
Connecticut, Torrington, provided information<br />
on the rediscovery of the type of Amorphopus cylindraceus<br />
in the Bell collection at Oxford University,<br />
England. C. F. Cowan, Cumbria, England, provided<br />
information on the dates of publication of<br />
Guerin's Iconographie. We thank M. Tiirkay, Natur-Museum<br />
Senckenberg, for bringing to our<br />
attention the account ofLambrus spinosissimus Osorio,<br />
1923, which we otherwise would have overlooked,<br />
and R. W. Ingle, British Museum (Natural<br />
History), for pointing out that Verany (1846)<br />
had introduced a generic and specific name for<br />
Brachynotus sexdentatus (Risso) (see synonymy for<br />
Brachynotus, p. 247).<br />
We thank two staff members of the Department<br />
of Invertebrate Zoology (Crustacea), National<br />
Museum of Natural History, <strong>Smithsonian</strong><br />
<strong>Institution</strong>, for helping with various aspects of<br />
preparation of the manuscript: Anne Cohen for<br />
help in checking the bibliography and Cynthia J.<br />
Hemmings for proofreading parts of the manuscript.<br />
The manuscript was read by our colleague<br />
at the <strong>Smithsonian</strong>, Fenner A. Chace, Jr., with<br />
his usual and welcome attention to detail. Lilly<br />
King Manning executed most of the original<br />
illustrations and prepared all of the illustrations<br />
for publication. Much of her work was done as a<br />
volunteer. Carolyn S. Hahn and Jack F. Marquardt<br />
of the <strong>Smithsonian</strong> Library helped to<br />
locate many obscure references.<br />
Family RANINIDAE de Haan, 1839<br />
RANINOIDEA de Haan, 1839:102 [emended to Raninidae by<br />
White, 1847a:56].
NUMBER 306<br />
NOTOPINAE Serene and Umali, 1972:22, 25, 29 [herein corrected<br />
to Notopodinae].<br />
EASTERN ATLANTIC GENERA.—Four, of which<br />
three, Cyrtorhina, Ranilia, and Raninoides, are represented<br />
by tropical species. The fourth genus,<br />
which occurs in the eastern Mediterranean, is<br />
Notopus de Haan (1841:138). Type-species: Cancer<br />
dorsipes Linnaeus, 1758, by monotypy; gender:<br />
masculine; name 1570 on Official List.<br />
EASTERN ATLANTIC SPECIES.—Four, three of<br />
which were recorded by Monod (1956) as follows:<br />
Name in Monod<br />
Ranilia atlantica<br />
Cyrtorhina granulosa<br />
Raninoides bouvieri<br />
Current Name<br />
Ranilia constrieta*<br />
Cyrtorhina granulosa<br />
Raninoides bouvieri*<br />
The fourth raninid occurring in the eastern<br />
Atlantic is Notopus dorsipes (Linnaeus, 1758): Eastern<br />
Mediterranean; an Indo-<strong>West</strong> Pacific species<br />
that has entered the Mediterranean via the Suez<br />
Canal, first recorded from the Mediterranean by<br />
Lewinsohn and Holthuis (1964).<br />
Subfamily NOTOPODINAE Serene and Umali,<br />
1972<br />
Genus Ranilia H. Milne Edwards, 1837<br />
Ranilia H. Milne Edwards, 1837:195 [type-species: Ranilia<br />
muricata H. Milne Edwards, 1837, by monotypy; gender:<br />
feminine].<br />
Raninops A. Milne Edwards, 1880:34 [type-species: Raninops<br />
constrictus A. Milne Edwards, 1880, by subsequent designation<br />
by Rathbun, 1937:17; gender: masculine].<br />
* Ranilia constricta (A. Milne Edwards, 1880)<br />
FIGURES 1, 2<br />
Raninops constrictus A. Milne Edwards, 1880:35.<br />
Notopus (Raninoides?) atlanticus Studer, 1883:17, pi. 1: figs.<br />
5a, b.<br />
Ranilia constricta.—A. Milne Edwards and Bouvier, 1923:302,<br />
pi. 1: figs. 11-13, pi. 3: figs. 2-5.—Rathbun, 1937:20, pi.<br />
4: fig. 5, pi. 5: figs. 1, 2.—Gomes Correa, 1970:2, pis. 1, 2,<br />
7: figs. 56-58.—Pequegnat, 1970:180.<br />
Notopus atlanticus.—Gurney, 1939:103 [listed].<br />
Ranilia atlantica.—Monod, 1956:47, 631, figs. 17, 18.—<br />
Longhurst, 1958:87.—Rossignol, 1962:113 [listed].—<br />
Gomes Correa, 1970:5 [discussion].<br />
Raninids.—Voss, 1966:50.<br />
Lyndeus.—Bayer, 1966:102.<br />
MATERIAL EXAMINED.—Pillsbury Material: Annobon:<br />
Sta 275, 9-69 m, rubble of coralline algae, 3
a<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 1.—Ranilia constricta (A. Milne Edwards), male cl ca. 15 mm, Pillsbury Sta 275: a, front;<br />
b, cheliped; c, fourth pereiopod; d, e, first pleopod of male;/, g, second pleopod of male.<br />
The largest specimen recorded in the literature is<br />
a male 41 mm long (Gomes Correa, 1970).<br />
REMARKS.—Both Monod (1956) and Gomes<br />
Correa (1970) have suggested that Ranilia atlantica<br />
(Studer, 1883) from Ascension Island and <strong>West</strong><br />
Africa was conspecific with R. constricta (A. Milne<br />
Edwards, 1880) from the western Atlantic, but,<br />
so far as we can determine, specimens from both<br />
sides of the Atlantic have not been compared<br />
directly. We have been able to compare the four<br />
specimens taken by the Pillsbury off Annobon with<br />
a female (cl 22.2 mm) from Bahia Honda, Cuba<br />
(USNM 48642), a female (cl 19.2 mm) from<br />
Barbados, 92-366 m (USNM 110223), a male (cl<br />
11.6 mm) and two females (cl 11.0-11.3 mm)<br />
from off Palm Beach, Florida, 55-73 m (USNM<br />
169698), and a male (cl 17.2 mm) and a female<br />
(cl 18.0 mm) from off Sombrero Light, Florida,<br />
92-110 m (USNM 169699), as well as with the<br />
syntypes of/?, atlantica from Ascension Island. We<br />
can find no significant differences between these<br />
specimens.<br />
The males from Annobon differ from the female<br />
in having slightly more divergent anterolateral<br />
spines on the carapace. Our female from<br />
Annobon resembles those from other localities in<br />
that the anterolateral spines do not diverge perceptibly<br />
from the lateral margins of the carapace.<br />
The illustrations published by A. Milne Edwards<br />
and Bouvier (1923) and the figures given<br />
by Rathbun (1937) differ in several respects, as<br />
pointed out by Monod (1956), who questioned<br />
whether the two specimens previously identified<br />
with R. constricta were conspecific. Gomes Correa<br />
(1970:5) commented on the resemblance of the<br />
figures given by Studer (1883) and Rathbun<br />
(1937).<br />
In all of our specimens, the rostral spine extends<br />
to or exceeds the adjacent frontal teeth by less<br />
than half its length; in no specimen is the rostrum<br />
so long as shown by A. Milne Edwards and<br />
Bouvier (1923, pi. 3: fig. 2) (Figure 2a). In addition,<br />
all of the specimens seen by us have a short<br />
but well-marked median carina on the rostrum.<br />
This carina is shown in the figures of A. Milne<br />
Edwards and Bouvier (1923, pi. 3: fig. 2), Rathbun<br />
(1937, pi. 5: fig. 1), and Studer (1883, pi. 1:<br />
fig 5a), but not by Monod (1956, figs. 17,18) or<br />
by Gomes Correa (1970, pi. 1: figs. 1,2).<br />
We have reproduced here the illustrations published<br />
by Studer (1883) and A. Milne Edwards<br />
and Bouvier (1923) (Figure 2). These figures,<br />
especially those by Studer, may be inaccessible to<br />
most workers. In addition, we have added<br />
sketches from our specimens (Figure 1), including<br />
a figure of the male pleopods.
NUMBER 306<br />
FIGURE 2.—Ranilia constricta (A. Milne Edwards): a, dorsal view (from A. Milne Edwards and<br />
Bouvier, 1923, pi. 3: fig. 2); b, cheliped (from A. Milne Edwards and Bouvier, 1923, pi. 1: fig.<br />
11); c, dorsal view; d, cheliped (from Studer, 1883, pi. 1: fig. 5a,b).<br />
BIOLOGY.—This species has been taken in <strong>West</strong><br />
<strong>African</strong> waters in depths between 40 and 69 m;<br />
off Ascension Island it occurs in 110 m. In the<br />
western Atlantic it has been recorded from the<br />
littoral zone to 366 or 481 m, usually in less than<br />
100 m. The record of 100-200 fm (183-366 m) in<br />
Pequegnat (1970) is somewhat puzzling. Pequegnat<br />
(1970:180) gave this depth in the species<br />
account, but in the station list (1970:6), published<br />
in the same volume, the station at which the<br />
species was collected, 65-A-9-15, is indicated as<br />
having a depth of 263 or 330 fm (481 or 604 m).<br />
The Pillsbury specimens were taken on rubble<br />
of coralline algae, whereas those from Sierra<br />
Leone were taken on muddy sand (Longhurst,<br />
1958). In the western Atlantic this species has<br />
been collected on a reef (Rathbun, 1937) and on<br />
a rocky reef off Palm Beach, Florida. Monod<br />
(1956) reported this species from the stomach<br />
contents of the fishes Syacium micrurum Ranzani<br />
and Trygon marmorata Steindachner. Nothing is<br />
known of the biology of this species. Ovigerous<br />
females have not been observed off <strong>West</strong> Africa.<br />
DISTRIBUTION.—Tropical Atlantic Ocean.<br />
Monod (1956), who reported material from Senegal,<br />
SW of Goree, 40 m, and between Joal and<br />
Sangomar, 42 m, as well as from Moyen Congo<br />
d<br />
[Congo], was the first to record the species from<br />
<strong>West</strong> Africa. Other records in the literature include:<br />
Eastern Atlantic. Sierra Leone: 07°43'N, 13°43'W, 40 m<br />
(Longhurst, 1958).<br />
Annobon Island: 01°24'S, O5°37'E to 01°24'S, 05°38'E,<br />
55-69 m (Bayer, 1966; Voss, 1966).<br />
Central Atlantic. Ascension Island: 60 fm (110 m) (Studer,<br />
1883).<br />
<strong>West</strong>ern Atlantic. Florida: Off Sombrero [reef], 47 fm<br />
(86 m) (A. Milne Edwards, 1880; A. Milne Edwards and<br />
Bouvier, 1923).<br />
Cuba: Bahia Honda (Rathbun, 1937).<br />
Gulf of Mexico: 23°OO'N, 86°48'W, 100-200 fm (183-366<br />
m) (or 481-604 m?) (Pequegnat, 1970).<br />
Brazil: Cabo Frio Island, littoral; Pai Island, Rio de<br />
Janeiro State, 20-30 m; off Guaritiba, Guanabara State, 35-<br />
40 m (all Gomes Correa, 1970).<br />
Subfamily RANININAE de Haan, 1839<br />
Genus Cyrtorhina Monod, 1936<br />
Cyrtorhina Monod, 1956:49 [type-species: Cyrtorhina granulosa<br />
Monod, 1956, by monotypy; gender: feminine].<br />
Cyrtorhina granulosa Monod, 1956<br />
Cyrtorhina granulosa Monod, 1956:49, figs. 19-31 [Ghana].—<br />
Gauld, 1960:68 [Ghana].—Monod, 1963, fig. 31 [no locality].—Forest<br />
and Guinot, 1966:42 [Principe].
10<br />
Cyrthorina granulosa.—Forest, 1959:23 [Principe; erroneous<br />
spelling].<br />
Cyrtorhyna granulosa.—Forest, 1959, pi. 2: fig. 1 [Principe;<br />
erroneous spelling].<br />
DISTRIBUTION.—Gulf of Guinea, from off<br />
Ghana and Ilha do Principe, in 5-6 and 12 m.<br />
Genus Raninoides H. Milne Edwards, 1837<br />
Raninoides H. Milne Edwards, 1837:196 [type-species: Ranina<br />
loevis Latreille, 1825, by monotypy; gender: masculine].<br />
* Raninoides bouvieri Capart, 1951<br />
Raninoides bouvieri Capart, 1951:59, fig. 17.—Monod, 1956:<br />
54, figs. 32-34.—Longhurst, 1958:87.—Forest, 1959:15.—<br />
Gauld, 1960:68.—Rossignol, 1%2:113.—Crosnier, 1964:<br />
35._Forest and Guinot, 1966:42.—Le LoeufT and Intes,<br />
1968, table 1.<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 68, 70 m, broken shell, 4d, 2$ (1 ov), 2 juv (L).<br />
Ivory Coast: Sta 62, 46 m, brown, branched and foliate<br />
Foraminifera, 1$ (L). Sta 64, 68 m, 16* (L).<br />
Ghana: Sta 28, 49-53 m, 1$ (L).<br />
Nigeria: Sta 241, 59-63 m, mud and shell, 2
NUMBER 306 11<br />
Liberia: 04°34.5'N, 08°31'W, 64 m (Forest and Guinot,<br />
1966).<br />
Ivory Coast: Off Fresco, 40 m (Le Loeuff and Intes, 1968).<br />
Ghana: Off Accra, 28-80 m (Gauld, 1960).<br />
Cameroon: No specific locality (Crosnier, 1964).<br />
Gabon: 00°38'25"S, 08°46'E, 5 m (Forest and Guinot,<br />
1966).<br />
Cabinda: W of Landana, 45 m (Rossignol, 1962).<br />
It has not previously been recorded from Nigeria<br />
or Dahomey, although both of these localities<br />
are well within its known range.<br />
Family HOMOLODROMIIDAE Alcock, 1899<br />
HOMOLODROMIDAE Alcock, 1899b: 127, 130 [corrected to<br />
Homolodromiidae by Stebbing, 1905:58; considered a<br />
subfamily of family Prosopidae von Meyer, 1860, by<br />
Glaessner, 1969:R486].<br />
EASTERN ATLANTIC GENERA.—One, which has<br />
not been recorded from tropical waters, is Dicranodromia<br />
A. Milne Edwards (1880:31). Typespecies:<br />
Dicranodromia ovata A. Milne Edwards,<br />
1880, by monotypy; gender: feminine.<br />
EASTERN ATLANTIC SPECIES.—One, Dicranodromia<br />
mahieuxii A. Milne Edwards, 1883. Bay of<br />
Biscay, Azores, and off the Sahara coast in depths<br />
between 454 and 1330 m (Zariquiey Alvarez,<br />
1968).<br />
Family DROMIIDAE de Haan, 1833<br />
DROMIACEA de Haan, 1833:ix [corrected to Dromiidae by<br />
Ortmann, 1892:541, 543; name 356 on Official List}.<br />
EASTERN ATLANTIC GENERA.—Two, Dromia and<br />
Sternodromia, both represented by species occurring<br />
off tropical <strong>West</strong> Africa.<br />
EASTERN ATLANTIC SPECIES.—Six, of which all<br />
but one are tropical. Three species were recorded<br />
by Monod (1956):<br />
Name in Monod Current Name<br />
Dromia caputmortuum<br />
Dromia nodosa [part]<br />
Dromia nodosa [part]<br />
Dromidiopsis spinirostris<br />
Dromia marmorea<br />
Dromia nodosa<br />
Dromia monodi*<br />
Sternodromia spinirostris*<br />
The fifth tropical species, D. bollorei, was named<br />
in 1974.<br />
The extralimital species is Dromia per sonata (Lin-<br />
naeus, 1758). Southern North Sea southward<br />
to Spanish Sahara, Mediterranean; sublittoral<br />
(Christiansen, 1969; Forest, 1974).<br />
Genus Dromia Weber, 1795<br />
Dromia Weber, 1795:92 [type-species: Cancer personatus Linnaeus,<br />
1758, by subsequent designation under the Plenary<br />
Powers of the International Commission on Zoological<br />
Nomenclature (Opinion 688 in Bulletin of Zoological Nomenclature,<br />
21:16-19); gender: feminine; name 1568 on Official<br />
List].<br />
Dromia bollorei Forest, 1974<br />
Dromia bollorei Forest, 1974:91, figs. Id, 2, 3d, 5, 6b, 7c,d, pi.<br />
2: figs. 1, 2, pi. 3: fig. 4, pi. 6: fig 1 [Mauritania and Ivory<br />
Coast].<br />
DISTRIBUTION.—Known only from off Mauritania<br />
and off the Ivory Coast in 100 m.<br />
Dromia marmorea Forest, 1974<br />
Dromia vulgaris.—Osorio, 1889:135, 139; 1898:193.—A.<br />
Milne Edwards and Bouvier, 1900:17, pi. 9: figs. 12-14<br />
[not fig. 15].—Rathbun, 1900a:300.—Balss, 1921:47.—<br />
Gordon, 1950:246 [part], figs. 24, 25.—Longhurst, 1958:<br />
87.—Buchanan, 1958:20. [Not Dromia vulgaris H. Milne<br />
Edwards, 1837 = D. personata (Linnaeus, 1758).]<br />
Hairy Brown Sea Crab.—Irvine, 1932:13, fig. 16.<br />
Dromia caput-mortuum.—Irvine, 1947:301, fig. 205.—Gauld,<br />
1960:68. [Not Cancer caputmortuum Linnaeus, 1767 = Dromia<br />
personata (Linnaeus, 1758).]<br />
Dromia caputmortuum.—Monod, 1956:59, figs. 35-51, 83a.—<br />
Rossignol, 1962:113. [Not Cancer caputmortuum Linnaeus,<br />
1767 = Dromia personata (Linnaeus, 1758).]<br />
Dromia nodosa.—Monod, 1956:65 [part]. [Not Dromia nodosa<br />
A. Milne Edwards and Bouvier, 1898.]<br />
Dromia personata.—Crosnier, 1967:321 [not Cancer personatus<br />
Linnaeus, 1758].<br />
Dromia marmorea Forest, 1974:79, 81, figs, lc, 2, 3b, 4d-f, j,k,<br />
5, pi. 1: figs. 2,4, pi. 3: fig. 2, pi. 4: fig. 7, pi. 5: figs. 3,4, pi.<br />
8: figs. 3,4.—Tiirkay, 1976b:61 [listed], 62.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Senegal: Dakar, 1949, R. Mauny, paratypes,<br />
1
12<br />
Ghana: No specific locality, 1966, F. R. Irvine, 1(5 (BM).<br />
Accra, 1938, F. R. Irvine, paratypes, 2? (1 ov) (BM).<br />
Gabon: Cap Lopez, 20 m, Dec 1956, A. Crosnier, paratype,<br />
19 ov (MP).<br />
Congo: Baie de Pointe-Noire, beach seine, Oct 1955, M.<br />
Rossignol, paratypes, 26 (MP). Baie de Pointe-Noire, widemouthed<br />
nets, Aug 1967, 1
NUMBER 306<br />
cellent figures of the present species, which show<br />
all the important details, and to which we have<br />
little to add. The only discrepancy that we can<br />
find is in the shape of the dactyli of the walking<br />
legs, which in Monod's figures 48 and 49 show<br />
minute ventral teeth, while in most of our specimens<br />
they bear very distinct spiniform teeth. The<br />
shape of these dactyli is very similar to that found<br />
in D. erythropus and D. personate<br />
Osorio (1889) reported "Dromia vulgaris" from<br />
Sao Tome and Ilha do Principe without giving<br />
morphological details of his material. In the same<br />
paper he also reported material of "Dromia spinirostris"<br />
from the islands; this latter material, as<br />
shown below, in all probability belongs to Dromia<br />
monodi. Therefore, it seems most likely that Osorio's<br />
Dromia vulgaris belongs to the other common<br />
shallow-water Dromia of the area, D. marmorea, the<br />
moreso as the latter species has often been confused<br />
with D. personata (= D. vulgaris and D.<br />
caputmortuum).<br />
Osorio (1898), Rathbun (1900a), and Balss<br />
(1921) just listed Osorio's 1889 records under<br />
Dromia vulgaris, and obviously did not see any new<br />
material. The references by Rathbun (1900a) and<br />
Balss (1921) to the occurrence of the species at St.<br />
Helena in all probability do not refer to the<br />
present species (see "Distribution," p. 14).<br />
The first original record of this species subsequent<br />
to that by Osorio (1889) is that by A. Milne<br />
Edwards and Bouvier (1900:17, pi. 9: figs. 12-<br />
14), who reported "Dromia vulgaris" from Senegal<br />
(16*) and from Porto da Praia (as La Praya), Cape<br />
Verde Islands (1 6 juv). Forest (1974), who examined<br />
both specimens, identified them with D.<br />
marmorea. A Milne-Edwards and Bouvier's description<br />
and illustrations of these <strong>West</strong> <strong>African</strong><br />
specimens of "Dromia vulgaris" contain some mistakes<br />
that have caused considerable confusion.<br />
The two French authors, in their account of the<br />
Talisman and Travailleur Brachyura reported two<br />
species of Dromia: Dromia vulgaris (1900:17) and D.<br />
nodosa (1900:18). The latter species was said in the<br />
text (1900:18) and in the explanation of the<br />
figures (1900:369) to be illustrated on plate 9:<br />
figs. 12-14, whereas Dromia vulgaris was said to be<br />
figured on plate 9: fig. 15. Actually, however, the<br />
reverse is true: plate 9: figs. 12-14 shows one of<br />
the two specimens of D. vulgaris, probably the<br />
larger one from Senegal, and figure 15 is based<br />
on the syntype of D. nodosa, which was figured<br />
again by Forest and Guinot (1966, fig. 2a). This<br />
mixup was the primary reason that Monod (1956)<br />
incorrectly interpreted D. nodosa.<br />
Irvine (1932) gave a short account and a recognizable<br />
figure of the present species, which he<br />
indicated as "Hairy Brown Sea Crab," based on<br />
material from the Gold Coast (Ghana), presumably<br />
from Accra. In a later publication, Irvine<br />
(1947) repeated this information and listed the<br />
species under the name Dromia caput-mortuum.<br />
According to Irvine (1947), this species is not<br />
considered to be edible in Ghana. Irvine also<br />
noted that the vernacular name of this crab in<br />
the Ga language of Ghana is "Tsitsikuntu," a<br />
name derived from the word "kuntu" for blanket<br />
and obviously referring to the wooly hair cover of<br />
the body.<br />
Gordon (1950:244-251), in her study of the<br />
morphology of the spermatheca in Dromiidae,<br />
figured and described the thoracic sternum of a<br />
female of "Dromia vulgaris" from Madeira. Forest<br />
(1974), who later examined the specimen, identified<br />
it with D. marmorea.<br />
Capart (1951:21, figs. 1, 3a) also reported<br />
"Dromia vulgaris" from <strong>West</strong> Africa. He mentioned<br />
a young female from Pulpito Bay, Rio de Oro<br />
(Spanish Sahara). Forest (1974), who examined<br />
the specimen, identified it with D. personata. Capart's<br />
illustrations were based on a specimen of<br />
D. personata from the English Channel, so that<br />
both his text and illustrations pertain to D. personata,<br />
not to the present species.<br />
The material reported upon by Sourie (1954a),<br />
Buchanan (1958), and Longhurst (1958) as<br />
Dromia vulgaris had been identified as such by<br />
Monod, and thus must be considered to be D.<br />
marmorea. As shown by the superb figures of<br />
"Dromia caputmortuum" in Monod's (1956) great<br />
work, the specimens that he considered to be that<br />
species (= D. vulgaris — D. personata) actually were<br />
D. marmorea.<br />
13
14<br />
It was Forest (1974) who first pointed out that<br />
the <strong>West</strong> <strong>African</strong> form is distinct from the Mediterranean<br />
and NE Atlantic D. personata, and Forest<br />
proposed the name Dromia marmorea for it, at<br />
the same time providing an excellent, well-illustrated<br />
description.<br />
BIOLOGY.—This species has been reported from<br />
depths between 0 and 96 m, with one doubtful<br />
record from 100 m, but more than 90% of the<br />
records are from depths of less than 42 m, and<br />
about 60% are from 20 m or less. Irvine (1947:<br />
302) remarked that the species "lives in deep<br />
water and is sometimes caught in bottom-nets a<br />
mile or more from land." It has been reported<br />
from the following types of bottom: mud (Crosnier,<br />
1967); very fine sand (Buchanan, 1958);<br />
sand and shells (Forest, 1974); muddy sand<br />
(Longhurst, 1958); rocks (Sourie, 1954a; Monod,<br />
1956; Forest, 1974). Monod (1956) reported a<br />
juvenile specimen from the hull of a ship. Sourie<br />
(1954a) classed this species as characteristic of the<br />
"hypobioses lapidicoles du sous-etage," while Buchanan<br />
considered it to belong to the active<br />
epifauna of the inshore fine sand community off<br />
Ghana.<br />
Osorio (1889) is the only author to report that<br />
this species carries a sponge. Sourie (1954a: 247)<br />
stated that it mostly bears didemnid or polycitorid<br />
ascidians. Two specimens from Bel-Air,<br />
Dakar, in the collection of the <strong>Smithsonian</strong> <strong>Institution</strong><br />
(USNM 152646) have balanids all over<br />
the central and anterior part of the carapace; if<br />
these specimens carried a sponge or ascidian it<br />
must have covered a small area on the posterior<br />
part of the body.<br />
Ovigerous females have been taken in September,<br />
November, and December (Monod, 1956;<br />
Forest, 1974).<br />
DISTRIBUTION.—Dromia marmorea is a <strong>West</strong> <strong>African</strong><br />
species, reported from the Azores, Madeira,<br />
and the Canary Islands south to Cabinda, and<br />
Sao Tome and Principe islands in the Gulf of<br />
Guinea. Monod (1956:62) reported two males<br />
and a female from "Cap Blanc, M. H. Routh<br />
coll., 1951 (B.M.)"; an examination of this lot in<br />
the British Museum revealed that Monod's local-<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
ity indication was erroneous, for the specimens<br />
actually originate from Gambia (see "Other Material"<br />
p. 11). The records of the species in the<br />
literature are as follows:<br />
Azores: Ilha do Muda [?], off Ilha das Flores, 22-30 m;<br />
Caldeira Inferno, Ilha do Faial, 10 m; Ponta Sao Diego [?],<br />
Ilha Terceira, 0-30 m; Ponta Delgada, 15-20 m, Caloura, 3<br />
m, and Ponta da Galera, 10-12 m, Ilha de Sao Miguel (all<br />
Forest, 1974).<br />
Madeira: No specific locality (Gordon, 1950; Forest, 1974;<br />
Tiirkay, 1976b). Funchal harbor (Tiirkay, 1976b).<br />
Canary Islands: Santa Cruz de La Palma, Isla de La<br />
Palma, 15 m; Playa de los Abrigos, Las Caletillas, 100 m<br />
(?), Puerto de la Cruz, intertidal, and Ensenada de Cristianos,<br />
intertidal, Isla de Tenerife; Playa Quemada, 2-3 m,<br />
and Arrecife, Isla de Lanzarote (all Forest, 1974).<br />
Cape Verde Islands: Porto da Praia (as La Praya), Sao<br />
Tiago, 10-30 m (A. Milne Edwards and Bouvier, 1900;<br />
Forest, 1974); Porto da Praia, Sao Tiago (as Porto Praia (I.<br />
Santiago)), the type-locality (Forest, 1974).<br />
Mauritania: No specific locality (Monod, 1956; Forest,<br />
1974).<br />
Senegal: No specific locality, in 80 m (A. Milne Edwards<br />
and Bouvier, 1900; Forest, 1974). Dakar (Monod, 1956;<br />
Forest, 1974). Goree Island, near Dakar, beach, 2-3 m and<br />
96 m; S of Goree, 33-35 m and 40 m (Monod, 1956; Forest,<br />
1974). Bel-Air, near Dakar, 5-10 m (Forest, 1974). Anse<br />
Bernard, Dakar; Goree, 23 m; Mbour, 25 m; Joal (Monod,<br />
1956).<br />
Gambia: Erroneously cited as Cap Blanc, Mauritania<br />
(Monod, 1956).<br />
Guinea-Bissau: 10°22'N, 16°22'W, 41 m (Forest, 1974).<br />
Guinea: 09°22'N, 13°42'W, 20-35 m; Guinea, 15-20 m;<br />
09°N, 13°50'W, 30 m (Monod, 1956). 09°20'N, 14°15'W,<br />
32 m (Forest, 1974).<br />
Sierra Leone: Off Freetown, 15 m (Forest, 1974).<br />
Ghana: Gold Coast (Irvine, 1932); same, 23-45 m<br />
(Longhurst, 1958). Near Accra, in deep water (Irvine, 1947);<br />
same (Monod, 1956); same, in 5.5-14.6 m (Buchanan, 1958);<br />
same, in shallow water to 25 m (Gauld, 1960); same (Forest,<br />
1974). Chorkor, near Accra, beach seine (Monod, 1956).<br />
Principe: No specific locality (Osorio, 1889, 1898).<br />
Sao Tome: Praia Lagarto (Osorio, 1889, 1898).<br />
Gabon: Cap Lopez, 20 m (Crosnier, 1967; Forest, 1974).<br />
Congo: Baie de Pointe-Noire, beach seine (Rossignol,<br />
1962; Crosnier, 1967; Forest, 1974). Off Pointe-Noire, 50 m<br />
(Crosnier, 1967; Forest, 1974).<br />
According to Forest (1974) the present species<br />
also occurs in Saint Helena, and he assigned<br />
material from there identified as Dromia vulgaris<br />
by Melliss (1875) and Cunningham (1910), as<br />
well as that reported upon as Dromia species? by
NUMBER 306<br />
Colman (1946) and as Dromia erythropus by Chace<br />
(1966), to the present species. The records by<br />
Rathbun (1900a) and Balss (1921) of Dromia vulgaris<br />
from St. Helena are not original, but based<br />
on Melliss' (1875) record. We have examined<br />
Chace's (1966) St. Helena material and found it<br />
impossible to confidently assign it to the present<br />
species, although it indeed is very similar, but so<br />
is it similar to Dromia erythropus and D. personata.<br />
Therefore, we have not included St. Helena in<br />
the range of D. marmorea. It seems possible that<br />
the St. Helena Dromia belongs to a species distinct<br />
from either D. marmorea and D. erythropus, but<br />
more material is needed to decide the status of<br />
that form; it also is possible, although less likely,<br />
that more than one species of Dromia occurs in St.<br />
Helena waters.<br />
* Dromia monodi Forest and Guinot, 1966<br />
FIGURE 3a, b<br />
Dromia spinirostris Osorio, 1889:136, 139; 1898:193—Balss,<br />
1921:47—Capart, 1951:23 [part, only specimens from A.S.<br />
141 and Mercator]. [Not Dromia spinirostris Miers, 1881 =<br />
Stemodromia spinirostris.]<br />
Dromia atlantica.—Rathhun, 1921:393, fig. 1, pi. 18: fig. 3<br />
[not Dromia atlantica Dofiein, 1904 = Stemodromia spinirostris<br />
(Miers, 1881).]<br />
Dromia nodosa.—Monod, 1956:65 [part], figs, 52-71, 83b.—<br />
Longhurst, 1958:87.—Gauld, 1960:68. [Not Dromia nodosa<br />
A. Milne Edwards and Bouvier, 1898.]<br />
Dromia monodi Forest and Guinot, 1966:43, fig. 1.—Crosnier,<br />
1967:321 [part, only the male specimens].—Le Loeuff and<br />
Intes, 1968:38, tables 1,5,9.—Forest, 1974:96, figs, le, 2,<br />
3f,g, 5, 7a,b, pi. 4: figs. 1-3, pi. 6: fig. 2.<br />
Dromia sp.—Forest and Guinot, 1966:46.—Forest, 1974:99.<br />
PDromia.—Maurin, 1968b, fig. 4.<br />
PDromia nodosa.—Maurin, 1968b:484, 489.<br />
Not Dromia nodosa.—Rossignol, 1962:113 [= Stemodromia spinirostris<br />
(Miers, 1881)].<br />
MATERIAL EXAMINED.—Pillsbury Material: Ghana:<br />
Sta 26, 27 m, shell bottom (scallops), 1$ ov (L).<br />
Nigeria: Sta 248, 33 m, 5c5, 5$ (2 ov) (L, W). Sta 250,<br />
24 m, brackish water, 19 (L).<br />
Other Material: Senegal: Dakar, 10 m, Dec 1951, E. Postel,<br />
1
16<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 3—Sternal grooves of females of various sizes. Dromia monodi Forest and Guinot, Pillsbury<br />
Sta 248: a, cb 14 mm, b, cb 38 mm. Sternodromia spinirostris (Miers), Pillsbury Sta 47: c, cb 21 mm.<br />
two ovigerous females, cl 14 and 18 mm, bridging<br />
nicely the existing gap. The smallest of these<br />
specimens has the sternal grooves rather straight<br />
and wide apart distally and each ends at the base<br />
of a tubercle; these tubercles are connected by a<br />
low wide ridge. The structure of the two tubercles<br />
and the ridge could very well be described as a<br />
strong transverse swelling, excavated in the middle.<br />
The specimen of 18 mm shows about the<br />
same situation, only the tubercles are less distinct<br />
and the ridge is absent, the surface between the<br />
tubercles being flush with the rest of the surface.<br />
In the ovigerous female of cl 47 mm the situation<br />
resembles that in the specimen of 18 mm, but<br />
here the tubercles are even less distinct as shown<br />
for a specimen 38 mm long in Figure 3b. We have<br />
not been able to find any consistent difference in<br />
the sharpness of the tubercles of the carpus of the<br />
cheliped in large and small, ovigerous and nonovigerous<br />
specimens, and believe that this character,<br />
to which Forest and Guinot themselves<br />
attached very little importance, indeed is of no<br />
taxonomic value here. We have come to the<br />
conclusion therefore that the specimens indicated<br />
by Forest and Guinot as Dromia sp. can safely be<br />
considered to belong to D. monodi. Forest (1974:<br />
100) considered it most likely that the specimens<br />
are precocious D. monodi, but did not exclude the<br />
possibility that they are a new species.<br />
An interesting feature of this species is the<br />
arrangement of the sternal grooves in the female.<br />
As already stated above, in the ovigerous females,<br />
whether large or small, these grooves end wide<br />
apart, each at the base of a low rounded tubercle,<br />
which in the small specimens is somewhat more<br />
distinct than in the large. From the genital opening<br />
the groove extends straight backward, parallel<br />
and rather close to the lateral margins of the<br />
thoracic sternum; the anterior ends of these<br />
grooves are definitely not turned inward. In the<br />
non-ovigerous females, again whether small or<br />
large, the tips of the grooves are distinctly curved<br />
inward towards each other, and no tubercles are<br />
visible; in these specimens the tubercle is replaced<br />
by a depression (Figure 3a, $ cb 14 mm; 3b, 9 cb<br />
38 mm). An excellent figure of the latter situation<br />
also is given by Rathbun (1921:393, fig. le).<br />
Forest and Guinot (1966) showed that the<br />
Dromia species from <strong>West</strong> Africa indicated by<br />
Monod (1956) as Dromia nodosa is specifically distinct<br />
from the type-specimen of the true Dromia<br />
nodosa A. Milne Edwards and Bouvier, 1898, and<br />
gave it the new name Dromia monodi. The source<br />
of the confusion was given by Forest and Guinot<br />
(1966:45) as "l'inexactitude du dessin ensemble<br />
(A. Milne Edwards et Bouvier, pi. IX; fig. 14),"<br />
evidently meaning figure 12, not 14; also Forest<br />
(1974:97) mentioned "le peu de fidelite du dessin<br />
publie par A. Milne Edwards et Bouvier (1900;<br />
pi. 9, fig. 12)." Examination of the text and
NUMBER 306<br />
figures published by A. Milne Edwards and Bouvier<br />
shows, however, that figure 12 is not inexact,<br />
but has only been given the wrong name. Comparing<br />
Forest and Guinot's figure 2a of the type<br />
of Dromia nodosa with A. Milne Edwards and<br />
Bouvier's (1900, pi. 9: fig. 15) of "Dromia vulgaris"<br />
it is clear that these two are made after the same<br />
specimen. What has happened, therefore, obviously<br />
is that A. Milne Edwards and Bouvier<br />
(1900) on their plate 9 gave as figures 12 to 14<br />
the entire animal and details of what in the text<br />
they called "Dromia vulgaris," and on figure 15 the<br />
type of Dromia nodosa in dorsal view, but that in<br />
the explanation of these figures (1900:369 and pi.<br />
9) they switched the names, and also in the text<br />
referred to the wrong figures. This is confirmed<br />
in the explanation (1900:369) of plate 9: figure<br />
15 ("Dromia vulgaris"), which is said to be of "un<br />
des petits exemplaires des lies du Cap-Vert,"<br />
which evidently refers to Dromia nodosa of which<br />
the authors had 5 specimens from the Cape Verde<br />
Islands, while in their material "Dromia vulgaris"<br />
was represented by only a single Cape Verde<br />
Islands specimen. A Milne Edwards and Bouvier's<br />
(1900, pi. 9: figs. 12-14) illustrations are<br />
actually made after a specimen they named<br />
Dromia vulgaris, probably the large male from<br />
Senegal, which was shown by Forest (1974:79) to<br />
belong to Dromia marmorea. It is not surprising<br />
therefore that Monod (1956) arrived at the wrong<br />
conclusion about the identity of Dromia nodosa,<br />
especially since A. Milne Edwards and Bouvier's<br />
(1900) figure 12 shows the second and third anterolateral<br />
tooth of the carapace quite close together<br />
as in Dromia monodi.<br />
As Forest and Guinot (1966:46) indicate, it is<br />
more likely that Osorio's (1889:136, 139) Dromia<br />
spinirostris from Sao Tome and Principe islands<br />
belongs to Dromia monodi rather than to Sternodromia<br />
spinirostris, because the latter species usually<br />
is found at greater depths than the 4 or 5 m from<br />
which Osorio reported his male. Monod (1956)<br />
referred the specimens listed by Balss (1921) under<br />
Dromia spinirostris to the present species, evidently<br />
because Balss did not find a median tubercle<br />
between the ends of the sternal grooves of<br />
his female. As Balss' specimens were collected at<br />
depths of 10 and 20 m, it seems highly likely that<br />
Monod's surmise is correct, even though the character<br />
of the sternal tubercle does not always hold<br />
good. Monod (1956) also referred the material<br />
that Odhner (1923) identified as Dromia spinirostris<br />
to the present species, but there are reasons to<br />
believe that Odhner's identification was correct<br />
(see p. 19).<br />
Rathbun's (1921) description and illustrations<br />
of what she thought to be Dromia atlantica Doflein<br />
leave no doubt that her material was actually D.<br />
monodi; already Monod (1956) pointed this out.<br />
Monod (1956) very extensively figured the<br />
present species and identified it with Dromia nodosa<br />
A. Milne Edwards and Bouvier, being followed<br />
in this by Longhurst (1958), Gauld (1960), Rossignol<br />
(1962), and possibly Maurin (1968b). Forest<br />
and Guinot (1966) pointed out that the present<br />
species is different from the true Dromia nodosa,<br />
and proposed the new name Dromia monodi for it,<br />
which name has been adopted by subsequent<br />
authors.<br />
Some of the specimens, mostly juveniles, that<br />
Monod (1956) identified as Dromia nodosa proved<br />
on examination to belong to neither that species<br />
nor to D. monodi. Forest (1974:79) found that the<br />
specimens indicated by Monod with the numbers<br />
1-2 (Mauritania), 5, 7, 12(1), 15, 16(4) (all from<br />
Goree), actually belong to Dromia marmorea. Examination<br />
of the specimen from Murray Town,<br />
Sierra Leone (Monod, 1956:69, no. 52), proved<br />
that it belongs to Stemodromia spinirostris.<br />
Crosnier (1967:321) assigned two males and<br />
two females to the present species. The females<br />
are both juvenile, one is from Dahomey, the other<br />
from "Pte Ste Clara" (= Cap Santa Clara), Gabon.<br />
The Gabon female had been reported on<br />
before by Rossignol (1962) under the name<br />
Dromia nodosa. The two females are now in the<br />
collection of the Museum national d'Histoire naturelle,<br />
Paris, and were examined by us. Although<br />
juveniles of Dromia monodi and Stemodromia spinirostris<br />
are usually difficult to distinguish, we believe<br />
that both specimens should be assigned to<br />
Stemodromia spinirostris rather than to Dromia mon-<br />
17
18<br />
odi. Forest (1974:101) also assigned the Dahomey<br />
specimen to Sternodromia spinirostris, but did not<br />
mention the Cap Santa Clara specimen. The two<br />
male specimens listed by Crosnier (1967), from<br />
Cap Lopez (Gabon) and Cabinda, are true Dromia<br />
monodi (Forest, 1974:96, and p. 15).<br />
Maurin (1968b:484, 489) reported Dromia nodosa<br />
from 200 m depth, dredged between Cabo<br />
Corbeiro and Cabo Blanco (= Cap Blanc), Spanish<br />
Sahara, and also from 60 and 70 m depth on<br />
the Bane d'Arguin, Mauritania. The species is<br />
not noted in Maurin's (1968a) more extensive<br />
treatise of the same area. As no morphological<br />
data are provided of the material, it is impossible<br />
to conclude whether the true Dromia nodosa or D.<br />
monodi is the species obtained by Maurin; the<br />
depth indications would indicate the former.<br />
Pechiiel-Loesche (1882) reported a specimen of<br />
Dromia from the "Banya" (= Crique Banjia, 00°<br />
14'N, 09°40'E), Gabon. As insufficient morphological<br />
details are provided by Pechiiel-Loesche,<br />
the identity of the species is uncertain, the description<br />
of its habits make it more likely that not<br />
Dromia but Phyllodorippe was meant (see p. 35).<br />
BIOLOGY.—As Forest and Guinot (1966)<br />
pointed out, this species occurs mostly between 0<br />
and 25 m depth. This is fully confirmed by the<br />
records below, all but one (60 m) of which are<br />
from less than 45 m and about 80% from 25 m or<br />
less.<br />
As to the bottom on which the species lives,<br />
this is given variously as green and black mud<br />
(Capart, 1951; Forest, 1974), "sable" (Osorio,<br />
1889), "sable vaseuse" (Le Loeuff and Intes,<br />
1968), "between sponges that surrounded Pinna"<br />
(Rathbun, 1921), sand, shelly sand and shelly<br />
mud (Longhurst, 1958), "roche, coraux" and<br />
"vase, coquilles" (Forest and Guinot, 1966),<br />
"fonds a Palythoa et Moigula, sable," "sable dur"<br />
(Monod, 1956). Specimens have been reported as<br />
collected with beach seines on sandy beaches<br />
(Monod, 1956; and p. 15).<br />
Like other species of Dromia the present species<br />
has been noted to carry sponges (Monod, 1956)<br />
and composite ascidians (Monod, 1956). Osorio<br />
(1889:136) reported on a specimen that carried a<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
piece of decaying wood, by which it was entirely<br />
covered, but "il ne marchait pas tout a son aise";<br />
it is possible that the collector mistook a sponge<br />
or ascidians for a piece of decaying wood.<br />
Ovigerous females have been collected in February,<br />
March, May, July, and October (Monod,<br />
1956; Pillsbury).<br />
DISTRIBUTION.—Dromia monodi has been reported<br />
from the west coast of Africa between<br />
Mauritania and Angola. The records from between<br />
Cabo Corbeiro and Cabo Blanco, Spanish<br />
Sahara, 200 m and Bane d'Arguin, Mauritania,<br />
60-70 m (both Maurin, 1968b) and Crique Banjia,<br />
Gabon, on the shore (Pechiiel-Loesche, 1882),<br />
discussed above, are excluded. Other records include<br />
the following:<br />
Mauritania: No specific locality (Forest, 1974).<br />
Senegal: Near Dakar, 15-20 m (type-locality) (Monod,<br />
1956; Forest and Guinot, 1966; Forest, 1974). SE of lie de la<br />
Madeleine, near Dakar, 35 m; near Dakar, 10 m and 34 m;<br />
Joal; Lagoba [?], 10-11 m (Monod, 1956; Forest, 1974). Bel-<br />
Air, near Dakar, 5-10 m (Forest, 1974). Near Goree, in 20 m<br />
(Balss, 1921); same, in 15-20 m (Monod, 1956). Anse Bernard,<br />
near Dakar, 13-15 m; between Goree and Rufisque,<br />
25 m; E of Bel-Air, 14 m; Ngazobil, 4 m; Joal, 4 m,<br />
10-11 m and 15 m (Monod, 1956).<br />
Guinea: 09°25'N, 13°55'W, 15 m; 09°16'N, 13°34'W,<br />
10 m; between Tamara and lie de Corail, 10-12 m (all<br />
Monod, 1956). Near Conakry, 20 m (Monod, 1956).<br />
09°51'N, 15°30'W (Capart, 1951; Forest, 1974).<br />
Sierra Leone: No specific locality, 10-22 m (Longhurst,<br />
1958). 08°38'-08°42'N, 8-12 m; Murray Town; Freetown<br />
(Monod, 1956). Off Freetown, 15 m (Forest, 1974).<br />
Ivory Coast: Grand-Bassam, 30 m, 60 m (Le Loeuff and<br />
Intes, 1968). Off San-Pedro, 20 m (Forest, 1974).<br />
Ghana: Near Accra, in 14, 35, 37, and 44 m (Monod,<br />
1956); same, in 0-45 m (Gauld, 1960). Chorkor, near Accra;<br />
Christiansborg; Ningo [?] (all Monod, 1956).<br />
Togo: No specific locality (Monod, 1956); Lome (Monod,<br />
1956).<br />
Cameroon: Douala (Balss, 1921).<br />
Principe: No specific locality (Osorio, 1889, 1898).<br />
Sao Tome: No specific locality (Balss, 1921). Praia de<br />
Santa Catarina, 00°16'N, 06°29'E, 3-10 m (Forest and<br />
Guinot, 1966). Sao Joao dos Angolares, 4-5 m (Osorio, 1889,<br />
1898).<br />
Gabon: No specific locality (Monod, 1956). Cap Lopez,<br />
20 m (Crosnier, 1967; Forest, 1974). Near Cap Lopez,<br />
OO°53'S, 08°40'E, in 33-55 m (Capart, 1951; Forest, 1974).<br />
Cabinda: No specific locality, in 25 m (Crosnier, 1967;<br />
Forest, 1974).
NUMBER 306<br />
Angola: Ambrizete (as Ambrizette), 10 m; Quicembo (as<br />
Kinsembo) (Balss, 1921). Luanda (as Sao Paulo de Loanda),<br />
a few feet deep (Rathbun, 1921).<br />
Dromia nodosa A. Milne Edwards and Bouvier,<br />
1898<br />
Dromia nodosa.—Monod, 1956:65 [part].—Forest, 1974:94,<br />
figs. 2, 3e, 5, 7e,f, pi. 2: figs. 3, 4, pi. 5: fig. 6 [references].<br />
REMARKS.—There is a male syntype of this<br />
species (USNM 22937), carapace length 7.2 mm,<br />
taken by the Talisman in the Cape Verde Islands<br />
in 75 m, 29 July 1883, in the collection of the<br />
<strong>Smithsonian</strong> <strong>Institution</strong>. Forest (1974:95) noted<br />
that only three of the original five specimens<br />
could then be located.<br />
DISTRIBUTION.—Cape Verde Islands, in 75 m,<br />
and off the Atlantic coast of Morocco.<br />
Genus Sternodromia Forest, 1974<br />
Stemodromia Forest, 1974:100 [type-species: Dromia spinirostris<br />
Miers, 1881, by original designation and monotypy; gender:<br />
feminine].<br />
* Sternodromia spinirostris (Miers, 1881)<br />
FIGURE 3C<br />
Dromia spinirostris Miers, 1881a:271, pi. 16: fig. 2.—Rathbun,<br />
1900a:300.—Odhner, 1923:15.—Capart, 1951:23 [part,<br />
not the specimens from A.S. 141 and Mercator], figs. 2,<br />
3b.—Rossignol, 1957:75.<br />
Dromia fulvo-hispida Miers, 1881a:270, pi. 16: fig. 1.<br />
Dromia fulvohispida.— Rathbun, 1900a: 300—Balss, 1921:47.<br />
Dromia atlantica Doflein, 1904:10, 189, 252, 253, pi. 7: figs. 3,<br />
4.<br />
Dromidtopsis spinirostris.—Monod, 1956:72, figs. 72-82, 83c—<br />
Longhurst, 1958:87.—Rossignol, 1962:113.—Forest and<br />
Guinot, 1966:47.—Crosnier, 1967:322.—Le Loeuff and<br />
Intes, 1968:38, 65, 67, 68, 71, fig. 62, tables 1, 6, 7, 8, 9;<br />
1969:64, 65.—Maurin, 1968a:48.—Maurin, 1968b:484,<br />
489, 491, fig. 7.—Tiirkay, 1975a: 71 [listed], 72.—Pastore,<br />
1976:114, fig. 1 [Mediterranean].—Lewinsohn, 1977:7<br />
[discussion].<br />
Dromia nodosa.—Monod, 1956:69 [part, specimen from Murray<br />
Town].—Rossignol, 1962:113. [Not Dromia nodosa A.<br />
Milne Edwards and Bouvier, 1898.]<br />
Dromia.— Voss, 1966:22.<br />
Dromia monodi.—Crosnier, 1967:321 [part, the two female<br />
specimens].<br />
Dromidiopsis.—Maurin, 1968b, figs. 4, 9.<br />
Dromidopsis spinifrons.—Maurin, 1968b:486 [erroneous spelling]-<br />
Stemodromia spinostris.—Forest, 1974:100, figs, If, 2, 3h, 5,<br />
6c,d, 7g-i, pi. 4: figs. 4,5, pi. 6: fig. 3, pi. 7: figs. 2, 4<br />
[erroneous spelling].<br />
Not Dromia spinirostris.—Osorio, 1889:136, 139; 1898:193.—<br />
Balss, 1921:47. [= Dromia monodi Forest and Guinot,<br />
1966.]<br />
Not Dromia atlantica.—Rathbun, 1921:393, fig. 1, pi. 18: fig.<br />
3 [= Dromia monodi Forest and Guinot, 1966].<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 68, 70 m, broken shell, 2 juv (L).<br />
Ivory Coast: Sta 46, 38-42 m, mud bottom with dense<br />
Jullienella, 6
20<br />
and a carapace width of 72 mm; in the largest<br />
female these values are 58 mm and 68 mm,<br />
respectively. Ovigerous (and impregnated) females<br />
have cl 48 to 58 mm and cb 55 to 58 mm.<br />
In juveniles the carapace width is slightly less (0.9<br />
times) than the carapace length, and the larger<br />
specimens become relatively wider. Even in the<br />
largest specimen however, the ratio between carapace<br />
width and carapace length does not go<br />
beyond 1.19, the females as a rule being on the<br />
average somewhat wider than the males. Our<br />
finds agree with those by Crosnier (1967) who<br />
found that this species is slightly wider on the<br />
average than Dromia monodi, although the difference<br />
is slight.<br />
REMARKS.—The present species greatly resembles<br />
Dromia monodi. Differences between adult<br />
specimens of the two species, some of which have<br />
already been pointed out by previous authors<br />
(Monod, 1956; Forest and Guinot, 1966; Crosnier,<br />
1967; Forest, 1974) are the following:<br />
1. The pubescence in Stemodromia spinirostris<br />
consists of a very dense cover of short hairs of<br />
equal length giving the animal a smooth and<br />
shiny, almost sealskin-like surface. In Dromia monodi<br />
the hairs are of two sizes, the shorter are less<br />
closely placed than in S. spinirostris and therefore<br />
do not produce the skin-like effect, while groups<br />
of longer hairs, especially in the anterior and<br />
lateral portions of the carapace give the pubescence<br />
a still more shaggy appearance.<br />
2. The interantennular spine in S. spinirostris<br />
is hardly visible in dorsal view; in Dromia monodi<br />
it is distinct and reaches beyond the middle of the<br />
rostral teeth.<br />
3. The anterolateral teeth of the carapace in<br />
D. monodi are far stronger and more conspicuous<br />
than in Stemodromia spinirostris; moreover, they are<br />
directed more outwards and are less appressed. In<br />
5. spinirostris the teeth are inconspicuous, especially<br />
the second and third, which are often hidden<br />
by the pubescence. Some of the teeth, especially<br />
the fourth and fifth are often continued<br />
backward as a marginal carina. The first anterolateral<br />
tooth is far more conspicuous than the<br />
second.<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
4. Between the first anterolateral tooth and<br />
the suprasutural tooth (Ihle, 1913:9, fig. 5) there<br />
is a marked groove in Dromia monodi; in Stemodromia<br />
spinirostris such a groove is hardly noticeable,<br />
while the teeth are less distinct.<br />
5. The chelipeds in Dromia monodi have strong<br />
tubercles: one on the outer surface of the palm at<br />
the base of the dactylus, and two near the anterior<br />
margin of the carpus. These tubercles are hardly<br />
noticeable in Stemodromia spinirostris being quite<br />
obscured by the pubescence.<br />
6. The outer surface of the carpus of the<br />
second and third pereiopods (= first and second<br />
walking legs) is evenly rounded in Stemodromia<br />
spinirostris, while in Dromia monodi it shows a distinct<br />
longitudinal ridge in the pubescence.<br />
7. The dactyli of the second and third pereiopods<br />
in D. monodi are shorter than in S. spinirostris<br />
and have stronger spines on the lower margin.<br />
8. The upper margin of the propodus of the<br />
fourth pereiopod is longer than the height of this<br />
segment in S. spinirostris, much shorter than the<br />
height in D. monodi. The dactylus is twisted in the<br />
latter species, lying in the same plane as the<br />
propodus in the former.<br />
9. The fifth pereiopod is more slender in D.<br />
monodi than in S. spinirostris.<br />
10. The female abdomen in non-ovigerous females<br />
is triangular with a rounded tip in D.<br />
monodi, more trapezoidal with a truncated tip in<br />
S. spinirostris.<br />
11. In the same non-ovigerous females the sternal<br />
grooves in S. spinirostris end somewhat closer<br />
together than in D. monodi.<br />
12. In our material of S. spinirostris the pubescence<br />
is grayish brown and the finger tips of the<br />
chelae "are white; in D. monodi the pubescence is<br />
more yellowish brown and the finger tips are<br />
pink.<br />
Young specimens of Stemodromia spinirostris differ<br />
from the adults in the following points:<br />
1. The pubescence of the carapace shows<br />
some longer hairs in the anterior part, by which<br />
the smooth appearance is lost and becomes more<br />
shaggy like in D. monodi.
NUMBER 306 21<br />
2. The interantennular spine is relatively<br />
longer and more distinct.<br />
3. The anterolateral teeth of the carapace are<br />
even less conspicuous than in the adults.<br />
These young specimens still can be distinguished<br />
from young of Dromia monodi by the character<br />
of the smaller anterolateral teeth and the<br />
more slender dactyli of the second and third<br />
pereiopods, in which the lower margin bears<br />
smaller spinules.<br />
The sternal grooves in the females collected by<br />
the Pillsbury Expedition, all of which are nonovigerous,<br />
very closely resemble those of non-ovigerous<br />
females of Dromia monodi, the only difference<br />
being that the anterior ends of the two<br />
grooves are placed somewhat closer together. In<br />
the Museum national d'Histoire naturelle, Paris,<br />
we examined five large females (cl 48 to 58 mm)<br />
of the present species, some of which are impregnated<br />
or ovigerous. In these specimens the sternal<br />
grooves converge anteriorly and end at a distinct<br />
median tubercle, as shown in Monod's (1956)<br />
figures 78 and 79. In a juvenile female in the<br />
Paris Museum (cl 34 mm) no tubercle was seen,<br />
but the grooves resemble those found in the Pillsbury<br />
specimens (Figure 3c, 9 cb 21 mm) in which<br />
the tubercle is visible. Evidently the arrangement<br />
of the sternal grooves changes drastically when<br />
sexual maturity is reached. It is debatable<br />
whether the sole character of the placement of<br />
the female sternal grooves is sufficient to distinguish<br />
genera. This seems the more doubtful when<br />
one considers the fact that Dromia monodi and<br />
Sternodromia spinirostris resemble each other so<br />
closely that they are difficult to separate, while<br />
both differ less from one another than from the<br />
other <strong>West</strong> <strong>African</strong> Dromiidae. Monod (1956:75)<br />
already commented that it does not seem logical<br />
that these two species should be placed in different<br />
genera. For the time being, however, we<br />
followed Forest (1974), who separated the present<br />
species from the other Dromidiopsis and Dromia<br />
species and erected a new genus, Sternodromia, for<br />
it. Lewinsohn (1977:7), when dealing with Dromiidae<br />
of the Red Sea, commented that in his view<br />
too much importance has been attached to the<br />
female sternal grooves as a generic character and<br />
that a revision of the family on a generic level is<br />
highly desirable.<br />
There can be little doubt that the type material<br />
of Miers' Dromia spinirostris belongs to the present<br />
species, and in the modern literature the epithet<br />
spinirostris has practically always consistently and<br />
correctly been used for it. A quite different matter<br />
is the identity of Dromia fulvo-hispida Miers, 1881.<br />
This species was based on a single juvenile specimen<br />
(8X11 mm), taken together with the types<br />
of Dromia spinirostris. The specimen was reexamined<br />
by Monod (1956:59), who gave additional<br />
details of it but did not arrive at a conclusion<br />
concerning its identity. We have also examined<br />
the specimen, which is in a poor condition; it is<br />
a juvenile female, which is partly damaged; therefore,<br />
the measurements 8X11 mm cannot be<br />
relied upon. We decided after studying this specimen<br />
that it is a juvenile specimen of Sternodromia<br />
spinirostris. The fact that the anterolateral margins<br />
show hardly any teeth (Miers described these<br />
margins as entire, but as Monod correctly pointed<br />
out there is at least one visible tooth) points<br />
strongly in this direction. Furthermore that it was<br />
taken at a depth of 9 to 15 fathoms (16.5-27 m)<br />
together with the types of Dromia spinirostris make<br />
the identity of the two highly likely. As the type<br />
specimen of Dromia fulvohispida shows no characters<br />
that make it impossible to identify it with D.<br />
spinirostris and as there are several indications that<br />
the two species are synonymous, we confidently<br />
sink the epithet Julvohispida as a synonym of spinirostris.<br />
Under Dromia monodi (p. 17) we have given the<br />
reasons why the specimens that Osoria (1889,<br />
1898) and Balss (1921) considered to be Dromia<br />
spinirostris actually should be referred to Dromia<br />
monodi. Also we point out that the female specimens<br />
from Dahomey and Cap Santa Clara, identified<br />
by Rossignol (1962) and Crosnier (1967) as<br />
Dromia monodi, be better referred to the present<br />
species. All this again clearly shows the close<br />
resemblance of the two species.<br />
The juvenile specimens from Angola that<br />
Odhner (1923) identified as Dromia spinirostris
22<br />
were referred by Monod (1956:72) to Dromia nodosa<br />
(= D. monodi). Although Monod examined<br />
one of Odhner's specimens, we are not convinced<br />
that his conclusion is correct. Odhner himself was<br />
convinced that his specimens were conspecific<br />
with Dromia atlantica Doflein, after he compared<br />
them with Doflein's description and figures;<br />
moreover Odhner sent his material to W. T.<br />
Caiman, who did not find any characters that<br />
would make them specifically different from<br />
Miers' types of Dromia spinirostris. Also the fact<br />
that Odhner's specimens came from depths of 72<br />
and 108 m makes it more likely that they belong<br />
to Sternodromia spinirostris than to Dromia monodi.<br />
As already suggested by Balss (1921), Odhner<br />
(1923), Monod (1956), and others, Dromia atlantica<br />
of Doflein (1904) is identical with the present<br />
species, as is clearly shown by Doflein's account.<br />
The juvenile male from Sta A.S. 141 and the<br />
juvenile male collected by the Mercator expedition,<br />
both of which were identified by Capart (1951)<br />
with the present species, were examined by Forest<br />
(1974:96) and found to be Dromia monodi.<br />
The specimen from Murray Town, Sierra<br />
Leone, assigned by Monod (1956:69) to Dromia<br />
nodosa, was examined by us and proved to belong<br />
to the present species.<br />
The frequent misidentification, especially of<br />
juvenile specimens, clearly shows the difficulty of<br />
distinguishing the various species of <strong>West</strong> <strong>African</strong><br />
Dromiidae.<br />
BIOLOGY.—The species has been reported from<br />
depths between 7.5-9.5 and 108 m. As already<br />
pointed out by Forest and Guinot (1966) it has<br />
its optimum occurrence in water deeper than 25<br />
m, this in contrast to Dromia monodi, which is<br />
hardly ever found in water deeper than 25 m.<br />
About 90% of the published records for Sternodromia<br />
spinirostris are from depths of 25 m or over<br />
and about 80% from depths between 25 and 75<br />
m. Le Loeuff and Intes (1968) indicate that the<br />
species "vit de 35 a 60 m et surtout a 40 et 50 m."<br />
The bottom on which the species has been taken<br />
has variously been described as "partly shelly and<br />
partly muddy" (Miers, 1881a); sand and shell<br />
fragments, sand and muddy sand with broken<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
shells and stones (Odhner, 1923); sand, mud,<br />
coral and rock, black mud, mud and brown sand,<br />
green mud and sand, brown mud, mud (Capart,<br />
1951); mud (Rossignol, 1957); muddy sand<br />
(Longhurst, 1958); mud, rock, calcareous algae,<br />
sand and Foraminifera, mud and sand, mud and<br />
shells, muddy sand and Foraminifera, mud with<br />
Area shells (Forest and Guinot, 1966); sandy mud<br />
and shells (Voss, 1966; and p. 19 herein); sandy<br />
mud with Foraminifera (Crosnier, 1967); "fonds<br />
detritiques en vases" (Maurin, 1968a, 1968b);<br />
sand or sand with shells and Suberites, liquid mud<br />
sometimes mixed with very fine sand (Maurin,<br />
1968b); mud, Foraminifera, dark red bryozoans,<br />
mud with Jullienella (p. 19). Le Loeuff and Intes<br />
(1968), who obviously carefully studied the species,<br />
stated that in Ivory Coast waters they only<br />
found it on sandy mud bottoms, and they ranged<br />
it among the "vasicoles."<br />
None of our specimens carried a sponge or<br />
ascidians, and we know of no record in the literature<br />
indicating such an association.<br />
Ovigerous females have been collected in February,<br />
April, May, and December (Capart, 1951;<br />
Forest and Guinot, 1966; Forest, 1974; and p.<br />
19).<br />
DISTRIBUTION.—Sternodromia spinirostris is a <strong>West</strong><br />
<strong>African</strong> species which occurs from the Spanish<br />
Sahara to Angola. Pastore (1976) recently recorded<br />
the species from the Gulf of Taranto in<br />
the Mediterranean. The <strong>West</strong> <strong>African</strong> records in<br />
the literature are as follows:<br />
Spanish Sahara: Islote Virginia (as Hot Virginie) (Forest,<br />
1974). Between Cabo Barbas and Cabo Blanco, 50-90 m<br />
(Maurin, 1968b). Between Cabo Corbeiro and Cabo Blanco,<br />
60-80 m (Maurin, 1968a). Cabo Blanco, 2O°37.7'N,<br />
17°24.4'W, 57 m (Tiirkay, 1975).<br />
Mauritania: 20 c 55'N, 17°22'W, 60 m; and 20°19.5'N,<br />
17°13.8'W, 20 m (Forest, 1974). Bane d'Arguin, 30-40 m<br />
(Maurin, 1968b).<br />
Cape Verde Islands: 15°16'28"N, 23°27'24"W, 40-45 m<br />
(Forest, 1974).<br />
Senegal: PSenegal, 50-62 m (Forest, 1974). Saint-Louis,<br />
35-40 m (Maurin, 1968b). Baie de Goree, 18-28 m (Miers,<br />
1881a). Between Goree and Cap Manuel, 7.5-9.5 m; Ngaparou<br />
(Forest, 1974). 13°01'N, 17°24'W, 51-55 m; and<br />
12°55.5'N, 17°33'W, 65-75 m (Forest and Guinot, 1966;<br />
Forest, 1974).
NUMBER 306<br />
Gambia: 13°12'N, 17°O3.5'W, 21 m (Forest, 1974).<br />
Guinea-Bissau: 10°19'N, 16°24'W, 60-73 m (Forest and<br />
Guinot, 1966; Forest, 1974).<br />
Guinea: 09°28'N, 14°58'W, 45 m (Forest, 1974). 09°15'N,<br />
14°50'W, 45 m (Monod, 1956).<br />
Sierra Leone: No specific locality, in 39 m (Longhurst,<br />
1958). Murray Town (Monod, 1956).<br />
Ivory Coast: No specific locality (Le Loeuff and Intes,<br />
1969). 04°35'N, 06°40'W to 04°35'N, 06°41'W, 64 m (Voss,<br />
1966). O5°O2.5'N, 05°25'W, 21-27 m (Forest and Guinot,<br />
1966; Forest, 1974). 05°11.5'N, 04°09'W, 40 m; 05°12.5'N,<br />
04°05'W, 40 m; and 05°04'N, 03°22.5'W, 50 m (Forest,<br />
1974). Fresco; Grand-Lahou, 40 m; Jacqueville; Grand-Bassam,<br />
35-60 m (overall depth range 15-60 m) (all Le Loeuff<br />
and Intes, 1968).<br />
Ghana: 05°18'N, 00°24'W, 35-37 m (Forest, 1974).<br />
Dahomey: 06°04'N, 01°38.5'E, 48 m (Forest, 1974). 06°<br />
10'N, 02°02'E, 45 m (Crosnier, 1967; Forest, 1974).<br />
Nigeria: 04°03'N, 06°12'E, 32 m (Forest and Guinot,<br />
1966; Forest, 1974).<br />
Cameroon: 03°54'N, 08°50'E, 65-70 m (Forest, 1974).<br />
Gabon: W of Cap Santa Clara, 20-40 m (Rossignol, 1962;<br />
Crosnier, 1967). Off Pointe Banda, 03°57.5'S, 10°36.5'E,<br />
85 m (Capart, 1951).<br />
Congo: Pointe-Noire, 100 m (Forest, 1974). Off Pointe-<br />
Noire, 50 m (Rossignol, 1957, 1962). Off Pointe-Noire, 04°<br />
23'S, 1 l°07'E, 88 m, and 05°S, 11°36'E, 50 m (Forest, 1974).<br />
Zaire: Off Banana, 05°54'S, 11°58.5'E, 50 m; 05°55'S,<br />
12°01'E, 25-30 m; and 05°56'S, 12°E, 50-60 m (Capart,<br />
1951; Forest, 1974).<br />
Angola: Off Moita Seca, 06°06'S, 12°14'E, 12-15 m<br />
(Capart, 1951; Forest, 1974), and 06°16'S, 12°07'E, 50 m<br />
(Capart, 1951). Off the mouth of the Congo River, 06°18'S,<br />
12°02'E, 44 m (Doflein, 1904). Near Ambriz, O7°57'S, 13°<br />
05'E, 40-50 m (Capart, 1951; Forest, 1974). Ambriz, 70 m<br />
(Forest, 1974). Porto Alexandre, 72 and 108 m (Odhner,<br />
1923).<br />
Family DYNOMENIDAE Ortmann, 1892<br />
DYNOMENIDAE Ortmann, 1892:538, 541, 543.<br />
EASTERN ATLANTIC GENERA.—One, Dynomene,<br />
represented in the tropical fauna.<br />
EASTERN ATLANTIC SPECIES.—One, Dynomene Jilholi,<br />
material of which was collected by the Pillsbury.<br />
Genus Dynomene Desmarest, 1823<br />
Dynomene Desmarest, 1823:422 [index] [a genus without included<br />
nominal species; type-species: Dynomene hispida<br />
Guerin-Meneville, 1832, by subsequent monotypy by<br />
Guerin-Meneville, 1832, in 1829-1844, pi. 14; gender:<br />
feminine].<br />
Maxillothrix Stebbing, 1921:456 [type-species: Maxillothrix actaeiformis<br />
Stebbing, 1921, a subjective junior synonym of<br />
Dynomene pilumnoides Alcock, 1899, by monotypy; gender:<br />
feminine].<br />
REMARKS.—So far as we can determine, the<br />
first latinized use of the generic name Dynomene<br />
was by Desmarest (1823:422) in the index to his<br />
article in the Dictionnaire des Sciences Naturelles. In<br />
the text of this article (182S:249, footnote) the<br />
name is used in the vernacular (DYNOMENE). Desmarest<br />
again used the vernacular form in his book<br />
Considerations Generates sur la Classe des Crustaces<br />
(1825:133, footnote), using DYNOMENE in the text<br />
(p. 133), DYNOMENE HISPIDE in the list of<br />
figures (p. 432), and Dynomene hispide on the<br />
plate itself (plate 18); in this work he gave the<br />
Latin name again in the index (p. 442). The<br />
specific epithet hispida, often attributed to Latreille<br />
or Desmarest, apparently was first used by<br />
Guerin-Meneville (14 July 1832: pi. 14). Guerin<br />
also used the Latin name in the text of his Iconographie<br />
which, however, was published as late as<br />
September 1844. We are much indebted to Col.<br />
C. F. Cowan (in litt., 29 September 1971), who<br />
provided the information that livraison 22 of<br />
Guerin's Iconographie, which was published on 14<br />
July 1832 (Cowan, 1971:29), contained Crustacea<br />
plates 12 and 14.<br />
* Dynomene filholi Bouvier, 1894<br />
Dynomene filholi.—Monod, 1956:76, figs. 84-88, 873.—Forest<br />
and Guinot, 1966:48.<br />
MATERIAL EXAMINED.—Pillsbury Material: Annobon:<br />
Sta 275, 9-69 m, rubble of coralline algae, 26, 1? (L). Sta<br />
282, 18-37 m, nodular coralline algae, 16, 59, 6 juv (L). Sta<br />
283, 51-55 m, nodular coralline algae, 25
24<br />
circular, about IV4 times broader than long, depressed,<br />
moderately convex. Fronto-orbital region<br />
forming on obtuse triangle, frontal margin thickened,<br />
fronto-orbital width about 2/3 carapace<br />
width. Regions well marked but obscured by coat<br />
of low setae. Several rounded prominences on<br />
surface, each with long tuft of setae: usually 1<br />
epigastric, 2 or 3 protogastric, 1 metagastric, 1<br />
intestinal, and 5 branchial. Anterolateral margins<br />
with 5 blunt teeth, each tufted.<br />
Chelipeds stout, subequal, larger in males, covered<br />
with low setae, ornamented with rounded,<br />
tufted prominences. Merus with subdistal prominence<br />
on posterior margin, smaller, lower naked<br />
tubercles on inner margin. Carpus with 3 tufted<br />
prominences on posterior margin, 1 dorsally, 1<br />
anteriorly; inner angle produced into slender,<br />
flattened lobe, almost spatulate. Propodus stout,<br />
high, slightly longer than movable finger, all with<br />
almost naked ventral surface ornamented with<br />
rounded prominences. Upper and outer surfaces<br />
of chela with longer setae, denser on inner face.<br />
Fingers gaping, movable finger with prominent<br />
tooth near midlength, naked apically, apices<br />
spatulate.<br />
Walking legs decreasing in length posteriorly,<br />
surface densely tomentose, dorsal surfaces with<br />
rounded, tufted prominences and some scattered<br />
longer setae. Dactyli curved, apices corneous, ventral<br />
margins toothed. Fifth legs very short, not<br />
extending much beyond midlength of merus of<br />
fourth, dactylus very small, subterminal, leg appearing<br />
chelate.<br />
Abdomen of 7 somites in both sexes, broadening<br />
posteriorly in males.<br />
Figures: A. Milne Edwards and Bouvier, 1900,<br />
pi. 3: fig. 3 (color); Monod, 1956, fig. 873.<br />
Male Pleopod: A. Milne Edwards and Bouvier,<br />
1900, pi. 8: fig. 13; Monod, 1956, figs. 84-88<br />
(both Cape Verde Islands).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 3 to 16 mm; the carapace widths<br />
of ovigerous females are 7 to 12 mm.<br />
REMARKS.—The male pleopods of our specimens<br />
have a large, triangular apical lobe, resembling<br />
that of the pleopod figured by A. Milne<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Edwards and Bouvier (1900, pi. 8: fig. 13) rather<br />
than that figured by Monod (1956, figs. 84-88).<br />
BIOLOGY.—The depth range of the species is<br />
from 23 to 1477 m (with one record of 9-47 m),<br />
but 80% of the records are from between 23 and<br />
75 m. As the Talisman and Travailleur records are<br />
notoriously unreliable, it is possible that the records<br />
from 110-180 and 150-275 m are incorrect.<br />
Also the depth of 1477 m (Monaco Expedition<br />
Sta 1209) needs confirmation.<br />
The Pillsbury specimens all were taken from a<br />
bottom of nodular coralline algae (diameter of<br />
the nodules about 100-150 mm; see Voss, (1966:<br />
50, fig. 14)). In the literature the species is reported<br />
from "fond dur" (Bouvier, 1922:90),<br />
"banes de corail rouge" (A. Milne Edwards and<br />
Bouvier, 1900:9) and from bottoms with calcareous<br />
algae or sand, rock, corals and calcareous<br />
algae (Forest and Guinot, 1966). Off <strong>West</strong> Africa,<br />
ovigerous females have been collected in May<br />
{Pillsbury).<br />
DISTRIBUTION.—This is an insular <strong>West</strong> <strong>African</strong><br />
species, known so far from the Cape Verde islands<br />
(the type-locality) and the Gulf of Guinea Islands,<br />
Principe, and Annobon; it has not been recorded<br />
from the mainland. Monod (1956) reported material<br />
from the Cape Verde Islands and Principe.<br />
Since 1956 Dynomene filholi has been recorded from<br />
the following localities:<br />
Principe: 01°35'N, O7°28'E, 45 m; and 01°38'35"N,<br />
07°21'35"E, 35 m (Forest and Guinot, 1966).<br />
Annobon: O1°27.5'S, 05°36.5'E, 35 m; N of San Antonio,<br />
23 m; 01°26'15"S, 05°35'40"E, 60 m (Forest and Guinot,<br />
1966).<br />
Family LATREILLIIDAE Stimpson, 1858<br />
LATREILLIDEA Stimpson, 1858c:226 [corrected to Latreilliidae<br />
by Stebbing, 1902:23; name 373 on Official List].<br />
EASTERN ATLANTIC GENERA.—One, Latreillia,<br />
represented in the tropical fauna.<br />
EASTERN ATLANTIC SPECIES.—One, Latreillia elegans,<br />
not represented in the Pillsbury collections.<br />
Genus Latreillia Roux, 1830<br />
Latreillia P. Roux, 1830, pi. 22 [type-species: Latreillia elegans
NUMBER 306<br />
Roux, 1830, by monotypy; gender: feminine; name 1630<br />
on Official List].<br />
Praetor Gistel, 1848:ix [erroneously substituted for Latreillia<br />
(see Rathbun, 1937:73); type-species: Latreillia elegans<br />
Roux, 1830; gender: masculine].<br />
Latreillia elegans Roux, 1830<br />
Latreillia elegans.—Monod, 1956:78 [references].—Zariquiey<br />
Alvarez, 1968:307 [Spain; references].<br />
Latreilla elegans.—Tiirkay, 1976a:25 [listed], 36, fig. 16 [Morocco;<br />
erroneous spelling].<br />
DISTRIBUTION.—Mediterranean Sea and adjacent<br />
eastern Atlantic, from Portugal, the Azores,<br />
off the Sahara coast, and the Cape Verde Islands,<br />
in depths between 100 and 475 m. Ascension<br />
Island (Stebbing, 1914); western Atlantic (Rathbun,<br />
1937).<br />
Family HOMOLIDAE de Haan, 1839<br />
HOMOLIDEA de Haan, 1839:102 [corrected to Homolidae by<br />
White, 1847a:55; name 243 on Official List, there attributed<br />
to White, 1847, in error].<br />
THELXIOPEIDAE Rathbun, 1937:62 [name 278 on Official<br />
Index].<br />
EASTERN ATLANTIC GENERA.—Three, two of<br />
which, Homola and Paromola, are represented by<br />
species occurring off tropical <strong>West</strong> Africa. The<br />
other genus is Homologenus A. Milne Edwards (in<br />
Henderson, 1888:20): A substitute name for Homolopsis<br />
A. Milne Edwards (1880:34), an invalid<br />
junior homonym of Homolopsis Bell, 1862 (typespecies:<br />
Homolopsis rostratus A. Milne Edwards,<br />
1880, by monotypy; gender: masculine).<br />
EASTERN ATLANTIC SPECIES.—Three, two of<br />
which were recorded by Monod (1956), as follows:<br />
Name in Monod Current Name<br />
Thelxiope barbata Homola barbata*<br />
Paromola cuvieri Paromola cuvieri<br />
The extralimital species is Homologenus rostratus<br />
(A. Milne Edwards, 1880): Eastern Atlantic from<br />
the Azores, Madeira, and off Morocco, 1435 to<br />
2195 m (Tiirkay, 1976a); western Atlantic (Rathbun,<br />
1937).<br />
Genus Homola Leach, 1815<br />
Thelxiope Rafinesque, 1814:21 [suppressed by the International<br />
Commission on Zoological Nomenclature, Opinion<br />
522 in Bulletin of Zoological Nomenclature, 19:211; type-species:<br />
Thelxiope palpigera Rafinesque, 1814, by monotypy;<br />
gender: feminine; name 1190 on Official Index].<br />
Homola Leach, 1815a:324 [type-species: Homola spinifrons<br />
Leach 1815, a subjective junior synonym of Cancer barbatus<br />
Fabricius, 1793, by monotypy; gender: feminine; name<br />
1301 on Official List].<br />
Homolax Alcock, 1899b: 124, 129, 156 [type-species: Homola<br />
megalops Alcock, 1894, by monotypy; gender: feminine].<br />
Moloha Barnard, 1947:371 [type-species: Latreillopsis alcocki<br />
Stebbing, 1920, by monotypy; gender: feminine].<br />
* Homola barbata (Fabricius, 1793)<br />
Thelxiope barbata.—Gordon, 1950:221, 230, 232, 239, 250,<br />
figs. 19, 26b-d.—Monod, 1956:79.—Maurin, 1968b:<br />
486.—Le Loeuff and Intes, 1968:31, table 1.<br />
Homola barbata.—Figueira, 1960:7.—Guinot and Ribeiro,<br />
1962:23.—Peres, 1964:20.—Forest and Guinot, 1966:<br />
48—Crosnier, 1967:322.—Maurin, 1968a: 19, 30, 43, 61,<br />
107 [p. 107 Balearic Isles]; 1968b:480, 484.—Zariquiey<br />
Alvarez, 1968:304, figs. 12g, 106c [Spain; references].—<br />
Rice and Provenzano, 1970:446 ff., figs. 1-15 [larvae].—<br />
Tiirkay, 1976a:25 [listed], 36 [Portugal, in part]; 1976b:<br />
61 [listed], 62.<br />
SYNONYMS.—Cancer cubicus Forskal, 1775 (suppressed<br />
by ICZN); Cancer novemdecos Sulzer, 1776<br />
(suppressed by ICZN); Thelxiope palpigera Rafinesque,<br />
1814; Homola spinifrons Leach, 1815; Dorippe<br />
spinosus Risso, 1816.<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 68, 70 m, broken shell, 19 ov (L).<br />
Nigeria: Sta 241, 59-63 m, mud and shell, 19 (W).<br />
Annobon: Sta 283, 51-55 m, nodular coralline algae, 1(5<br />
(W).<br />
Other Material: Congo: Off Pointe-Noire, 04°56'S, 11°<br />
31'E, 95-97 m, trawl, 21-22 Sep 1965, 2
26<br />
Lateral margin of carapace with line of spinules<br />
decreasing in size posteriorly. Linea anomurica<br />
indistinct. Front, under orbit, with 3 spines forming<br />
triangle. Pterygostomian region with curved<br />
row of 6 spines dorsally, lower margin with curved<br />
row of 5-6 spines, and between these, 1-2 spines;<br />
below and posterior to these larger spines are<br />
numerous small spines. Raised anterior margin of<br />
epistome terminating before meeting in midline,<br />
leaving smooth median gap. Eyes elongate, stalk<br />
greatly enlarged proximal to cornea. Basal segment<br />
of antennal peduncle with outer spine.<br />
Chelae slender, equal. Merus with 2 lines of<br />
spines below, 1 above, lower, outer spines largest.<br />
Carpus with 1 large and 1 small inner spines,<br />
outer surface with 6-8 spines arranged in 2 irregular<br />
rows and 2 distal spines. Palm with low,<br />
blunt spine on outer margin. Fingers shorter than<br />
palm, cutting edges corneous, tips of fingers crossing<br />
when closed, movable finger with low, obtuse<br />
prominence proximally on cutting edge.<br />
Walking legs compressed, thin, elongate. Meri<br />
with row of fixed spines above. Propodi elongate,<br />
each shorter than respective merus, with row of<br />
4 slender, movable spines below and 2 distal<br />
spines. Dactyli elongate, each equal to or slightly<br />
longer than respective carpus, more than half as<br />
long as respective propodus, with 11-13 spines<br />
below. Fifth leg with dorsal spine on coxa, 1 or 2<br />
spines on ischium; merus with 4 large posterior<br />
spines and 1 terminal anterior spine; carpus with<br />
numerous small spines anteriorly; propodus with<br />
numerous spines on opposable margin; dactylus<br />
with 4 spines on opposable margin.<br />
Abdomen of 7 somites in both sexes. Second<br />
segment of abdomen with prominent conical<br />
tooth.<br />
Figures: Rathbun, 1937, fig. 16, pi. 15: figs. 1,<br />
2; Rice and Provenzano, 1970, fig. 2.<br />
Male Pleopod: Gordon, 1950, figs. 26b-d (Madeira).<br />
MEASUREMENTS.—Our specimens have carapace<br />
lengths of 16 to 28 mm; the carapace length<br />
of the ovigerous female is 22 mm.<br />
REMARKS.—In addition to the specimens collected<br />
by Pillsbury, we have been able to examine<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
two larger males (cl 29 and 30 mm) reported off<br />
Pointe-Noire in 95-97 meters by Crosnier (1967:<br />
322). The male pleopods of these specimens and<br />
that from a male (cl 32 mm, Leiden Crust.<br />
D.I2989) from Naples, the type-locality, closely<br />
resemble that illustrated from a small male, cl 14<br />
mm, from Madeira by Gordon (1950). The Mediterranean<br />
and eastern Atlantic populations of<br />
this species apparently are conspecific.<br />
On the basis of their study of larvae identified<br />
with this species, Rice and Provenzano (1970)<br />
reinforced earlier suggestions by Rice (1964) and<br />
Rice and Von Levetzow (1967) that there might<br />
be several taxa, each with a distinctive larval<br />
form, currently placed in H. barbata. The recognized<br />
three groups of larvae are the western Atlantic,<br />
eastern Atlantic (including the Mediterranean<br />
and some specimens from South Africa),<br />
and a group found only off South Africa. Rice<br />
and Provenzano examined postlarval specimens<br />
from the western North Atlantic, Brazil, the Mediterranean,<br />
and South Africa but could find no<br />
consistent differences between specimens from<br />
these areas. However, they suggested that eastern<br />
Atlantic representatives of H. barbata matured<br />
faster and grew larger than specimens from other<br />
areas.<br />
BIOLOGY.—Homola barbata inhabits moderate<br />
depths, having been taken in 55-679 m in the<br />
western Atlantic (Rathbun, 1937) and between<br />
50 and 400 m in the eastern Atlantic (Monod,<br />
1956); records since 1956 indicate that the species<br />
lives in depths between 10-30 and 470-500 m in<br />
the eastern Atlantic. The Pillsbury specimens were<br />
taken on broken shell in 70 m, on mud and shell<br />
in 59-63 m, and in nodular coralline algae in 51-<br />
55 m. This species also has been taken on mud in<br />
215 m (Guinot and Ribeiro, 1962); reddish gravel<br />
with shell debris in 210 m (Peres, 1964); calcareous<br />
algae in 73 m (Forest and Guinot, 1966);<br />
muddy sand in 95-98 m (Crosnier, 1967); muddy<br />
sand or sandy detritus in 10-30 m, slightly sandy<br />
mud with funiculines in 200-250 m, alcyonarians<br />
in mud in 90-150 m, and muddy sand in 470-<br />
500 m (all Maurin, 1968a); and on muddy shell<br />
in 50 m, mud in 40-60 m, muddy detritus in 200
NUMBER 306<br />
m, and mud or sandy mud in 90-100 m (all<br />
Maurin, 1968b).<br />
Off <strong>West</strong> Africa, ovigerous females have been<br />
collected in June and September (Crosnier, 1967;<br />
Pillsbury).<br />
DISTRIBUTION.—Mediterranean, eastern and<br />
western Atlantic, and off South Africa, in depths<br />
between 10-30 m and 679 m. In the western<br />
Atlantic it has been recorded from localities between<br />
Massachusetts and Brazil (Rathbun, 1937;<br />
Rice and Provenzano, 1970). In the eastern Atlantic<br />
it is known from scattered localities in the<br />
Mediterranean and in the Atlantic from Portugal<br />
and Spain southward to South Africa, including<br />
the Azores, the Cape Verde Islands, and Madeira<br />
(Barnard, 1950; Gordon, 1950; Monod, 1956;<br />
Zariquiey Alvarez, 1968). Since 1956 it has been<br />
recorded from the following:<br />
Spain: Off Guadiana, 470-500 m (Maurin, 1968a).<br />
Azores: Off Horta, Ilha do Faial, in deep water (Figueira,<br />
1960).<br />
Madeira: Funchal, 100-ca. 250 m; Ilhas Desertas (Tiirkay,<br />
1976b).<br />
Morocco: Bane de Spartel, 35°54'N, 06°14'W, 210 m<br />
(Peres, 1964). Between Cap Rhir and Cap Draa, 10-30 m<br />
(Maurin, 1968a). Off Cap Juby, 50 m (Maurin, 1968b).<br />
31°01'N, 10°16'W, 360-375 m (Tiirkay, 1976a).<br />
Spanish Sahara: Off Medano de Aaiun and Cabo Bojador,<br />
200-300 m; Morro Garnet, 300-350 m (Maurin, 1968a);<br />
between Cabo Corbeiro and Cabo Blanco, 200 m (Maurin,<br />
1968b).<br />
Mauritania: S Bane d'Arguin, 90-150 m (Maurin,<br />
1968a). Bane d'Arguin, 40-60 and 90-100 m (Maurin,<br />
1968b).<br />
Ivory Coast: 05°06'N, 03°49'W, 50 m (Crosnier, 1967).<br />
In 40-50 m (Le Loeuff and Intes, 1968).<br />
Principe: 01°43'10"N, O7°28'2O"E, 73 m (Forest and<br />
Guinot, 1966).<br />
Congo: Off Pointe-Noire, 05°00'S, 11°32'E, 98 m, and<br />
04°56'S, 11°31'E, 95-97 m (Crosnier, 1967).<br />
Angola: Baia dos Tigros, 215 m (Guinot and Ribeiro,<br />
1962).<br />
It has not been recorded previously from Liberia,<br />
Nigeria, or Annobon, although these localities<br />
are within the known range of the species.<br />
Genus Paromola Wood-Mason and Alcock,<br />
1891<br />
Paromola Wood-Mason and Alcock, 1891:267 [type-species:<br />
Dorippe cuvieri Risso, 1816, by monotypy; gender: feminine;<br />
name 1641 on Official List].<br />
Paromola cuvieri (Risso, 1816)<br />
Paromola cuvieri.—Capart, 1951:25, fig. 4.—Monod, 1956:79,<br />
fig. 89.—Guinot and Ribeiro, 1962:23.—Rossignol, 1962:<br />
113.—Forest, 1963:628.—Peres, 1964:27, 29.—Figueira,<br />
1964:69, pis. 1,2.—Zariquiey Alvarez, 1968:301, fig. 106b<br />
[Spain; references].—Maurin, 1968a:29, 33, 50, 64, 106,<br />
114, 121 [p. 106, 114, 121 Mediterranean], fig. 22; 1968b:<br />
479, 480, 482, 484, 489, 491, 492, figs. 3, 6.—Christiansen,<br />
1969:24, fig. 8, map 2 [Scandinavia].—Crosnier, 1969:<br />
529.—Tiirkay, 1976a:25 [listed], 36; I976b:61 [listed],<br />
62.—Intes and Le Loeuff, 1976:103.<br />
Paromola.—Maurin, 1968a, fig. 23; 1968b, figs. 1, 9.<br />
Parhomola.—Maurin, 1968a, fig. 29 [erroneous spelling].<br />
Paramola.—Maurin, 1968b, fig. 4 [erroneous spelling].<br />
SYNONYM.—Maia dumerili Risso, 1816.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Morocco: Off Cap de Mazagan, 33°40'N,<br />
08°45'W, 570 m, Agassiz trawl, 28 Mar 1976, Onversaagd Sta<br />
159, 19 (L).<br />
DESCRIPTION.—Capart, 1951:27.<br />
Figures: Capart, 1951, fig. 4; Monod, 1956, fig.<br />
89; Christiansen, 1969, fig. 8.<br />
Male Pleopod: Sankarankutty, 1968, fig. la,b<br />
(North Sea).<br />
Color: "Couleur generate jaune orange, plus<br />
rouge a l'avant de la carapace et sur les epines<br />
anterieures; les pattes un peu plus foncees" (Capart,<br />
1951:27).<br />
BIOLOGY.—Paromola cuvieri is a deep water species,<br />
generally occurring on soft bottoms in depths<br />
between 150 and about 1000 m; there is one<br />
record from a depth of 952-1038 m (Tiirkay,<br />
1976a), and another from a depth of 10 m (Doflein,<br />
1904), but generally the species occurs beyond<br />
shelf depths. It has been recorded from mud<br />
or sandy mud.<br />
Off <strong>West</strong> Africa, ovigerous females have been<br />
recorded in March and October (Capart, 1951;<br />
Monod, 1956).<br />
DISTRIBUTION.—Eastern Atlantic, from the Hebrides<br />
and southern Scandinavia southward to<br />
Angola, including the Azores, the Cape Verde<br />
Islands, and the Mediterranean, in depths between<br />
10 and more than 1000 m, usually deeper<br />
27
28<br />
than 150 m. Monod (1956) summarized earlier<br />
<strong>West</strong> <strong>African</strong> records and reported material from<br />
Senegal. Since 1956 the species has been recorded<br />
from:<br />
Azores: Ilha Terceira (Figueira, 1964).<br />
Madeira: No specific locality; fish market, Madeira (Tiirkay,<br />
1976b).<br />
Morocco: Foum Agouitir (as Puerto-Cansado), 350-450<br />
m (Maurin, 1968a). 35°19'N, O6°32'W to 35°28.8'N,<br />
06°39.2'W, 333-360 m, and 35°17.5'N, 06°10.3'W to<br />
35°13.9'N, 06°36.2'W, 295-340 m (PeVes, 1964). 33°37.5'N,<br />
09°02.2'W, 952-1038 m (Tiirkay, 1976a). Asilah (as Arzila)<br />
to Larache, 300-350 m (Maurin, 1968a).<br />
Spanish Sahara: Off Medano de Aaiiin and W of Cabo<br />
Bojador, 300-500 m; off Cabo Bojador and Morro Garnet,<br />
530-720 m (Maurin, 1968b). Off Villa Cisneros, 300-500 m<br />
(Maurin, 1968a,b).<br />
Mauritania: Off Nouakchott, 350-400 to 600 m (Maurin,<br />
1968a). Bane d'Arguin, 200-300 m, and Tamzak (as<br />
Tamxat), 350-600 m (Maurin, 1968b).<br />
Senegal: Fosse de Kayar, 300-350 to 600 m (Maurin,<br />
1968b).<br />
Ivory Coast: No specific locality (Intes and LeLoeuff,<br />
1976). 04°32'30"N, 06°31'W, 300-455 m, and 04°54'N,<br />
03°23'W, 380-400 m (Forest, 1963).<br />
Ghana: 04°39'N, 02°46'W, 300-400 m (Forest, 1963).<br />
Congo: W of Banga (as Banda), 250 m (Rossignol, 1962).<br />
Off Pointe-Noire, 04°54'S, 11°19'E, 300 m (Crosnier, 1969).<br />
Angola: Baia dos Tigres, 453-478 m (Guinot and Ribeiro,<br />
1962). 06°24'N, 11°34'E, 325 m (Crosnier, 1969).<br />
Family CYCLODORIPPIDAE Ortmann, 1892<br />
CYCLODORIPPIDAE Ortmann, 1892:552.<br />
TYMOLINAE Alcock, 1896:273, 274.<br />
REMARKS.—This family is not represented in<br />
the eastern Atlantic fauna. It was considered by<br />
Glaessner (1969:R492, as Tymolinae) to be a<br />
subfamily of the Dorippidae.<br />
Family CYMONOMIDAE Bouvier, 1898<br />
CYMONOMAE Bouvier, 1898:55, 59 [corrected to Cymonomidae<br />
by Glaessner, 1969:R627].<br />
EASTERN ATLANTIC GENERA.—One, occurring<br />
to the north of the tropical region, Cymonomus A.<br />
Milne Edwards (1880:26). Type-species: Cymonomus<br />
quadratus A. Milne Edwards, 1880, by mono-<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
typy; gender: masculine; name 1618 on Official<br />
List.<br />
EASTERN ATLANTIC SPECIES.—Two, both occurring<br />
north of the tropical region:<br />
Cymonomus granulatus (Thomson, 1873). Ireland<br />
southward to Sahara coast, Mediterranean; sublittoral,<br />
300-1350 m (Zariquiey Alvarez, 1968).<br />
Cymonomus normani Lankester, 1903. Between<br />
Iceland and the Faroes, 875-1269 m, and off<br />
Portugal, 1370-1430 m (see Turkay, 1976a:36,<br />
for references).<br />
Family DORIPPIDAE MacLeay, 1838<br />
DORIPPINA MacLeay, 1838:69 [corrected to Dorippidae by<br />
White, 1847a:53; name 355 on Official List, there attributed<br />
to de Haan, 1841, in error.]<br />
ETHUSINAE Guinot, 1977:1052.<br />
EASTERN ATLANTIC GENERA.—Four, Ethusa,<br />
Ethusina, Medorippe, new genus, and Phyllodorippe,<br />
new genus, each represented by tropical species.<br />
EASTERN ATLANTIC SPECIES.—Ten, six of which<br />
were recorded by Monod (1956), as follows:<br />
Name in Monod<br />
[Cymonomus granulatus<br />
(Dorippidae)]<br />
Ethusina abyssicola<br />
Ethusa mascarone<br />
Ethusa rosacea<br />
Ethusa rugulosa<br />
Dorippe lanata<br />
Dorippe armata<br />
Current Name<br />
[Cymonomus granulatus<br />
(Cymonomidae)]<br />
Ethusina alba<br />
Ethusa vossi, new species*<br />
Ethusa rosacea*<br />
Ethusa rugulosa*<br />
Medorippe lanata*<br />
Phyllodorippe armata*<br />
A seventh <strong>West</strong> <strong>African</strong> species, Ethusina beninia,<br />
new species, is represented in the Pillsbury collections.<br />
One species assigned to the Dorippidae by<br />
Monod (1956), Cymonomus granulatus (Thomson,<br />
1873), is included under the Cymonomidae<br />
(above), following Garth and Haig (1971:6.7).<br />
The extralimital species are as follows:<br />
Ethusa mascarone (Herbst, 1785). Mediterranean<br />
and adjacent Atlantic, including Canary Islands<br />
(Miers, 1881a; A. Milne Edwards and Bouvier,<br />
1900), Portugal (Nobre, 1936), and Bay of Biscay<br />
(Zariquiey Alvarez, 1968). For possible references
NUMBER 306<br />
to E. mascarone from the NW <strong>African</strong> coast, see 44 off Cap Rhir, Morocco, 29°52'N, 14°07'W of<br />
account of E. vossi, new species (p. 39).<br />
Paris (11°47'W of Greenwich), 2083 m, 25 June<br />
Ethusa microphthalma Smith, 1881. Azores, in 1883.<br />
1000 fm (1830 m) (Miers, 1886:329); western REMARKS.—Usually the genera Dorippe sensu<br />
Atlantic (Rathbun, 1937).<br />
lato, Ethusa, and Ethusina are placed in the nom-<br />
Ethusina talismani A. Milne Edwards and Bouinate subfamily of the Dorippidae. Recently<br />
vier, 1897. Off the Azores and Cap Rhir, Mo- Guinot (1977:1052) recognized a second subfamrocco,<br />
in 2075 to 2235 m (Monod, 1956). A male ily, Ethusinae (p. 38), for Ethusa and Ethusina, an<br />
syntype (USNM 22940) in the collections of the<br />
action with which we concur.<br />
<strong>Smithsonian</strong> <strong>Institution</strong> was taken at Talisman sta<br />
Key to Subfamilies and Genera of Dorippidae<br />
1. Afferent branchial orifices narrow and placed somewhat before bases of<br />
chelipeds, being separated from chelipeds by narrow process of carapace<br />
reaching down to sternum. Last two pereiopods distinctly subchelate,<br />
dactylus closing against tubercle on propodus (Dorippinae) 2<br />
Afferent branchial orifices wide and placed immediately against bases of<br />
chelipeds. Last two pereiopods not subchelate; dactylus short and not<br />
closing against tubercle on propodus (Ethusinae) 8<br />
2. Carapace with distinct epibranchial tooth 3<br />
Carapace without epibranchial tooth 5<br />
3. Male gonopod short, broad, and straight. Gastric region of carapace with<br />
V-shaped ridge. Dactylus of second and third pereiopod with upper<br />
margin naked or with few hairs in extreme basal part (less than VA of<br />
length) 4<br />
Male gonopod slender, S-shapedly curved. Gastric region without V-shaped<br />
ridge. Dactylus of second and third pereiopod with fringe of short hairs<br />
over more than basal third of dorsal margin [Carapace broader than<br />
long.] Phyllodorippe, new genus<br />
4. Meri of second and third pereiopods with dorsal row of spines or spinules;<br />
dactyli of these legs with few hairs and spinules in extreme basal part.<br />
Carapace broader than long with some low elevations. Abdomen of male<br />
and female without teeth Medorippe, new genus<br />
Meri of second and third pereiopods without dorsal spines or spinules;<br />
dactyli of these legs naked and smooth. Carapace longer than wide, with<br />
distinct tubercles. Abdomen of male and female with distinct teeth on<br />
third and fourth somites (also on second in male) Dorippe<br />
5. Male gonopod short, stubby, with one or more narrow appendages, one or<br />
more of which are T-, hammer-, or cross-shaped. Carapace distinctly<br />
wider than long. Front reaching about as far forward as outer orbital<br />
teeth 6<br />
Male gonopod very slender and curved, ending in 3 or 4 rounded or<br />
subacute lobes. Carapace longer than wide or almost as long as wide.<br />
Front reaching beyond outer orbital teeth 7<br />
29
30<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
6. Male gonopod straight, with single twisted L- or T-shaped appendage at<br />
apex, and lobe at base Dorippoides<br />
Male gonopod with sharp angular bend in middle, with several appendages<br />
at top, without lobe at base Paradorippe<br />
7. Carapace distinctly longer than wide, flattened dorsally, without tubercles,<br />
but with grooves. Front reaching far beyond outer orbital teeth. Dactyli<br />
of second and third pereiopods with fringe of long hairs along both<br />
upper and lower margins Neodorippe<br />
Carapace slightly wider than long, convex, smooth or with tubercles. Front<br />
reaching only slightly beyond outer orbital teeth. Dactyli of second and<br />
third pereiopods with dorsal and ventral fringes of short hairs. Nobilum<br />
8. Eyes movable. Basal segment of antennules normal Ethusa<br />
Eyes immovable. Basal segment of antennules very large and swollen<br />
Ethusina<br />
Subfamily DORIPPINAE MacLeay, 1838<br />
REMARKS.—The two eastern Atlantic species<br />
belonging to this subfamily are present in the<br />
Pillsbury collections. Until recently both would<br />
have been placed in the genus Dorippe, but in<br />
1969, Serene and Romimohtarto in a paper dealing<br />
with Indo-<strong>West</strong> Pacific species, recognized<br />
instead of the old genus Dorippe three genera<br />
{Dorippe, Neodorippe, and Paradorippe) and two new<br />
subgenera, Dorippoides (in Dorippe) and Nobilum (in<br />
Neodorippe).<br />
We tried to accommodate the Atlantic species<br />
of Dorippe in the genera and subgenera recognized<br />
by Serene and Romimohtarto, but encountered<br />
several difficulties. Therefore a study first had to<br />
be made of the Indo-<strong>West</strong> Pacific species of Dorippinae.<br />
During this study we could only confirm<br />
the correctness of the taxonomic views of Serene<br />
and Romimohtarto, and even had to raise some<br />
of their subgenera to the rank of full genera.<br />
Furthermore, we found it necessary to establish<br />
two new genera, one each for the Atlantic species.<br />
It is our intention to discuss the Indo-<strong>West</strong> Pacific<br />
species of Dorippe, sensu lato, in a future paper,<br />
but our main conclusions can be found in the<br />
following list of the genera of Dorippinae that we<br />
at present recognize:<br />
Dorippe Weber (1795:93). Type-species: Cancer<br />
quadridens Fabricius, 1793, a subjective junior synonym<br />
of Cancer frascone Herbst, 1785, by subse-<br />
quent selection by Latreille, 1810:96, 422; gender:<br />
feminine. So far Dorippe contains only the typespecies.<br />
Dorippoides Serene and Romimohtarto (1969:3,<br />
4, 8). Type-species: Cancer facchino Herbst, 1785,<br />
by original designation and monotypy; gender:<br />
masculine. So far only containing the type-species.<br />
Medorippe, new genus. Type-species: Cancer lanatus<br />
Linnaeus, 1767, by present designation and<br />
monotypy; gender: feminine.<br />
Neodorippe Serene and Romimohtarto (1969:3,<br />
4, 11). Type-species: Dorippe astuta Fabricius,<br />
1798, a subjective junior synonym of Cancer facchino<br />
Herbst, 1785, by original designation; gender:<br />
feminine. An examination of three syntypes<br />
of Dorippe astuta Fabricius, 1798, in the collection<br />
of the Leiden Museum, and photographs of two<br />
of the four syntypes in the Zoological Museum,<br />
Copenhagen, showed that Dorippe astuta Fabricius,<br />
1798, is a subjective junior synonym of Cancer<br />
facchino Herbst, 1785, as defined by the lectotype<br />
selection for the latter species by Serene and<br />
Romimohtarto (1969:9). The specimens usually<br />
assigned by authors to Dorippe astuta and also<br />
described and figured under that name by Serene<br />
and Romimohtarto (1969:11, figs. 3, 7, 12, 17, pi.<br />
Id, pi. 4a,b), prove to be Dorippe callida Fabricius,<br />
1798. Photographs of the two type-specimens of<br />
Dorippe callida in the collection of the Copenhagen<br />
Museum, which could be examined by us, clearly
NUMBER 306<br />
showed the identity of the species. Thus, Neodorippe<br />
is a genus based on a misidentified typespecies,<br />
and in order to comply with Article 70(a)<br />
of the International Code of Zoological Nomenclature,<br />
we will submit this case to the International<br />
Commission on Zoological Nomenclature, requesting<br />
that under their plenary powers Dorippe<br />
callida Fabricius, 1798, be designated the typespecies<br />
of the genus Neodorippe.<br />
Although Serene and Romimohtarto assigned<br />
two species and a subspecies (the type-species,<br />
plus Dorippe japonica Von Siebold and D. japonica<br />
taiwanensis Serene and Romimohtarto) to the<br />
nominate subgenus of Neodorippe, we prefer to<br />
consider Neodorippe a genus with a single species,<br />
Neodorippe callida (Fabricius, 1798).<br />
Nobilum Serene and Romimohtarto (1969:3, 5,<br />
14). Type-species: Dorippe histrio Nobili, 1903, by<br />
original designation and monotypy; gender: neuter.<br />
We prefer to place Dorippe japonica in this<br />
genus rather than in Neodorippe. The gonopods of<br />
the two species, Dorippe japonica and D. histrio, in<br />
our opinion do not differ in essential details, while<br />
the general morphology of the body shows the<br />
species to be much closer to one another than to<br />
Neodorippe callida. Therefore we consider the genus<br />
Nobilum at present to consist of Nobilum histrio<br />
(Nobili, 1903), N. japonicum japonicum (Von Siebold,<br />
1824) and N. japonicum taiwanense (Serene<br />
and Romimohtarto, 1969).<br />
Paradorippe Serene and Romimohtarto (1969:3,<br />
5, 15). Type-species: Dorippe granulata de Haan,<br />
1841, by original designation; gender: feminine.<br />
This genus contains three species: Paradorippe granulata<br />
(de Haan, 1841), P. australiensis (Miers,<br />
1884), and P. polita (Alcock and Anderson, 1894).<br />
Phyllodorippe, new genus. Type-species: Dorippe<br />
armata Miers, 1881, by present designation and<br />
monotypy; gender: feminine.<br />
The new genera, Medorippe and Phyllodorippe,<br />
are restricted to the eastern Atlantic. The other<br />
five genera inhabit the Indo-<strong>West</strong> Pacific region.<br />
The subfamily, so far, has not been found in<br />
American waters (neither in the western Atlantic<br />
nor in the eastern Pacific).<br />
Genus Medorippe, new genus<br />
TYPE-SPECIES.—Cancer lanatus Linnaeus, 1767.<br />
ETYMOLOGY.—The name is formed from the<br />
Greek prefix me- (not) and the feminine generic<br />
name Dorippe.<br />
DIAGNOSIS.—Carapace broader than long, pubescent,<br />
with distinct epibranchial spine on either<br />
side. Extra-orbital teeth slightly surpassing front.<br />
Tubercles on dorsal surface of carapace few and<br />
low, grooves conspicuous; V-shaped ridge present<br />
on cardiac region. Eyes not reaching beyond<br />
extra-orbital teeth. Lower margin of orbit, between<br />
extra- and infra-orbital teeth, smooth. Second<br />
and third pereiopods with dorsal margin of<br />
merus carrying row of spines. Dactylus of these<br />
pereiopods narrow and somewhat twisted, not<br />
fringed with hairs (at most with very short row of<br />
hairs in extreme basal part of dorsal margin).<br />
Abdomen of male with blunt and low elevations,<br />
but without teeth or spines. First pleopod of male<br />
short, stubby and straight, with lobe at outer<br />
margin of base; apex acute, turned inward<br />
abruptly (almost at right angle), without distal<br />
appendages.<br />
REMARKS.—In the genus Medorippe, the shape<br />
of the male gonopod is closest to those of Dorippe<br />
and Dorippoides, but differs from that in both<br />
genera in the sharp straight apex which is turned<br />
inward. From the other genera, Medorippe can be<br />
distinguished by the short, straight gonopod without<br />
distal appendages. Like Dorippe and Phyllodorippe,<br />
Medorippe has a distinct epibranchial spine,<br />
which in Dorippe, however, is preceded by a row<br />
of granules. The presence of spines on the dorsal<br />
margin of the meri of pereiopods 2 and 3 distinguishes<br />
Medorippe from the others.<br />
* Medorippe lanata (Linnaeus, 1767),<br />
new combination<br />
FIGURE Aa-h<br />
Dorippe lanata.—Capart, 1951:30, fig. 6.—Monod, 1956:90,<br />
fig. 103 [as Dorippe armata; not fig. 102 = Phyllodorippe<br />
armata].—Rossignol, 1957:75, pi. 3: fig. 3.—Forest and<br />
Gantes, 1960: 350—Gauld, 1960:68.—Guinot and Ri-<br />
31
32<br />
beiro, 1962:25.—Rossignol, 1962:114.—Crosnier, 1964:<br />
34.—Forest and Guinot, 1966:50.—Guinot, 1967a:244<br />
[listed; Indian Ocean].—Zariquiey Alvarez, 1968:312,<br />
figs. 2f, 14b, 105a,b, 106b [Spain; references].—Maurin,<br />
1968a:30,41; 1968b:480, 486.—Le Loeuffand Intes, 1968:<br />
38, table 1, figs. 47, 62; 1969:63, 65.—Serene and Romimohtarto,<br />
1969:6 [discussion].—Bas, Arias, and Guerra,<br />
1976, table 3.—Tiirkay, 1976a:25 [listed], 37.<br />
Dorippe armata.—Monod, 1956, fig. 103.—Crosnier, 1964, fig.<br />
on pi. A.—Maurin, 1968b, figs. 5, 7. [Not Phyllodorippe<br />
armata (Miers, 1881).]<br />
Dorripe lanata.—Rossignol, 1957:126 [key].—Crosnier, 1970:<br />
1215. [Erroneous spelling.]<br />
SYNONYM.—Dorippe qffinis Desmarest, 1823.<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 68, 70 m, broken shell, 16\ 1$ ov, 6 juv (L).<br />
Ivory Coast: Sta 42, 62-75 m, mud with brown, branched<br />
Foraminifera, 2 juv (W). Sta 46, 38-42 m, mud with dense<br />
Jullienella, lie?, 11$, 27 juv (L). Sta 47, 37 m, bottom with<br />
Jullienella, 46*, 19 (W). Sta 60, 79-82 m, coral or rock, 1(5, 1<br />
juv (L). Sta 62, 46 m, brown, branching and foliate Foraminifera,<br />
19(5, 89 (W). Sta 64, 68 m, 19 (L). Sta 65, 46-49 m,<br />
26, 49 (W).<br />
Ghana: Sta 16, 46 m, mud with Foraminifera, shells, 1(5<br />
(W). Sta 23, 42 m, foliate brown to orange bryozoans, 36, 29<br />
(L). Sta 24, 35-37 m, dark red bryozoans, 1(5 (L). Sta 28, 49-<br />
53 m, 26, 19 (W). Sta 30, 61-64 m, coral, 1(5, 19 (L).<br />
Nigeria: Sta 241, 59-63 m, mud and shell, 4c?, 19 (W). Sta<br />
248, 33 m, 26 (L).<br />
Cameroon: Sta 260, 46 m, 1(5, 19 (W).<br />
Undaunted Material: Angola: Sta 102, 54 m, 19 (L). Sta<br />
103, 90 m, 16 (L).<br />
Other Material: Morocco: Off Cap Hadid, 31°55'N, 09°-<br />
52'W, 78 m, muddy sand, 5m beam trawl, 25 Mar 1976,<br />
Onversaagd Sta 127, 1(5 (L).<br />
DESCRIPTION.—Monod, 1933b:490-494; Capart,<br />
1951:30.<br />
Figures: Capart, 1951, fig. 6; Monod, 1956, fig.<br />
103.<br />
Male Pleopod: Monod, 1933b, fig. 3f,g (Morocco,<br />
Syria) (Figure 4g,h).<br />
Color: Capart (1951:31) noted that his material<br />
was "uniforme ocre rose, la carapace souvent<br />
encrassee de vase; les doigts de la pince roses."<br />
The Pillsbury specimens also were uniform pinkish<br />
in color, without obvious bands on the pereiopods.<br />
MEASUREMENTS.—Carapace lengths of our<br />
specimens range from 4 to 28 mm; the single<br />
ovigerous female has a carapace length of 28 mm.<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
REMARKS.—The <strong>West</strong> <strong>African</strong> specimens of<br />
Medorippe lanata closely resemble those from localities<br />
in the Mediterranean as well as those from<br />
the Atlantic coasts of Spain and Portugal with<br />
which they have been compared. The male pleopods<br />
of our specimens apparently are identical<br />
with those of Mediterranean specimens, resembling<br />
those illustrated by Monod (1933b, fig. 3f,<br />
6* from Morocco) (Figure 4g); the pleopods of<br />
immature males are like that figured by Monod<br />
(1933b: fig. 3g) (Figure Ah) from a specimen from<br />
Syria; Monod questioned whether his specimens<br />
from the eastern Mediterranean actually were<br />
mature.<br />
In spite of their close resemblance, some differences<br />
can be observed between our material from<br />
the Gulf of Guinea and specimens from more<br />
northern localities, but these observed differences<br />
overlap so broadly that we believe that the Gulf<br />
of Guinea and the Mediterranean populations of<br />
this species cannot be separated nomenclaturally<br />
at this time. Perhaps with additional material,<br />
distinct northern and southern subspecies can be<br />
recognized. In general, the Mediterranean specimens<br />
appear to be more pubescent, the spines on<br />
the merus of the walking legs are larger and give<br />
the impression of being less numerous, the teeth<br />
on the cutting edges of the chelae are sharper,<br />
and the walking legs appear to be longer and<br />
slenderer. The second and third pereiopods of a<br />
specimen from Tunisia (Figure Ac,d) and one<br />
from the Ivory Coast (Figure AeJ) are shown<br />
here. Of these apparent differences, the first three<br />
vary widely and cannot be used to distinguish<br />
representatives of the two populations. Mediterranean<br />
specimens have 7-15 (usually 10-11)<br />
spines on the dorsal edge of the merus, whereas<br />
the Gulf of Guinea specimens have 7 to approximately<br />
20 (usually 12) spines. The most striking<br />
difference is in the proportion of length to width<br />
of the merus of the walking legs (Table 1). The<br />
Mediterranean specimens appear to have much<br />
slenderer and longer legs, and this is in general<br />
supported by measurements, although for each<br />
size range the ratios overlap to some extent.<br />
The Gulf of Guinea population may mature at
NUMBER 306<br />
FIGURE 4.—Medorippe lanata (Linnaeus): a, dorsal view (from Monod, 1956, fig. 103); b, dorsal<br />
view of carapace (from Monod, 1933b, fig. 4c); c, second pereiopod, male, cl 23 mm, Tunis; d,<br />
third pereiopod, same specimen; e, second pereiopod, male, cl 19 mm, Pillsbury Sta 47;/, third<br />
pereiopod, same specimen; g, h, male, first pleopods (from Monod, 1933b, fig. 3f,g). Phyllodorippt<br />
armata (Miers): i, dorsal view (from Monod, 1956, fig. 102);^, dorsal view of carapace (from<br />
Monod, 1933b, fig. 4a); k, I, male first pleopods and apexes (from Monod, 1933b, fig. 3h).<br />
b<br />
33
34<br />
TABLE 1.—Length-width ratios of the propodus of the third<br />
pereiopods for M. lanala from the Gulf of Guinea and the<br />
Mediterranean<br />
Carapace<br />
length<br />
(mm)<br />
8-10<br />
11-13<br />
14-16<br />
17-19<br />
20-22<br />
23-25<br />
26-28<br />
Number<br />
of<br />
specimens<br />
3<br />
11<br />
12<br />
1<br />
3<br />
-<br />
-<br />
Gulf of Guinea<br />
Range<br />
(mm)<br />
4.2-4.9<br />
3.8-4.9<br />
3.7-4.7<br />
-<br />
3.3-3.8<br />
-<br />
-<br />
Mean<br />
(mm)<br />
4.6<br />
4.4<br />
4.1<br />
3.9<br />
3.5<br />
-<br />
-<br />
Number<br />
of<br />
specimens<br />
1<br />
-<br />
4<br />
2<br />
13<br />
11<br />
8<br />
Mediterranean<br />
Range<br />
(mm)<br />
_<br />
-<br />
4.5-5.5<br />
4.3-4.6<br />
3.8-5.1<br />
3.8-4.9<br />
4.1-4.8<br />
Mean<br />
(mm)<br />
4.6<br />
-<br />
5.1<br />
4.5<br />
4.5<br />
4.4<br />
4.3<br />
a smaller size. A 22 mm long male from Angola<br />
has the right cheliped greatly inflated, whereas in<br />
specimens from Spain males with carapace<br />
lengths of 23-26 mm show only slight inflation of<br />
the chela and smaller ones, 18-22 mm long, have<br />
chelae similar to those of females.<br />
Monod (1933b:492) suggested that the eastern<br />
Mediterranean population of this species might<br />
be subspecifically distinct from those in the western<br />
Mediterranean and the Atlantic. One of the<br />
differences he noted was in the shape of the male<br />
pleopod, but, as already pointed out, his Syrian<br />
specimens were juveniles and the pleopods were<br />
not fully developed. He also noted that the Syrian<br />
specimens differed from others he examined in<br />
having a more spinulose sternal border; this feature<br />
varies widely in our material, too.<br />
As indicated by Forest and Guinot (1966:50),<br />
figures 102 and 103 of Monod's (1956) monograph<br />
are interchanged, figure 102 showing Phyllodorippe<br />
armata, figure 103 Medorippe lanata. Crosnier's<br />
(1964) figure of "Dorippe armata" on his<br />
plate A, as well as Maurin's (1968b) figures 5D.a.<br />
and 7D.a., all are copies of Monod's (1956) figure<br />
103, and thus do not represent Phyllodorippe armata,<br />
but Medorippe lanata.<br />
BIOLOGY.—The depth at which this species<br />
occurs is usually indicated as sublittoral to about<br />
100 m (Bouvier, 1940:200; Monod, 1956:92). The<br />
depth of 952-1038 m off Morocco given by Tiirkay<br />
(1976a) is far in excess of the depths at which<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
the species normally is found and is probably<br />
erroneous. Zariquiey Alvarez (1968:313) indicated<br />
that this species is usually taken in Spanish<br />
waters in depths of 40 or 50 to 100 m; and Pesta<br />
(1918:287) gave these depths for the Adriatic<br />
population, with the reservation that the species<br />
may occur in depths of 10 m. So far as <strong>West</strong><br />
Africa is concerned, evidently the same situation<br />
is found: Crosnier (1964) listed the species as<br />
occurring in cold water in depths greater than 50<br />
m off Cameroon, whereas Le Loeuff and Intes<br />
(1968) stated that although Medorippe lanata is<br />
sometimes taken in Ivory Coast waters in depths<br />
between 25 and 30 m (in their table they give 15<br />
m as least depth; this may be an error for 25), the<br />
majority of the specimens there were found in<br />
depths between 35 and 50 m and as deep as 100<br />
m. Of the specimens examined by us only one<br />
was found at a depth less than 35 m (viz. 33 m),<br />
10 specimens (55%) were found between 35 and<br />
55 m, and the remainder were from deeper water,<br />
55-90 m. Also, more than 80% of the material<br />
listed by Monod (1956) for which depth is indicated<br />
was from depths of 35 to 55 m, whereas<br />
only two specimens, the remainder, were taken at<br />
96 and 100 m. Capart's <strong>West</strong> <strong>African</strong> material for<br />
the larger part (14 of the 17 lots for which depth<br />
was recorded) came from between 30 and 92 m,<br />
and only two lots were from less than 30 m (12-<br />
15 m and 20-24 m). Most other <strong>West</strong> <strong>African</strong><br />
records also conform to this pattern: the far<br />
greater part is from deeper than 35 m. Only<br />
Maurin (1968a) reported the species from a depth<br />
of 10-30 m, and only Tiirkay (1976a) has reported<br />
it from much deeper water, as noted<br />
above.<br />
Capart (1951) reported the occurrence of this<br />
species on coastal bottoms of mud or muddy sand<br />
generally in depths between 25 and 100 m, with<br />
one capture at 15 (12-15) m. It was taken only<br />
on one occasion with the shallower Phyllodorippe<br />
armata.<br />
The bottom types on which this species was<br />
collected by the Pillsbury include: mud with Foraminifera,<br />
46 m; foliate brown to orange bryozoans<br />
in 42 m and dark red bryozoans in 35-37
NUMBER 306<br />
m; coral in 61-64 m and coral and rock in 79-82<br />
m; broken shell in 70 m; and mud and shell in<br />
59-63 m. Most of the Pillsbury specimens were<br />
taken on mud bottom with foliate Foraminifera,<br />
probably Jullienella, in depths between 37 and<br />
75 m.<br />
Other bottom types on which the species has<br />
been collected include: mud and sand in 65-<br />
75 m, mud in 50-64 m, mud, stones, calcareous<br />
algae, sand, and Foraminifera in 51 to 55 m and<br />
gravel, shell, and Foraminifera in 50 m (all Forest<br />
and Guinot, 1966); sand and muddy sand in 10-<br />
30 m and muddy sand in 40-50 m (Maurin,<br />
1968a); shelly mud, in 50 m (Maurin, 1968b);<br />
and mud and sandy mud in 15[25?]-100 m (Le<br />
Loeuff and Intes, 1968). Le Loeuff and Intes<br />
(1969:63) indicated that this species was a mud<br />
dweller off the Ivory Coast. It seems that this<br />
species, like E. vossi, prefers a soft bottom with<br />
hard particles in it.<br />
Off <strong>West</strong> Africa, ovigerous females have been<br />
collected in March, April, May, June, August,<br />
and October (Capart, 1951; Monod, 1956;<br />
Guinot and Ribeiro, 1962; Pillsbury).<br />
DISTRIBUTION.—Eastern Atlantic, from the<br />
Mediterranean Sea and Portugal to South Africa<br />
and Mozambique (Barnard, 1955). Monod<br />
(1956) enumerated earlier records from <strong>West</strong> Africa;<br />
since 1956 M. lanata has been recorded from<br />
the following localities:<br />
Morocco: 33°37.5'N, 09°02.2'W, 952-1038 m (?) (Tiirkay,<br />
1976a). Agadir, 60-130 m (Forest and Games, 1960);<br />
Agadir (Maurin, 1968a). Between Cap Rhir (as Cap Ghir)<br />
and Cap Draa (as Cap Noun), 10-30 m (Maurin, 1968a).<br />
Off Essaouira, 32°08'N, 09°02'W, 33 m, and 31°37'N, 09°-<br />
54'W, 70 m (Forest and Games, 1960). SW of Cap Juby,<br />
40-50 m (Maurin, 1968a). Cap Juby, 50 m (Maurin, 1968b).<br />
Spanish Sahara: 21°44'N, 17°03'W to 21°46.5'N, 17°-<br />
04'W, 40 m, and 21°3O'N, 17°24'W to 21°27'N, 17°24'W,<br />
95-137 m (Bas, Arias, and Guerra, 1976).<br />
Mauritania: Bane d'Arguin, 40-60 m and 90-100 m<br />
(Maurin, 1968b).<br />
Senegal: 13°01'N, 17°24'W, 51-55 m, and 12°55.5'N,<br />
17°33'W, 65-75 m (Forest and Guinot, 1966).<br />
Sierra Leone: 07°15.5'N, 12°51'W, 64 m (Forest and<br />
Guinot, 1966).<br />
Ivory Coast: No specific locality (Le Loeuff and Intes,<br />
1969). Off Sassandra, off Fresco, off Grand-Lahou, off Jac-<br />
queville, and off Grand-Bassam, 15[25?]-100 m (Le Loeuff<br />
and Intes, 1968).<br />
Ghana: Off Accra, 37-51 m (Gauld, 1960). 04°40'N,<br />
02°08'W to 04°39'N, 02°05'W, 50 m, and 04°36.5'N, 01°-<br />
31'W, 50 m (Forest and Guinot, 1966).<br />
Cameroon: No specific locality, more than 50 m (Crosnier,<br />
1964).<br />
Congo: Near Pointe-Noire (Rossignol, 1957, 1962).<br />
Angola: 17°O2'S, ll°40'E, 54 m, and 17°06'S, 11°35'E,<br />
90 m (Crosnier, 1970).<br />
Genus Phyllodorippe, new genus<br />
TYPE-SPECIES.—Dorippe armata Miers, 1881.<br />
ETYMOLOGY.—The name is formed by a combination<br />
of the Greek word phyllon (leaf) and the<br />
feminine name Dorippe.<br />
DIAGNOSIS.—Carapace distinctly broader than<br />
long, with sparse pubescence dorsally and distinct<br />
epibranchial spine on either side. Extra-orbital<br />
teeth reaching slightly farther forwards than<br />
front. Tubercles on dorsal surface low, grooves<br />
distinct. Cardiac region without V-shaped ridge.<br />
Eyes not reaching beyond extra-orbital teeth.<br />
Lower margin of orbit between extra- and infraorbital<br />
teeth smooth. Dorsal margin of pereiopods<br />
2 and 3 without spines on dorsal margin of merus.<br />
Dactyli of these pereiopods narrow and slightly<br />
twisted; with fringe of short hairs present in basal<br />
half of upper margin. Abdomen of male with<br />
blunt low elevations, but without teeth or spines.<br />
First pleopod of male long, slender, S-shapedly<br />
curved, with 2 short appendages in distal part,<br />
lacking lobe in proximal part.<br />
REMARKS.—In the shape of the male gonopod,<br />
Phyllodorippe is closest to the genera Neodorippe and<br />
Nobilum, in both of which the male gonopod is<br />
also slender and curved; the gonopod of Phyllodorippe<br />
differs from those of the other genera<br />
because it ends in a narrow acute point and<br />
carries two lobiform appendages. The presence of<br />
epibranchial spines distinguishes the present genus<br />
from all other dorippine genera except Dorippe<br />
and Medorippe. Furthermore, in Neodorippe and<br />
Nobilum the front reaches farther forward than the<br />
anterolateral teeth. So far the genus is represented<br />
by a single species.<br />
35
36<br />
* Phyllodorippe armata (Miers, 1881),<br />
new combination<br />
FIGURE 4/-/<br />
?Dromia Pechiiel-Loesche, 1882:288.<br />
Dorippe Senegalensis Monod, 1933b:548 [footnote; nomen nudum,<br />
published in synonymy].<br />
Dorippe armato.—Monod, 1933b:548 [footnote; erroneous<br />
spelling].<br />
Dorippe armata.—Capart, 1951:33, fig. 7.—Monod, 1956:92,<br />
figs. 102 [as Dorippe lanata], 104-107 [not fig. 103 =<br />
Medorippe lanata].—Rossignol, 1957:75.—Longhurst, 1958:<br />
87— Lebour, 1959:131, 132, 134-136, fig. 19 [larvae].—<br />
Gauld, 1960:68—Rossignol, 1962:114.—Guinot and Ribeiro,<br />
1962:24.—Crosnier, 1964:32, 38 [not fig. on pi. A<br />
= Medorippe lanata].—Forest and Guinot, 1966:50.—Le<br />
Loeuff and Intes, 1968:38, figs. 47, 61, tables 1, 3-6, 8;<br />
1969:63-65.—Maurin, 1968a:48, 59, 62; 1968b:484, 486,<br />
489, 491 [not figs. 5D.a. and 7D.a. = Medorippe lanata].—<br />
Serene and Romimohtarto, 1966:6—Monod, 1970:66.<br />
Dorrippe armata.—Rossignol, 1957:126 [key; erroneous spelling]<br />
Dorippe armate.—Lebour, 1959:136 [larvae; erroneous spelling]<br />
Dorippe.—Voss, 1966:19.—Maurin, 1968b, figs. 4, 9.<br />
MATERIAL EXAMINED.—Pillsbury Material: Ivory<br />
Coast: Sta 46, 38-42 m, mud with dense Jullienella, 3$ (L).<br />
Sta 47, 37 m, bottom with Jullienella, 96, 4$ ov (L). Sta 48,<br />
22 m, 126\ 16$ (13 ov) (W). Sta. 49, 73-77 m, 1
NUMBER 306<br />
28 specimens of the species were obtained) and<br />
accidentally remained in the net.<br />
Of the post-1956 records listed under "Distribution,"<br />
those by Maurin (1968a, 1968b) are the<br />
deepest (1 record of 35-40 m, 6 records between<br />
50 and 100 m, and 1 of about 200 m). Forest and<br />
Guinot (1966) reported the species from depths<br />
between 4 and 50 m (all but one of these 9 records<br />
from 34 m or less). Le Loeuff and Intes (1968:38)<br />
remarked that "Z). armata est la plus cotiere [of<br />
the two Dorippe species] et ne depasse pas les 35<br />
m," although in their table 1 they give the depth<br />
of the species as 15-50 m. Guinot and Ribeiro<br />
(1962) reported the species from very shallow<br />
localities (5 and 8 m), but also from 42 m. Other<br />
authors indicated depth ranges up to 43 m<br />
(Gauld, 1960) or even 60 m (Longhurst, 1958). It<br />
seems likely that the species has its greatest concentration<br />
below 35 m, but also occurs occasionally<br />
as deep as 60 m. The very deep records of<br />
the species need verification.<br />
Inasmuch as Maurin (1968b) followed Monod<br />
(1956) and reversed the figures of M. lanata and<br />
P. armata and Maurin's records for the shallow<br />
water species, P. armata, are deeper than usual<br />
and his records for the deeper living species, M.<br />
lanata, are shallower than usual, it seems likely<br />
that the two species were reversed throughout his<br />
accounts (1968a,b).<br />
Phyllodorippe armata is usually found on a sand<br />
or mud bottom, with shell, calcareous algae, or<br />
Foraminifera. It seems to be able to tolerate<br />
low salinities. Balss (1921:48) reported it from<br />
"Flusswasser" (in Nigeria), from the "Nyanga-<br />
Fluss, Salzwasser" (Gabon), and "Siisswasser des<br />
Kongoflusses" (near Boma [05°51'S, 13°03'E],<br />
Zaire). Voss (1966:37) remarked that Pillsbury Sta<br />
250, at which P. armata was taken, had "a typically<br />
brackish water or estuarine fauna due to our<br />
proximity to the mouths of the Niger, or Kwara,<br />
which strongly influence this section of the coast."<br />
Sourie (1954b) found it on coarse shelly sand<br />
bottom with Area and Pyura, in 10-12 m in the<br />
Baie de Dakar.<br />
Crosnier (1964) found this species on mud or<br />
sandy mud off Cameroon and characterized it as<br />
a warm water species, living in 0-30 m. Longhurst<br />
(1958) reported if from muddy sand, shelly sand,<br />
and shelly mud in 10-60 m off Sierra Leone. Le<br />
Loeuff and Intes (1969:63) characterized this species<br />
as a coastal species indifferent to the nature<br />
of the substrate.<br />
Ovigerous females have been collected in January,<br />
February, March, April, May, June, August,<br />
September, November, and December (Capart,<br />
1951; Monod, 1956; Forest and Guinot,<br />
1966; and p. 36).<br />
Pechiiel-Loesche (1882:288) described the behavior<br />
of a crab that he observed at the Crique<br />
Banjia, Gabon, as follows: "Urn vieles komischer<br />
nimmt sich aber eine andere Krabbenart<br />
(Dromia?) aus, die ich am Banya beobachtete;<br />
beim Spazierengehen pflegt sie mittelst der hinteren,<br />
am Riicken entspringenden Beinpaare einen<br />
Sonnenschirm iiber sich zu halten, welcher<br />
gewohnlich aus einem halben Mangrovenblatte<br />
besteht. Fliichtet sie eilig, oder geht sie ins Wasser,<br />
so lasst sie das wunderliche Schutzdach fallen."<br />
As no morphological details of this crab are provided<br />
by Pechiiel-Loesche, it is impossible to state<br />
with certainty which species was observed by<br />
him. We know of no Dromia carrying leaves, but<br />
Serene and Romimohtarto (1969:12), when dealing<br />
with Neodorippe callida, remarked that "in<br />
Singapore the species is very common on the<br />
mangrove swamps and generally all specimens<br />
hook a leaf of a mangrove tree on their backs." If<br />
Pechiiel-Loesche's specimen actually is a dorippid,<br />
it is more likely to be Phyllodorippe armata than<br />
Medorippe lanata, as the former is known to live in<br />
shallower depths than the latter. New field observations<br />
may solve the problem and could add<br />
much of interest to the knowledge of the biology<br />
of these little known animals.<br />
DISTRIBUTION.—Off <strong>West</strong> Africa, where it has<br />
been recorded from many localities between Cabo<br />
Corbeiro, Spanish Sahara and Angola. Monod<br />
(1956) summarized earlier records and added<br />
many new ones. Since 1956, P. armata has been<br />
recorded from the following localities:<br />
Spanish Sahara: Between Cabo Corbeiro and Cabo<br />
Blanco, 60-80 m (Maurin, 1968a), in about 200 m (Maurin,<br />
37
38<br />
1968b). Between Cabo Barbas and Cabo Blanco, 50-90 m<br />
(Maurin, 1968b).<br />
Mauritania: N Bane d'Arguin, 20°20'N-20°40'N, 90-100<br />
m (Maurin, 1968a); same, less than 45 m, 40-60 m, 60-70<br />
m, and 90-100 m (Maurin, 1968b). Tamzak (as Tamxat),<br />
70-75 m (Maurin, 1968a), in 70 m (Maurin, 1968b).<br />
Senegal: Mboro, 35-40 m (Maurin, 1968b).<br />
Sierra Leone: No specific locality, 10-60 m (Longhurst,<br />
1958). O7°2O'N, 12°39'W, 30-34 m (Forest and Guinot,<br />
1966).<br />
Ivory Coast: No specific locality (Le LoeufT and Intes,<br />
1969). 05°00'N, 05°28.5'W, 27 m, and 05°03'N, 05°25'W,<br />
20-25 m (Forest and Guinot, 1966). Off Sassandra, off<br />
Fresco, off Grand-Lahou, and off Grand-Bassam, 15-50 m<br />
(Le Loeuff and Intes, 1968).<br />
Ghana: 04°40'N, 02°08'W to 04°39'N, 02°05'W, 50 m<br />
(Forest and Guinot, 1966). Off Accra and Takoradi, 10-43<br />
m (Gauld, 1960).<br />
Cameroon: No specific locality, 0-30 m (Crosnier, 1964).<br />
Principe: Between Ponta da Mina and Ilheu Santana,<br />
10-12 m; off (Cais de) Santana, 11 m; between Ponta da<br />
Mina and Ponta Novo Destino, 6 m (Forest and Guinot,<br />
1966).<br />
Sao Tome: Off Ponta Diogo Nunes, 4 m (Forest and<br />
Guinot, 1966).<br />
Gabon: ?Crique Banjia (as Banya) (Pechiiel-Loesche,<br />
1882).<br />
Congo: Off Pointe-Noire (Rossignol, 1957, 1962).<br />
Angola: Porto Amboim, 42 m; Baia de Benguela, 5 m;<br />
Baia da Caota, Benguela, 8 m (Guinot and Ribeiro, 1962).<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
* Ethusa rosacea A. Milne Edwards and<br />
Bouvier, 1897<br />
Ethusa rosacea.—Capart, 1951:28, fig. 5.—Monod, 1956:88.—<br />
Rossignol, 1957:126 [key].—Crosnier, 1967:323; 1969:530,<br />
figs. 20-22.<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 74, 641-733 m, l?ov(L).<br />
Ivory Coast: Sta 41, 641-842 m, 26* (W). Sta 44, 403-586<br />
m, hard, dark gray mud, lc? (L).<br />
DESCRIPTION.—Capart, 1951:28.<br />
Figure: Capart, 1951, fig. 5.<br />
Male Pleopod: Crosnier, 1969, figs. 20-22 (Angola).<br />
Color: Capart (1951:28) gave the following account<br />
of color pattern: "Couleur brun clair, sauf<br />
la face ventrale et les pinces, qui sont blanc-rose."<br />
MEASUREMENTS.—Our male specimens have<br />
carapace lengths of 8 to 11 mm; the ovigerous<br />
female has a carapace length of 12 mm.<br />
REMARKS.—Crosnier (1969) figured the first<br />
male pleopod of this species and commented on<br />
its distinctness from that of E. rugulosa, figured in<br />
Monod (1956:140). Some of the apparent differences<br />
between the first pleopods of these two<br />
species may be the result of their having been<br />
Lebour (1959:136) recorded larvae of this spe- illustrated from slightly different angles.<br />
cies from several localities in the Gulf of Guinea: BIOLOGY.—Ethusa rosacea is a deep-water spe-<br />
Ivory Coast, 04°29'N, 06°41'W; Ghana, 05°29'N, cies, occurring generally in depths below 100 m.<br />
00°20'E and 05°37'N, 00°38'E; and Nigeria, off One of the Pillsbury samples was taken on hard,<br />
Lagos, off Bonny River, and 05°58'N, 04°38'E. dark gray mud in 403-586 m. Material of this<br />
species reported by Crosnier (1969) was taken on<br />
mud in depths between 545 and 602 m. Capart<br />
Subfamily ETHUSINAE Guinot, 1977 (1951) reported the species from brown muddy<br />
sand in 125 m; brown, black mud in 100 m;<br />
Genus Ethusa Roux, 1830<br />
sandy mud in 290-390 and 500 m; and green<br />
sandy mud in 280-420 m.<br />
Ethusa P. Roux, 1830, pi. 18 [type-species: Cancer mascarone Ovigerous females have been recorded in<br />
Herbst, 1785, by subsequent designation by Fowler, 1912: March, April, and June (Capart, 1951; Crosnier,<br />
590; gender: feminine; name 1622 on Official List]. 1969; Pillsbury).<br />
?Pndope Nardo, 1847a:2 [nomen nudum; without included<br />
DISTRIBUTION.—Eastern Atlantic, where it has<br />
species).<br />
Pridope Nardo, 1869:307 [type-species: Pridope typica Nardo,<br />
been recorded from localities between the Canary<br />
1869, a subjective junior synonym of Cancer mascarone Islands and Luanda, Angola, in depths between<br />
Herbst, 1785, by original designation by use of typica and 100 and 1113 m (Monod, 1956). Since 1956 it has<br />
monotypy; gender: feminine).<br />
been recorded from the following localities.
NUMBER 306 39<br />
Congo: Off Pointe-Noire, 05°04'S, 1 l°20'E, 500 m (Crosnier,<br />
1967).<br />
Angola: 08°35'S, 12°51'E, 545-555 m; 09°27'S, 12°38'E,<br />
545-555 m; 11°55'S, 13°20'E, 545-552 m; and 11°57'S,<br />
13°29'E, 595-602 m (all Crosnier, 1969).<br />
It has not been recorded previously from the<br />
Ivory Coast or Liberia.<br />
* Ethusa rugulosa A. Milne Edwards and<br />
Bouvier, 1897<br />
Ethusa rugulosa.—Monod, 1956:89, figs. 99-101.—Longhurst,<br />
1958:87.—Rossignol, 1962:114.—Forest and Guinot,<br />
1966:49.—Crosnier, 1970:1215.<br />
MATERIAL EXAMINED.—Pillsbury Material: Ivory<br />
Coast: Sta 49, 73-77 m, 1$ (L). Sta 50, 128-192 m, 29 (W).<br />
Sta 60, 79-82 m, coral or rock, 1 juv (L).<br />
Other Material: Ivory Coast: Off Grand-Lahou, 20 m,<br />
dredge, 31 Mar 1964, Guinean Trawling Survey, Tr 24, Sta<br />
1, 1
40<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 5.—Ethusa vossi, new species, dorsal view (from Monod, 1956, fig. 90).<br />
56, 17$ (L). Sta 47, 37 m, bottom with Jullienella, 1(5, 1$ ov<br />
(W).<br />
Ghana: Sta 22, 51 m, rough bottom, 1$ ov (W). Sta 23,<br />
42 m, foliate brown to orange bryozoans, 1
NUMBER 306 41<br />
FIGURE 6.—Ethxtsa vossi, new species, male paratype, cl 9 mm, Pillsbury Sta 248: a, dorsal view;<br />
b, terminal two somites of abdomen; c, palm of chela. Ethusa mascarone (Herbst), male, cl 10 mm,<br />
Naples: d, dorsal view; e, terminal two somites of abdomen; /, palm of chela.<br />
and merus of third and fourth legs. Terminal<br />
somite of male abdomen shorter than penultimate<br />
(Figure 6b).<br />
Male Pleopod: Illustrated in Figure la,b.<br />
Color: Our specimens had the legs banded with<br />
red or orange and white. In males the enlarged<br />
right cheliped is white, with the tips of the fingers<br />
brown. Monod (1956) recorded one specimen<br />
with irregular white spots on the branchial regions.<br />
MEASUREMENTS.—Our specimens have carapace<br />
lengths of 4 to 11 mm. The ovigerous females<br />
have carapace lengths of 10 mm.<br />
REMARKS.—Ethusa vossi closely resembles E.<br />
mascarone (Herbst) from the Mediterranean and<br />
adjacent Atlantic; this new species had been identified<br />
with E. mascarone in the past, but it differs<br />
in several important respects. In E. vossi the terminal<br />
somite of the male abdomen is shorter than<br />
the penultimate somite (Figure 6b), whereas in E.<br />
mascarone the terminal somite is the longer (Figure<br />
6
42<br />
the orbit is smoother in E. mascarone (Figure 6d)<br />
than in E. vossi (Figure 6a), and females of the<br />
two species can be distinguished by this feature.<br />
Miers (1881a:270), in identifying material of<br />
this new species from Goree, Senegal, with E.<br />
mascarone, noted that "the larger chelipede in the<br />
male has the palm deeper and externally somewhat<br />
more convex than the male from the Mediterranean<br />
in the [British] Museum collection."<br />
In addition to the tropical <strong>West</strong> <strong>African</strong> records<br />
cited below (and assigned to E. vossi), E. mascarone<br />
has been recorded from the following localities<br />
outside of the Mediterranean: Canary Islands (A.<br />
Milne Edwards and Bouvier, 1900; Miers,<br />
1881a); Cape Verde Islands (A. Milne Edwards<br />
and Bouvier, 1900); and Cabo Blanco, Spanish<br />
Sahara (Cap Blanc, Mauritania) (Monod, 1933b;<br />
Tiirkay, 1975a). These materials will have to be<br />
reexamined to determine whether E. mascarone or<br />
E. vossi is involved.<br />
TYPE-LOCALITY.—Gulf of Guinea, off Nigeria,<br />
04°03'N, 05°41'E to 04°07'N, 05°40'E, Pillsbury<br />
Sta 248.<br />
DISPOSITION OF TYPES.—The holotype (Crust.<br />
D. 31538), a male, cl 8 mm, is in the Rijksmuseum<br />
van Natuurlijke Historie, Leiden. The other specimens<br />
are paratypes; they have been divided<br />
between the Rijksmuseum van Natuurlijke Historie<br />
in Leiden and the National Museum of<br />
Natural History, <strong>Smithsonian</strong> <strong>Institution</strong>, Washington.<br />
ETYMOLOGY.—It is a pleasure to name this<br />
species for Gilbert L. Voss of the Rosenstiel<br />
School of Marine and Atmospheric Science, University<br />
of Miami, Florida, who organized and led<br />
the Pillsbury expeditions to <strong>West</strong> Africa in 1964<br />
and 1965.<br />
BIOLOGY.—This species has been recorded from<br />
depths between 6 and 96 m, but 80% of the<br />
records are from depths of less than 45 m, and<br />
somewhat more than half from between 30 and<br />
45 m. The bottoms on which the species has been<br />
found have been described as mud with shells<br />
(Miers, 1881a); coarse shelly sand, bottom with<br />
Area and Pyura (Sourie, 1954b); partly shelly and<br />
partly muddy and shelly sand (Longhurst, 1958);<br />
mud and shells, mud, calcareous algae and shells,<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
algae and calcareous algae, gravel, shells and<br />
Foraminifera, and mud, calcareous algae and<br />
shells (all Forest and Guinot, 1966). The Pillsbury<br />
specimens were taken on bottoms with JullieneIla<br />
(Sta 46, 47) (Voss, 1966), broken shell (Sta 68),<br />
and branched Foraminifera (Sta 70). It seems,<br />
therefore, that the species prefers a muddy bottom<br />
on which solid particles (shells, Foraminifera,<br />
etc.) occur.<br />
Ovigerous females have been recorded in January,<br />
February, April, May, August, and September<br />
(Monod, 1956; Forest and Guinot, 1966; Pillsbury).<br />
DISTRIBUTION.—Off tropical <strong>West</strong> Africa,<br />
where it has been recorded from localities between<br />
Senegal and Angola; there is one record<br />
from Mauritania, which requires corroboration.<br />
Records in the literature include:<br />
Mauritania: Cap Blanc (?) (Monod, 1933b).<br />
Senegal: S of Mbao (Monod, 1956). Near Dakar,<br />
14°38.5'-14°41'N, 17°20.5'-17°23.5'W, 22-34 m (Monod,<br />
1956). Anse de Hann (as Baie de Hann) (Monod, 1956).<br />
Between Goree and Thiaroye-sur-Mer, ca. 15 m (Monod,<br />
1956). Near Goree, 25 m (Balss, 1921); same, 17-% m<br />
(Monod, 1956). Baie de Goree, 18-28 m (Miers, 1881a).<br />
Near Bane du Seminole, Baie de Goree, 38 m (Monod,<br />
1956). Near lie de la Madeleine, 35 and 40 m (Monod,<br />
1956).<br />
Guinea: 09°40'N, 14°05'W, 18 m (Forest and Guinot,<br />
1966). Near Conakry, 09°N, 13°5O'W, 30 m; 09°22'N, 13°-<br />
37^, 10 m; 3 mi NW of Tamara, 12 m; between Tamara<br />
and lie de Corail, 10 m (all Monod, 1956).<br />
Sierra Leone: No specific locality, 44 m (Monod, 1956);<br />
6-44 m (Longhurst, 1958).<br />
Ivory Coast: 05°07'N, 04°32'W to O5°O7'N, 04°36'W,<br />
43-38 m (Voss, 1966).<br />
Ghana: Off Accra, 16-44 m (Monod, 1956; Gauld, 1960).<br />
04°40'N, 02°08'W to 04°39'N, 02°05'W, 50 m, and<br />
04°36.5'N, 01°31'W, 50 m (Forest and Guinot, 1966).<br />
Principe: O1°37'N, 07°22'E, 30 m; 01°38'25"N, 07°-<br />
22'05"E, 31 m; and 01°43'10"N, 07°28'20"E, 73 m (Forest<br />
and Guinot, 1966).<br />
Sao Tome: 00°25'40"N, 06°40'10"E, 50 m (Forest and<br />
Guinot, 1966).<br />
Cabinda: No specific locality, 38-39 m (Guinot and<br />
Ribeiro, 1962).<br />
Angola: Off the mouth of the Congo River, 06°18'S,<br />
12°O2'E, 44 m (Doflein, 1904).<br />
Genus Ethusina Smith, 1884<br />
Ethusitu Smith, 1884:349 [type-species: Ethusina abyssicola<br />
Smith, 1884, by monotypy; gender: feminine].
NUMBER 306 43<br />
Key to Eastern Atlantic Species of Ethusina<br />
1. Outer orbital spine very small, directed laterally, not overreaching eye<br />
K alba<br />
Outer orbital spine large, directed anteriorly, overreaching eye 2<br />
2. Outer orbital spine extending to base of inner orbital spine . E. talismani<br />
Outer orbital spine extending to or beyond apex of inner orbital spine ...<br />
E. beninia, new species<br />
Ethusina alba (Filhol, 1884)<br />
FIGURES 8, 9d, 10<br />
Ethusa alba Filhol, 1884:199 [23 February; nomen nudum],<br />
230 [8 March; nomen nudum], 232 [8 March; available<br />
name].—Anonymous, 1884:7 [20 April].—Filhol, 1885b:<br />
129 [1 December].<br />
Ethusina abyssicola.—Bouvier, 1898:66 [listed].—A. Milne Ed-<br />
wards and Bouvier, 1899:18;-1900:29, pi. 1: fig. 6 [color;<br />
outline based on E. abyssicola Smith from Smith, 1884: pi.<br />
2: fig. 1].—Bouvier, 1922:53, pi. 2: fig. 1 [color].—Monod,<br />
1956:85[references only]. [Not Ethusina abyssicola Smith,<br />
1884.]<br />
Ethusa abyssicola.—Bouvier, 1922:91 [not Ethusina abyssicola<br />
Smith, 1884].<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: North Atlantic: Azores to France, 42° 15'N,<br />
23°37'W (of Paris, = 21°17'W of Greenwich), 3975 m, to<br />
42°19'N, 23°36'W (of Paris, = 21°16'W of Greenwich), 4060<br />
m, soft white mud, Talisman Sta 133, 134, 24 Aug 1883, Id,<br />
(holotype) 1$ (L,W).<br />
DESCRIPTION.—Carapace (Figures 8a, 10) longer<br />
than broad in male (width = 0.95 X length),<br />
broader than long in female (width = 1.03 X<br />
length), noticeably narrowing anteriorly. Carapace<br />
coarsely granular, at most lightly pubescent<br />
dorsally, few longer hairs present laterally. Cervical<br />
and cardiobranchial grooves distinct. Branchial<br />
regions moderately inflated. Front with 4<br />
upturned spines, submedians broad, blunt, laterals<br />
narrower, sharper, falling short of submedians<br />
in male (Figure 8a), overreaching submedians<br />
in female (Figure 8*). Front less than half greatest<br />
width of carapace. Outer orbital spine small,<br />
triangular, scarcely overreaching eye, not extending<br />
to front. Eyes short, stout, fixed, with terminal<br />
black cornea, narrower than stalk, visible in dorsal<br />
view. Basal segment of antennule inflated,<br />
surface irregular, not tuberculate. Basal article of<br />
antenna short, not extending to front. Antennal<br />
flagellum elongate, extending to posterior third<br />
of carapace. Ventral surface of body less granular<br />
but more pubescent than dorsal.<br />
Outer maxillipeds slender, merus narrower<br />
than, but more than half as long as, ischium.<br />
Chelipeds similar, equal, small, extending with<br />
merus to cornea in both sexes; propodus slightly<br />
larger in male than in female but not markedly<br />
inflated. Fingers (Figure 8b) longer than palm,<br />
slender, flattened, with shallow longitudinal<br />
groove, cutting edges crenulate.<br />
Second and third pereiopods similar, third<br />
longer, more than twice as long as chela, about<br />
2 1 A times as long as carapace. On each leg dactylus<br />
longer than propodus but shorter than car-<br />
FIGURE 8.—Ethusina alba (Filhol). Male, cl 10.4 mm, North<br />
Atlantic: a, front, dorsal view; b, chela; c, propodus and<br />
dactylus of third pereiopod, posterior view; d, abdomen.<br />
Female, cl 9.3 mm, North Atlantic: e, front, dorsal view; /,<br />
front, lateral view; g, abdomen.
44<br />
FIGURE 9.—Male pleopods: a, Ethusina beninia, new species,<br />
holotype, cl 14.0 mm, Pillsbury Sta 18; b, Ethusina robusta<br />
(Miers), cl 13.9 mm, Galapagos Islands; c, Ethusina talismani<br />
A. Milne Edwards and Bouvier, syntype, carapace broken,<br />
Morocco; d, Ethusina alba (Filhol), cl 10.4 mm, North Atlantic;<br />
e, Ethusina abyssicola Smith, cl 11.9 mm, Massachusetts.<br />
apace, subequal to or slightly shorter than merus,<br />
flattened dorsoventrally, grooved longitudinally,<br />
spatulate, slightly twisted, apex corneous; dactyli<br />
of both legs shorter than carapace in our material.<br />
Fourth and fifth pereiopods dissimilar, much<br />
shorter than second and third, both overreaching<br />
merus of third by length of dactylus; dactyli of<br />
last 2 legs very short, curved, apices corneous,<br />
with row of spinules on ventral margin; ischium<br />
of fifth pereiopod longer than that of fourth.<br />
Pereiopods with light surface pubescence, especially<br />
on fourth and fifth, latter also with some<br />
longer hairs.<br />
Male abdomen with 5 somites, that of female<br />
with 7 (Figure 8dg).<br />
Male Pleopod: Illustrated in Figure 9d.<br />
Color: "Cephalothorax bleuatre sur le vivant,<br />
avec une legere teinte violacee; pattes et abdomen<br />
d'un blanc jaunatre, avec les doigts de couleur<br />
rose" (A. Milne Edwards and Bouvier, 1900:30).<br />
Bouvier (1922:53) noted that material from the<br />
Cape Verde Islands was "jaune d'ocre pale avec<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
les doigts et les antennules roses" and that in<br />
material collected earlier by the Talisman the<br />
carapace was "uniformement bleuatre." The<br />
color of this species has been illustrated by A<br />
Milne Edwards and Bouvier (1900, pi. 1: fig. 6)<br />
and Bouvier (1922, pi. 2: fig. 1).<br />
MEASUREMENTS.—The single male examined<br />
has a carapace length of 10.4 mm, a carapace<br />
width of 9.9 mm; the only female examined<br />
measures 9.3 X 9.6 mm.<br />
REMARKS.—The present species was identified<br />
by A. Milne Edwards and Bouvier (1900:29) with<br />
Ethusina abyssicola Smith. Originally, however, A.<br />
Milne Edwards must have thought that the species<br />
was new and gave it provisionally the manuscript<br />
name Ethusa alba. Filhol (1884:199, 230,<br />
232), in a popular article on the Travailleur and<br />
Talisman expeditions, introduced, obviously unintentionally,<br />
the name Ethusa alba into the literature;<br />
A. Milne Edwards was cited as the author.<br />
In that paper, Filhol used the name on three<br />
occasions; in two of these (pp. 199, 230) the name<br />
was not accompanied by sufficient data to make<br />
it an available name. On page 232, however,<br />
Filhol remarked that Ethusa alba differed from<br />
other species of the genus by the presence of eyes<br />
("dans un meme genre Ton trouve des especes<br />
aveugles et d'autres qui ne le sont pas, ainsi<br />
YEthusa granulata [= Cymonomus granulatus] qui vit<br />
dans les mers du Nord entre 200 et 1300 metres<br />
FIGURE 10.—Ethusina alba (Filhol) (from Bouvier, 1922, pi. 1:<br />
fig. 1).
NUMBER 306 45<br />
est aveugle, alors que YEthusa alba que nous avons<br />
pris dans 1'Ocean par 5000 metres ne Test pas").<br />
Hereby the name Ethusa alba becomes available<br />
as from 8 March 1884 and Filhol has to be cited<br />
as the author. Since the publication of Filhol's<br />
paper, the name Ethusa alba has only been mentioned<br />
a few times in the literature. First in an<br />
anonymous guide to the "exposition sous-marine<br />
du Travailleur et du Talisman" issued for the<br />
"visiteurs de l'Exposition organisee sous la direction<br />
de M. Alph. Milne-Edwards, president de la<br />
Commission des dragages". This guide was a<br />
"publication du journal La Nature" and consists<br />
of an abstract of Filhol's 1884 paper, probably<br />
made by Gaston Tissandier, chief editor of the<br />
journal. The dates of the exposition are given on<br />
page 16 of the guide as follows:<br />
Cette exposition des collections du Travailleur et du Talisman<br />
a ete ouverte le 26 Janvier 1884 et fermee le 16 mars.<br />
On 1'a ouverte de nouveau le 7 avril, a l'occasion de la<br />
reunion des societes savantes a la Sorbonne, et elle sera<br />
fermee le 20 du meme mois.<br />
The fact that the last verb is written in the<br />
future tense shows that the catalog was written<br />
and probably published before 20 April 1884.<br />
The remark on Ethusa alba in the guide is verbally<br />
the same as the one made by Filhol (1884:199),<br />
viz., "les Ethuses dont une espece nouvelle,<br />
YEthusa alba, a ete prise a 5000 metres de profondeur."<br />
Filhol (1885b: 129) in his book La vie au<br />
fonds des mers, does not add anything new to the<br />
knowledge of the species. We have been unable<br />
to find any later reference to Ethusa alba in the<br />
literature, probably because A. Milne Edwards<br />
and Bouvier (1899) identified their specimens<br />
with Ethusina abyssicola S. I. Smith and used that<br />
name ever since.<br />
Apart from the fact that the species has eyes,<br />
the only information on it provided by Filhol is<br />
that it was taken by the Talisman at 5000 m<br />
(Filhol, 1884:199, 232; 1885b: 129) or "entre<br />
quatre et cinq mille metres" (Filhol, 1884:230).<br />
A. Milne Edwards and Bouvier (1900) in their<br />
account of the Talisman and Travailleur Brachyura,<br />
listed the following Ethusinae collected by the<br />
Talisman: Ethusa mascarone (Herbst) in 38 and 60<br />
m (Sta 56); E. rugulosa A. Milne Edwards and<br />
Bouvier in 150-275 m (Sta 103); E. rosacea A.<br />
Milne Edwards and Bouvier in 930 and 1013—<br />
1113m (Stas 85,87); Ethusina abyssicola S. I. Smith<br />
in 2995, 3655, 3975, and 4060 m (Stas 102, 131,<br />
133, 134); and E. talismani A. Milne Edwards and<br />
Bouvier in 2075, 2083, 2115, 2212, and 2235 m<br />
(Stas 40, 41, 43, 44, 130). Of Ethusina abyssicola<br />
they remarked (1900:30): "C'est, de tous les<br />
Crabes, le plus abyssal." Although none of the<br />
crabs of the Talisman expedition was collected at<br />
5000 m, as stated for Ethusa alba, Ethusina abyssicola<br />
is the only species then taken in a depth of more<br />
than 4000 m and would fit Filhol's statement that<br />
Ethusa alba was taken "entre quatre et cinq mille<br />
metres." The only Talisman station of 5000 m<br />
depth is no. 137 of 27 August 1883 (4975-5005<br />
m), but no crabs are recorded from it. Filhol's<br />
statement that Ethusa alba was taken at 5000 m<br />
either is an exaggeration or an error (he may have<br />
confused stations 134 and 137), or a careless<br />
abbreviation of 4000-5000 m. Confusion in the<br />
station numbers of the Talisman expedition occurred<br />
very frequently (the expedition is rather<br />
notorious because of it). So de Folin (1887:332)<br />
when dealing with station 136 (26 August 1883,<br />
depth 4255 m) mentioned from it "des Ethuses<br />
differentes de celles deja connues," although A.<br />
Milne Edwards and Bouvier (1900) do not mention<br />
any Ethusinae from that station. However<br />
this may be, there can be little doubt that the<br />
present species at first was thought by A. Milne<br />
Edwards to be a new species, Ethusa alba, and that<br />
he later changed his views and considered it<br />
identical with S. I. Smith's Ethusina abyssicola.<br />
Filhol's use of the name Ethusa alba in print made<br />
that name available and it has to be used, now<br />
that the specific distinctness of the Talisman specimens<br />
from S. I. Smith's Ethusina abyssicola has<br />
been demonstrated.<br />
In identifying material of this species collected<br />
by the Talisman at several different localities, A.<br />
Milne Edwards and Bouvier (1900:29) noted:<br />
"Nos specimens ressemblent completement a ceux<br />
qu'a decrits ou figures S.-I. Smith. Nous ferons<br />
remarquer, toutefois, que les doigts de leurs
46<br />
grandes pattes ambulatoires sont manifestement<br />
tordus, que leurs saillies rostrales sont tantot obtuses,<br />
tantot acuminees, enfin que leurs epines<br />
susorbitaires ont une longueur extremement variable.<br />
L'aire cardiaque est toujours assez distincte<br />
et les dents laterales du 3° segment abdominal<br />
sont peu saillantes."<br />
Our material ofE. alba differs from E. abyssicola<br />
in having much shorter frontal spines with their<br />
apices only slightly reflected dorsally. In E. abyssicola<br />
the frontal spines are much more prominent<br />
and their apices, especially of the outer pair, are<br />
strongly reflected dorsally, being situated at an<br />
angle of almost 45° from the longitudinal axis of<br />
the carapace. The most important difference between<br />
the two species is in the structure of the<br />
male pleopod, which in E. alba (Figure 9d) is<br />
slenderer, not sharply bent basally, with a distinct,<br />
corneous angled prominence apically; the<br />
apex of the pleopod in E. abyssicola is rounded or<br />
flattened distally (Figure 9*).<br />
As in E. abyssicola, the female of E. alba is<br />
comparatively broader and has much shorter<br />
frontal spines than the male.<br />
The figure of this species given by A. Milne<br />
Edwards and Bouvier (1900, pi. 1: fig. 6) is based<br />
on an outline of E. abyssicola published by Smith<br />
(1884).<br />
Coordinates for Talisman station 133 given by<br />
A. Milne Edwards and Bouvier (1900:30), 42°-<br />
45'N, 23°37 / W of Paris (- 21 O 17'W of Greenwich),<br />
may be in error, for that station was made<br />
on the same day as station 134 and the latitude<br />
for station 133 is given as 42°15'N in a list of<br />
Talisman stations prepared by the Paris Museum.<br />
These stations were so close to each other that the<br />
material from them may have been combined.<br />
The male specimen from Leiden has the following<br />
label data: 42°19 / N, 23°37'W [Paris], 4060 m,<br />
Talisman Sta. 134 [(it should read longitude<br />
23°36'W (Paris)], whereas that from Washington<br />
bears the coordinates 42°15'N, 23°37'W (Paris),<br />
3975-4060 m; both sets of data were originally<br />
based on longitude of Paris and are corrected to<br />
Greenwich (see "Distribution").<br />
BIOLOGY.—Ethusina alba, like E. abyssicola, is a<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
true abyssal species, living in depths exceeding<br />
1000 m. It has been collected on white mud with<br />
Foraminifera in 4261 m (A. Milne Edwards and<br />
Bouvier, 1899); soft white mud in 2995, 3975 and<br />
4060 m, and gray mud in 3655 m (A. Milne<br />
Edwards and Bouvier, 1900); and on muddy<br />
volcanic sand with Foraminifera in 3890 m (Bouvier,<br />
1922).<br />
Ovigerous females have not been recorded.<br />
DISTRIBUTION.—Eastern Atlantic, from the<br />
North Atlantic N of the Azores to the Cape Verde<br />
Islands, in depths between 2995 and 4261 m.<br />
Records in the literature include:<br />
North Atlantic between the Azores and France: 42°15'N,<br />
23°37'W of Paris (= 21°17'W of Greenwich), 3975 m, and<br />
42°19'N, 23°37'W of Paris (= 21°17'W of Greenwich), 4060<br />
m (A. Milne Edwards and Bouvier, 1900).<br />
Azores: 38°38'N, 27°26'W of Paris (= 25°06'W of Greenwich),<br />
2995 m (A. Milne Edwards and Bouvier, 1900).<br />
36°55'N, 24°43'W of Paris (= 22°23'W of Greenwich), 4261<br />
m (A. Milne Edwards and Bouvier, 1899).<br />
Cape Verde Islands: 16° 12'N, 24°43'45"W, 3890 m (Bouvier,<br />
1922). 15°48'N, 22°43'W of Paris (= 20°23'W of Greenwich),<br />
3655 m (A. Milne Edwards and Bouvier, 1900).<br />
* Ethusina beninia, new species<br />
FIGURES 9a, 11<br />
MATERIAL EXAMINED.—Pillsbury Material: Ghana:<br />
Sta 18, 3047-3129 m, soft, dark gray clay, 16* (holotype), 19<br />
(L). Sta 34, 1948-1984 m, mud, 19 (W).<br />
DESCRIPTION.—Carapace (Figure 1 la) slightly<br />
broader than long (1.05 times broader in male,<br />
1.01 to 1.05 times in females), noticeably narrowing<br />
anteriorly. Surface finely granular, lightly<br />
pubescent. Cervical and cardiobranchial grooves<br />
distinct, latter deeper, better defined. Branchial<br />
regions inflated. Intestinal region with pair of<br />
rounded prominences. Front (Figure 1 \a-d) with<br />
4 upturned spines, inner pair broader and shorter<br />
than outer, with sinus between inner spines<br />
broader and deeper than that between inner and<br />
outer spines. Inner frontal spines shorter, blunter<br />
in female than in male. Front less than half<br />
greatest width of carapace. Outer orbital spine<br />
strongly developed, extending to or beyond base<br />
of frontal sinus, apex curved upward, directed
NUMBER 306 47<br />
a d<br />
FIGURE 11.—Ethusina beninia, new species. Holotype, male, cl 14.0 mm, Pillsbury Sta 18: a, dorsal<br />
view; b, front, dorsal view; c, front, lateral view; g, abdomen. Paratype, female, cl 16.1 mm,<br />
Pillsbury Sta 18: d, front, ventral view; e, chela;/, dactylus of third pereiopod; h, abdomen.<br />
anterolaterally, broader and shorter in female<br />
than in male. Eyes (Figure 1 lc,d) short, stout,<br />
fixed, with terminal black cornea; eyes tapering<br />
distally, scarcely or not at all visible in dorsal<br />
view. Basal segment of antennule inflated, with<br />
1 or 2 anterior tubercles. Basal article of antenna<br />
short, not extending to front. Antennal flagellum<br />
elongate, extending to posterior fourth of carapace.<br />
Ventral surface of body more granular than<br />
dorsal.<br />
Outer maxillipeds slender, merus narrower<br />
than, but more than half as long as, ischium.<br />
Chelipeds similar, equal, small, slender, with<br />
propodus slightly stouter than carpus. Palm 1.44
48<br />
times longer than high in male, 1.33-1.34 times<br />
in female. Left chela broken on only intact male,<br />
meri of chelipeds on that male subsimilar in size<br />
and shape, intact cheliped slightly larger than<br />
those of female (Figure 11
NUMBER 306 49<br />
30°03'N, 14°02'W of Paris (= 11°42'W of Greenwich),<br />
in 2212 m, was shown on plate 10: figure<br />
9. Apparently the same male was figured in color<br />
on plate 3: figure 6, although according to the<br />
text a female was figured there (only one female<br />
was reported in the collection). According to the<br />
figure and to the account given in the text (A.<br />
Milne Edwards and Bouvier, 1900:31), "les femelles<br />
se distinguent des males par leur carapace<br />
plus large, par leurs pointes frontales plus courtes<br />
et plus obtuses . . . ," the female differed from the<br />
males in having the frontal spines shorter and<br />
broader, the lateral spines originating more anteriorly<br />
and extending further forward, and the<br />
eyes were not visible in dorsal view. In these<br />
features the female more closely resembles E.<br />
beninia, differing mainly in having shorter exorbital<br />
spines which are directed anteriorly. The<br />
female reported by A. Milne Edwards and Bouvier<br />
and E. beninia both differ from males of E.<br />
talismani in having eyes that are hidden in dorsal<br />
view and in having the exorbital spines originating<br />
nearer the anterior margin of the carapace.<br />
There is no indication of dimorphism to such an<br />
extent in our material of E. beninia, so it seems<br />
likely that the female from the Talisman collection<br />
may represent a distinct species.<br />
TYPE-LOCALITY.—Bight of Benin, off Ghana,<br />
05°01'N, 00° 12'E, in 3047-3129 m.<br />
DISPOSITION OF TYPES.—The male holotype<br />
(Crust. D. 31781) and female paratype from Pillsbury<br />
Sta 18 are in the Rijksmuseum van Natuurlijke<br />
Historie, Leiden. A female paratype from<br />
Pillsbury Sta 34 is in the National Museum of<br />
Natural History, <strong>Smithsonian</strong> <strong>Institution</strong>, Washington.<br />
ETYMOLOGY.—The specific name, beninia, alludes<br />
to its occurrence in the Bight of Benin.<br />
BIOLOGY.—This is a deep water species occurring<br />
between 1984 and 3047 m (the two stations<br />
at which it was collected being from 1948-1984<br />
m and 3047-3129 m). The bottom where the<br />
species was obtained in 3047-3129 m was soft,<br />
dark gray clay.<br />
DISTRIBUTION.—This species is known only<br />
from the Bight of Benin, off Ghana, at 05°01'N,<br />
00°12'E and 03°53'N, 02°33'W.<br />
Family CALAPPIDAE de Haan, 1833<br />
CALAPPIDEA de Haan, 1833:ix [corrected to Calappidae by<br />
White, 1847a:44; name 371 on Official List].<br />
MATUTOIDEA de Haan, 1835:36.<br />
ORITHYINAE Dana, 1852b:391.<br />
EASTERN ATLANTIC GENERA.—Four, Acanthocarpus,<br />
Calappa, Cycloes, and Matuta, all of which have<br />
representatives in tropical waters.<br />
EASTERN ATLANTIC SPECIES.—Seven, all of<br />
which were recorded by Monod (1956), as follows:<br />
Name in Monod<br />
Matuta michaelseni<br />
[Osachila stimpsoni<br />
(Calappidae)]<br />
Calappa gallus<br />
Calappa peli<br />
Calappa granulata<br />
Calappa rubroguttata<br />
Acanthocarpus brevispinis<br />
Cryptosoma cristatum<br />
Current Name<br />
Matuta michaelseni<br />
[Sakaila africana,<br />
new genus, new species*<br />
(Part henopi dae) ]<br />
Calappa gallus<br />
Calappa pelii*<br />
Calappa granulata<br />
Calappa rubroguttata*<br />
Acanthocarpus brevispinis*<br />
Cycloes cristata<br />
As noted above, we have placed <strong>West</strong> <strong>African</strong><br />
material identified with Osachila stimpsonii in Sakaila<br />
africana, new genus, new species, which has<br />
been assigned to the Subfamily Aethrinae, Family<br />
Parthenopidae (p. 322).<br />
We believe that one other species reported in<br />
the <strong>West</strong> <strong>African</strong> fauna by Monod (1956:151) is<br />
based on an erroneously labeled specimen. It is<br />
Hepatus princeps (Herbst, 1794): a western Atlantic<br />
species reported from "Guinea" by Rathbun<br />
(1937:237) based upon a single specimen in the<br />
Zoological Museum, Copenhagen.<br />
Subfamily CALAPPINAE de Haan, 1833<br />
Genus Acanthocarpus Stimpson, 1871<br />
Acanthocarpus Stimpson, 1871a: 152 [type-species:- Acanthocarpus<br />
alexandri Stimpson, 1871, by monotypy; gender: masculine].
50<br />
* Acanthocarpus brevispinis Monod, 1946<br />
Acanthocarpus africanus Capart, 1951:35, fig. 8.—Rossignol,<br />
1957:127 [key].<br />
Acanthocarpus brevispinis Monod, 1956:109, figs. 125-132.—<br />
Guinot-Dumortier and Dumortier, 1960:129 [stridulation].—Guinot<br />
and Ribeiro, 1962:27.—Forest, 1963:<br />
628.—Monod, 1963, fig. 32.—Maurin, 1968b:484, 492,<br />
fig. 6— Le Loeuffand Intes, 1968, table 1; 1969:66.<br />
Acanthocarpus bispinosus.—Longhurst, 1958:87 [not Acanthocarpus<br />
bispinosus A. Milne Edwards, 1880].<br />
Acanthocarpus brevispinnis.—Maurin, 1968b:489 [erroneous<br />
spelling].<br />
MATERIAL EXAMINED.—Pillsbury Material: Ivory<br />
Coast: Sta 51, 329-494 m, 1
NUMBER 306 51<br />
March and October (Capart, 1951; Monod,<br />
1956).<br />
DISTRIBUTION.—<strong>West</strong> Africa, where it has been<br />
recorded from a few localities between Cap Juby,<br />
Morocco [the type-locality] and Angola, in depths<br />
between 100 and 500 m. Monod (1956) studied<br />
material from Senegal, Gambia, and Guinea.<br />
Since 1956 it has been recorded from the<br />
following:<br />
Spanish Sahara: Off Villa Cisneros, 300-500 m; off<br />
Tamzak (as Tamxat), 200-400 m (both Maurin, 1968b).<br />
Mauritania: Bane d'Arguin, 200-300 m (Maurin, *968b).<br />
Senegal: Off Saint-Louis, 300 m; Fosse de Kayar, 250 to<br />
300-350 m (both Maurin, 1968b).<br />
Sierra Leone: No specific locality, 180-400 m (Longhurst,<br />
1958).<br />
Ivory Coast: No specific locality (Le Loeuff and Intes,<br />
1969); Off Grand-Bassam, 200 m (Le Loeuff and Intes,<br />
1968). 04°32'30"N, 06°31'W, 300-455 m (Forest, 1963).<br />
Ghana: 04°39'N, 02°46'W, 300-400 m (Forest, 1963).<br />
Angola: Ponta de Sao Jose, Baia Farta, Benguela, 235-<br />
310 m (Guinot and Ribeiro, 1962).<br />
Genus Calappa Weber, 1795<br />
Calappa Weber, 1795:92 [type-species: Cancer granulatus Linnaeus,<br />
1758, by subsequent designation by Latreille, 1810:<br />
422; gender: feminine; name 1611 on Official List].<br />
Camara de Haan, 1837:67 [type-species: Calappa fomicata Fabricius,<br />
1798, a subjective junior synonym of Calappa<br />
calappa Linnaeus, 1758, by monotypy; gender: feminine].<br />
Lophos de Haan, 1837:67 [type-species: Cancer lophos Herbst,<br />
1785, by tautonymy; gender: masculine].<br />
Gallus de Haan, 1837:67 [type-species: Cancer gallus Herbst,<br />
1803, by tautonymy and monotypy; gender: masculine].<br />
Pistor Gistel, 1848:ix [substitute name for Gallus de Haan,<br />
1837; type-species: Cancer gallus Herbst, 1803; gender:<br />
masculine].<br />
Calappa gallus (Herbst, 1803)<br />
Calappa gallus.—Monod, 1956:100, figs. 115, 116 [Cape<br />
Verde Islands, Senegal, Sierra Leone, Ghana].—Rossignol,<br />
1957:76, 127 [key], fig. 1 [Congo].—Longhurst, 1958:<br />
87 [Sierra Leone]—Gauld, 1960:68 [Ghana].—Guinot and<br />
Ribeiro, 1962:26 [Cape Verde Islands].—Rossignol, 1962:<br />
114 [Congo].—Ribeiro, 1964:4 [Cape Verde Islands].—<br />
Forest and Guinot, 1966:51 [Principe, Sao Tome, Annobon].—Guinot,<br />
1967a:245 [Indian Ocean, listed].—<br />
Monod, 1967:178 [no locality].—Ribeiro, 1973:5 [Cape<br />
Verde Islands].<br />
SYNONYMS.—Calappa galloides Stimpson, 1859;<br />
Calappa squamosa Desbonne and Schramm, 1867;<br />
Calappa gallus var. capellonis Laurie, 1906.<br />
DISTRIBUTION.—Tropical Atlantic and Indo-<br />
<strong>West</strong> Pacific. In the eastern Atlantic it has been<br />
recorded from scattered localities between the<br />
Cape Verde Islands and Senegal to Angola, in-<br />
cluding the offshore islands of the Gulf of Guinea;<br />
from shore to a depth of about 50 m.<br />
Calappa granulata (Linnaeus, 1758)<br />
Calappa granulata.—Gruvel, 1913:168 [listed].—Chapman<br />
and Santler, 1955:374.—Monod, 1956:105 [references<br />
only].—Figueira, 1960:8.—Peres, 1964:28.—Zariquiey Alvarez,<br />
1968:315, figs. 105c, 107a [Spain; references].—<br />
Maurin, 1968a: 18 [Portugal], 43, 107 [Mediterranean],<br />
fig. 4; 1968b:479, 480, 482, 484, fig. 2.—Ribeiro, 1973:4,<br />
fig. 1.—Turkay, 1976a:25 [listed], 36, fig. 17 [Portugal in<br />
part]; 1976b:61 [listed], 62.—Bas, Arias, and Guerra, 1976,<br />
table 3.<br />
Calappa.—Maurin, 1968b, fig. 1.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Morocco: Off Cap Hadid, 31°54'N, 09°<br />
55^, 85 m, muddy sand, 5 m beam trawl, 25 Mar 1976,<br />
Onversaagd Sta 126, 19 (L). Casablanca fishmarket, 31 Mar<br />
1976, 1
52<br />
Geer ex Algiria mihi," showing that De Geer's<br />
specimen(s), originating from Algeria, were examined<br />
by Linnaeus himself. Linnaeus' (1758)<br />
description evidently is entirely or partly based<br />
on this Algerian material. In this description,<br />
namely, Linnaeus described "Rostrum bilobum, obtusum"<br />
while Catesby's figure shows a narrowly<br />
rounded truncate rostrum, which certainly cannot<br />
be called bilobed. As Cancer granulatus is a<br />
composite species, its identity can only be fixed<br />
by a lectotype selection. As this, so far as is known<br />
to us, has so far not been done, we now select as<br />
the lectotype of Cancer granulatus Linnaeus, 1758,<br />
the specimen (or if more than one specimen is<br />
represented in the material, the largest specimen)<br />
from Algeria in the De Geer collection.<br />
Monod (1956:105) noted that "la seule mention<br />
possible de l'espece dans notre dition est celle<br />
d'A. MILNE-EDWARDS et BOUVIER [1900:59-60].<br />
Mais comme ces auteurs considerent rubroguttata<br />
comme probablement synonyme de granulata, rien<br />
ne prouve qu'ils aient bien eu cette derniere<br />
espece sous les yeux."<br />
The confirmation of the early record of this<br />
species from the Cape Verde Islands (A. Milne<br />
Edwards and Bouvier, 1900) by Ribeiro (1973),<br />
the records of Maurin (1968a,b) from Morocco,<br />
Spanish Sahara, and Mauritania, and the present<br />
records from Morocco suggest that this species is<br />
more widely distributed in the Atlantic than previously<br />
believed.<br />
BIOLOGY—Calappa granulata is a sublittoral species<br />
which, according to Bouvier (1940) and Zariquiey<br />
Alvarez (1968), lives in depths between 30<br />
and 150 m. If the recent records in the literature<br />
are accurate, the species occurs in much deeper<br />
water off the northwestern coast of Africa. Tiirkay<br />
(1976a) recorded it from depths of 160-250 m off<br />
Morocco and (1976b) in 13 m and 20-30 m off<br />
Madeira. Peres (1964) found it off Morocco in<br />
260-500 m on compact bathyal mud. Maurin<br />
(1968a,b) recorded it from the following depths<br />
and bottom types off Spanish Sahara: 200-300 m<br />
and 200-350 m, slightly sandy mud with Pennatulacea;<br />
400-700 m, sandy mud; 50-90 m, on<br />
muddy detritus; and 60 m, shelly, muddy sand,<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
as well as on bathyal mud with funiculines and<br />
Isidella in 350-450 m off Morocco.<br />
DISTRIBUTION.—Eastern Atlantic, from the<br />
Mediterranean, the Atlantic coasts of Portugal<br />
and Spain, southward to Cap Blanc, Mauritania,<br />
including the Azores, Madeira, and the Cape<br />
Verde Islands, in depths between 13 and 400-700<br />
m (Zariquiey Alvarez, 1968). Monod (1956) summarized<br />
the literature but reported no material.<br />
In addition, the following species localities have<br />
been recorded.<br />
North Atlantic: Conception Bank, 200 and 237 m<br />
(Maurin, 1968b).<br />
Azores: Faial (Figueira, 1960). Horta Harbor, Faial<br />
(Chapman and Santler, 1955).<br />
Madeira: No specific locality; Funchal harbor, 13 m;<br />
Ponta dos Reis Magos, 20-30 m (Tiirkay, 1976b).<br />
Morocco: Foum Agoiiitir (as Puerto-Cansado), 350-450<br />
m (Maurin, 1968b). 35°05.5'N, 09°18'W, 160-250 m (Turkay,<br />
1976a). 34°39.6'N, 06°54.5'W to 34°33.5'N, 06°56'W,<br />
260-500 m (Peres, 1964).<br />
Spanish Sahara: 23°30'N, 16°06'W to 23°28.5'N,<br />
16°09.5'W, 24-29 m (Bas, Arias and Guerra, 1976). Off<br />
Medano de Aaiun and Cabo Bojador, 200-300 m (Maurin,<br />
1968a,b). Between Cabo Bojador and Morro Garnet, 200-<br />
350 m; off Morro Garnet, 400-700 m; between Cabo Corbeiro<br />
and Cabo Blanco, 50-90 m; and off peninsula of Cabo<br />
Blanco, 60 m (all Maurin, 1968b).<br />
Cape Verde Islands: Boavista, 15°58'N, 22°43'W, 80 m<br />
(Ribeiro, 1973).<br />
* Calappa pelii Herklots, 1851<br />
FIGURE 12a<br />
Calappa pelt.— Capart, 1951:39, figs. 9, 10.—Monod, 1956:<br />
102, figs. 117-121.—Rossignol, 1957:75, 127 [key].—<br />
Longhurst, 1958:87.—Gauld, 1960:69.—Guinot and Ribeiro,<br />
1962:26.—Rossignol, 1962:114.—Crosnier, 1964:34,<br />
35, fig. on pi. B.—Forest and Guinot, 1966:52—Monod,<br />
1967:178 [no material].—Maurin, 1968a:48, 59, 64, fig.<br />
23; 1968b:484, 486, 489, 491, figs. 5, 9.—Le Loeuff and<br />
Intes, 1968:40, table 1, figs. 48, 61, 63; 1969:63, 64.—<br />
Crosnier, 1970:1215 [listed], 1216.<br />
Calappa.—Voss, 1966:19.—Maurin, 1968b, figs. 1, 4[?].<br />
Calappa Pelii.—Holthuis, 1968:29 [listed].<br />
SYNONYM.—Calappa piscatorum Caiman, 1914.<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 68, 70 m, broken shell, 39, 16 juv (L).<br />
Ivory Coast: Sta 42,62-75 m, mud with brown, branched
NUMBER 306<br />
Foraminifera, 2 juv (L). Sta 46, 38-42 m, mud with dense<br />
Jullienella, 3 juv (W). Sta 47, 37 m, bottom with Jullienella,<br />
1 juv (L). Sta 49, 73-77 m, 2 juv (W). Sta 62, 46 m, brown,<br />
branching and foliate Foraminifera, 7 juv (L). Sta. 64, 68 m,<br />
1$, 1 juv (W).<br />
Ghana: Sta 17, 48 m, fine sand and green mud, 1? (L).<br />
Sta 22, 51 m, rough bottom, Id (W). Sta 23, 42 m, foliate<br />
brown to orange bryozoans, 3 juv (L).<br />
Nigeria: Sta 237, 101 m, 16* (W). Sta 241, 59-63 m, mud<br />
and shell, 8 juv (L). Sta 253, 33-40 m, mud, 1 juv (L).<br />
Cameroon: Sta 259, 59 m, mud and broken shell, 2 juv<br />
(W). Sta 260, 46 m, 7 juv (L).<br />
Gerommo Material: Gabon: Sta 212, 200 m, 1
54<br />
The deepest records for this species that we<br />
have found are those of Maurin (1968a,b), who<br />
reported material from depths of 200 m off Spanish<br />
Sahara and 200-350 and 200-400 m off Mauritania.<br />
The species generally is taken in depths of<br />
50 to 150 m.<br />
Ovigerous females have been collected in<br />
March, May, August, October, and December<br />
(Capart, 1951; Guinot and Ribeiro, 1962).<br />
DISTRIBUTION.—Off <strong>West</strong> Africa, from localities<br />
between Spanish Sahara and Angola, in<br />
depths between 8-20 and 400 m, usually between<br />
50 and 150 m. Monod (1956) recorded material<br />
from localities between Port-Etienne, Mauritania,<br />
and Luanda, Angola. Since 1956 this species<br />
has been recorded from the following localities.<br />
<strong>West</strong> Africa: No specific locality (Monod, 1967).<br />
Spanish Sahara: Between Cabo Corbeiro and Cabo<br />
Blanco, 60-80 m (Maurin, 1968a); in 200 m (Maurin,<br />
1968b). Between Cabo Barbas and Cabo Blanco, 50-90 m<br />
(Maurin, 1968b).<br />
Mauritania: N of Bane d'Arguin, 20°20'N to 20°40'N,<br />
90-100 m (Maurin, 1968a). Bane d'Arguin, 40-60 m, 60-70<br />
m, 90-100 m (Maurin, 1968b). Between Cap Timiris and<br />
Tamzak (as Tamxat), 200-350 m (Maurin, 1968a). Off<br />
Tamzak (as Tamxat), 200-400 m (Maurin, 1968b).<br />
Senegal: Saint-Louis, 35-40 m (Maurin, 1968b). Off<br />
Mboro, 35-40 m (Maurin, 1968b). 13°01'N, 17°24'W, 5U<br />
55 m, and 12°55.5'N, 17°33'W, 65-75 m (Forest and Guinot,<br />
1966).<br />
Guinea-Bissau: 10°19'N, 16°34'W, 60-73 m (Forest and<br />
Guinot, 1966).<br />
Sierra Leone: No specific locality, in 12-120 m (Longhurst,<br />
1958).<br />
Ivory Coast: No specific locality (Le Loeuff and Intes,<br />
1969). Off Sassandra, off Fresco, off Grand-Lahou, off Jacqueville,<br />
and off Grand-Bassam, 22-200 m (Le Loeuff and<br />
Intes, 1968). 05°07'N, 04°32'W to O5°O7'N, 04°36'W, 38-<br />
42 m (Voss, 1966).<br />
Ghana: Off Accra, 37 m (Gauld, 1960). 04°36.5'N, 01°<br />
31'W, 50 m (Forest and Guinot, 1966).<br />
Cameroon: No specific locality, in more than 50 m (Crosnier,<br />
1964).<br />
Principe: 01°38'25"N, 07°22'05"E, 31 m (Forest and<br />
Guinot, 1966).<br />
Gabon: 00°25'N, 09°00'E, 73 m (Forest and Guinot,<br />
1966).<br />
Congo: Pointe-Noire, 40-75 m (Rossignol, 1957). W of<br />
Pointe-Noire, 80-100 m (Rossignol, 1962).<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Angola: 16°41'S, 11°21'E, 162 m (see "Remarks"); 16°<br />
37'S, 11°22'E, 122 m; 17°O2'S, 1 l°40'E, 54 m; and 17°06'S,<br />
11°35'E, 90 m (all Crosnier, 1970). Luanda, 110-113 m;<br />
Mussulo Grande, Luanda, 86-90 m; Porto Amboim, 85 m;<br />
Baia da Caota, Benguela, 8-20 m, 13 m, 17 m, 30 m;<br />
Sombreiro, 13 m; between Ponta da Caruita and Sombreiro,<br />
29 m; Baia Farta, 22 m, 100-144 m; Baia de Mocamedes;<br />
Baia dos Tigres, 107 m (all Guinot and Ribeiro, 1962).<br />
*Calappa rubroguttata Herklots, 1851<br />
FIGURES 126, 13<br />
Calappa rubroguttata.—Biiuikofer, 1890:466, 487.—Johnston,<br />
1906: 862.—Postel, 1950:25, 26.—Monod, 1956:106, figs.<br />
122-124.—Rossignol, 1957:76.—Longhurst, 1958:87.—<br />
Buchanan, 1958:23.—Gauld, 1960:69.—Rossignol, 1962:<br />
144.—Guinot and Ribeiro, 1962:27.—Crosnier, 1964:32,<br />
35, fig. on pi. B.—Monod, 1967:178, pi. 16: fig. 1 [no<br />
material].—Forest and Guinot, 1966:53.—Maurin, 1968b:<br />
491, fig. 9.—Le Loeuff and Intes, 1968:40, table 1, figs.<br />
48, 61, 63; 1969:63, 64, 65.—Uschakov, 1970:439, 455<br />
[listed].<br />
?Calappa granulata.—Bouvier, 1911:226 [not C. granulata (Linnaeus,<br />
1767)].<br />
Calappa rubroguttatus.—Capan, 1951:43, figs. 10, 11.—-Rossignol,<br />
1957:127 [key]. [Erroneous spelling.]<br />
Calappa.— Bassindale, 1961:499.<br />
Callappa rubroguttata.—Bott, 1968:170 [erroneous spelling].<br />
SYNONYM.—Calappa bocagei De Brito Capello,<br />
1871.<br />
MATERIAL EXAMINED.—Pilhbury Material: Ivory<br />
Coast: Sta 47, 37 m, bottom with Jullienella, 1? ov (L).<br />
Ghana: Sta 24, 35-37 m, dark red bryozoans, 2 juv (W).<br />
Nigeria: Sta 248, 33 m, 2 juv (W).<br />
Other Material: Liberia: No specific locality, 1879-1882, J.<br />
Biittikofer, 3cJ, 1? (L) (smallest male with huge Chelonibia<br />
patula (Ranzani) on posterior margin of carapace).<br />
Ghana: Butre (04°49'N, 01°55'W), 1841-1851, H. S. Pel,<br />
lectotype, \6 (L); paralectotypes, 8
NUMBER 306<br />
pattern in this species: "Coloration typique: la<br />
carapace beige clair, marquee des bandes et<br />
taches rondes carminees. La partie anterieure a<br />
fond carmin clair. La pince marquee de trois<br />
taches rondes disposees en triangle; le carpe de<br />
deux taches. Les pattes de couleur claire."<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 14 to 104 mm; the carapace width of<br />
the single ovigerous female examined is 95 mm.<br />
REMARKS.—The lectotype of this species, selected<br />
here, is a male, cb 104 mm, from Butre<br />
(= Boutry), east of Dixcove, Ghana, collected<br />
between 1841 and 1851 by H. S. Pel (Crust. D.<br />
772). There are nine paralectotypes, a female, cb<br />
51 mm, and eight males, cb 82 to 99 mm, two of<br />
which are preserved dry (Crust. D. 771).<br />
A sketch of the carapace of a juvenile is shown<br />
in Figure \2b along with that of a young C. pelii<br />
(Figure 12a). We also reproduce here an original<br />
illustration of this species prepared by Herklots<br />
(Figure 13).<br />
BIOLOGY.—This species lives in shallower water<br />
than does C. pelii, from shore to a maximum<br />
depth of about 90 m. It apparently can live on all<br />
level substrates, but may prefer sand. Gauld<br />
(1960:69) reported that off Ghana this species was<br />
"very common on sand from shallow water to 30<br />
m. Catches of up to 250 have been taken in a<br />
single net; large catches have been taken only<br />
beyond 10 m." Crosnier (1964) characterized this<br />
species as a warm water crustacean, living in 0-<br />
30 m off Cameroon; there it replaces C. pelii in<br />
FIGURE 13.—Calappa ruboguttata Herklots (sketch by J. A.<br />
Herklots).<br />
shallow water. Off the Ivory Coast, Le Loeuff and<br />
Intes (1968:40) found this species to be very<br />
abundant all year on all sediments in depths less<br />
than 40 m, usually between 20 and 35 m. In<br />
depths between 30 and 40 m it occurred together<br />
with C. pelii. In 1969 they noted that it appeared<br />
to be indifferent to the nature of the substrate<br />
and that it was eurythermic.<br />
More than 85% of the occurrences reported in<br />
the literature for which depth information is<br />
available are from depths of 40 m or less. There<br />
are about equal numbers of records of the species<br />
in depths below 20 m and between 20 and 40 m.<br />
Ovigerous females have been collected in May,<br />
September, and October (Capart, 1951; Forest<br />
and Guinot, 1966; Pillsbury).<br />
DISTRIBUTION.—Off <strong>West</strong> Africa, between Senegal<br />
and Angola, usually in depths of less than 50<br />
m. Monod (1956) summarized the literature prior<br />
to 1956 and reported numerous specimens from<br />
localities between Senegal and Luanda, Angola.<br />
In addition this species has been recorded from<br />
the following localities.<br />
<strong>West</strong> Africa: No specific locality (Monod, 1967).<br />
Senegal: No specific locality, 30-80 m (Postel, 1950). Off<br />
Saint-Louis, 35-40 m; off Mboro, 35-40 m (Maurin, 1968b).<br />
13°01'N, 17°24'W, 51-55 m (Forest and Guinot, 1966). Baie<br />
de Rufisque (Bouvier, 1911).<br />
Guinea: No specific locality, depths greater than 20 m<br />
(Uschakov, 1970).<br />
Sierra Leone: No specific locality, 10-41 m (Longhurst,<br />
1958).<br />
Liberia: No specific locality (Biittikofer, 1890; Johnston,<br />
1906).<br />
Ivory Coast: No specific locality (Le Loeuff and Intes,<br />
1969). 05°00'N, 05°28.5'W, 27 m; 05°03.5'N, 05°25'W, 20-<br />
25 m; 05°02.5'N, 05°25'W, 21-27 m (Forest and Guinot,<br />
1966). Off Sassandra, off Fresco, off Grand-Lahou, off Jacqueville,<br />
and off Grand-Bassam, 8-40 m (Le Loeuff and<br />
Intes, 1968).<br />
Ghana: Off Accra (Bassindale, 1961). In 3-8 fm (5-15<br />
m) (Buchanan, 1958). In shallow water to 30 m (Gauld,<br />
1960).<br />
Cameroon: No specific locality, 0-30 m (Crosnier, 1964).<br />
Principe: Between Ponta da Mina and Ponta Novo Destino,<br />
6 m (Forest and Guinot, 1966).<br />
Gabon: 00°38'25"N, 08°46'E, 5 m (Forest and Guinot,<br />
1966).<br />
55
56<br />
Congo: Baie de Pointe-Noire, beach (Rossignol, 1957).<br />
Pointe-Noire, beach (Rossignol, 1962).<br />
Angola: Luanda (Bott, 1968). Bai'a de Benguela, shore<br />
(Guinot and Ribeiro, 1962).<br />
Genus Cycloes de Haan, 1837<br />
Cydoes de Haan, 1837:67, 68, 69 [type-species: Cycloes granulosa<br />
de Haan, 1837, by monotypy; gender: feminine].<br />
Cryptosoma Brulle, 1837:16 [an invalid junior homonym of<br />
Cryptosoma Bert hold, 1827 (Coleoptera); type-species: Cryptosoma<br />
cristatwn Brulle, 1837, by monotypy; gender: neuter].<br />
REMARKS.—Chace (1968:610) discussed the use<br />
of the two names previously applied to this genus<br />
and gave convincing reasons for using Cycloes<br />
rather than Cryptosoma. He also provided a key to<br />
the species of Cycloes.<br />
Cycloes cristata (Brulle, 1837)<br />
Cryptosoma cristatum.—Monod, 1956:114, fig. 133 [Cape<br />
Verde Islands; references].—Tiirkay, 1976b:61 [listed].<br />
Cycloes cristata.—Guinot-Dumortier and Dumortier, 1961:<br />
561, figs. 1-4 [Cape Verde Islands].—Guinot and Ribeiro,<br />
1962:27, figs. 3, 4 [Cape Verde Islands].<br />
Cycloes cristata.—Chace, 1968:610 [key].—Guinot, 1968b, fig.<br />
13 [morphology].<br />
Cycloes cristata.—Ribeiro, 1964:4 [Cape Verde Islands].<br />
Crytosoma cristatwn.—Tiirkay, 1976b:62 [Madeira, Porto<br />
Santo, Ilhas Desertas; erroneous spelling].<br />
SYNONYM.—Cryptosoma dentatum Brulle, 1839.<br />
DISTRIBUTION.—Eastern Atlantic, from Madeira,<br />
the Canary Islands, and the Cape Verde<br />
Islands; sublittoral (Monod, 1956).<br />
Subfamily MATUTINAE de Haan, 1835<br />
Genus Matuta Weber, 1795<br />
Matuta Weber, 1795:92 [type-species: Cancer victor Fabricius,<br />
1786, a subjective junior synonym of Cancer lunaris Forskal,<br />
1775, by subsequent designation by Latreille, 1810:422;<br />
gender: feminine].<br />
Matutinus MacLeay, 1838:70 [type-species: Cancer victor Fabricius,<br />
1786, a subjective junior synonym of Cancer lunaris<br />
Forskal, 1775, by monotypy; gender: masculine].<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Matuta michaelseni Balss, 1921<br />
Matuta michaelseni.—Capart, 1951:45, fig. 12.—Monod, 1956:<br />
98, figs. 108-114.—Rossignol, 1957:77, 127 [key].—Buchanan,<br />
1958:20.—Longhurst, 1958:87.—Gauld, 1960:<br />
68.—Guinot and Ribeiro, 1962:25.—Rossignol, 1962:<br />
114.—Forest and Guinot, 1966:51.—Le Loeuff and Intes,<br />
1968, table 1.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Liberia: Off St. Paul River, Monrovia,<br />
trawl, 6 Jan 1953, G. C. Miller, \6 (W).<br />
Ivory Coast: Off Sassandra, 11 m, 3 Apr 1969, Guinean<br />
Trawling Survey, Tr 22, Sta 1, 19 (L).<br />
Ghana: Takoradi, 14 Aug 1961, Amegah, 16 (W).<br />
Congo: Beach at Pointe-Noire, beach seine, Jul 1963, A.<br />
Crosnier, 26, 79 (W).<br />
DESCRIPTION.—Capart, 1951:46.<br />
Figures: Monod, 1956, figs. 108-114.<br />
Male Pleopod: Monod, 1956, figs. 110-114 (Senegal,<br />
Togo).<br />
Color: "Blanc-rose uniforme" (Rossignol, 1957:<br />
77).<br />
MEASUREMENTS.—Our specimens have carapace<br />
lengths of 9 to 14 mm.<br />
BIOLOGY.—This species inhabits primarily<br />
sandy bottoms in shallow water, from the shore<br />
to a depth of about 30 m. Buchanan (1958) found<br />
it in the inshore fine sand community, in 3-8 fm<br />
(5-15 m) off Accra, Ghana, and Longhurst<br />
(1958) found it on sand and muddy sand, but<br />
noted that it occurred mostly on shelly sand in 5-<br />
30 m off Sierra Leone. Guinot and Ribeiro (1962)<br />
reported material collected on a beach at low<br />
tide.<br />
Ovigerous females have been collected in January,<br />
March, April, and September (Capart,<br />
1951; Monod, 1956; Guinot and Ribeiro, 1962).<br />
DISTRIBUTION.—<strong>West</strong> Africa, from Senegal to<br />
Angola, in shallow water, shore to about 30 m.<br />
Monod (1956) summarized earlier records and<br />
reported material from localities between Senegal<br />
and Togo. Since 1956 the species has been recorded<br />
from:<br />
Sierra Leone: No specific locality, 5-30 m (Longhurst,<br />
1958).<br />
Ivory Coast: Off Sassandra, 15 m (Le Loeuff and Intes,<br />
1968). 05°03'N, 05°25'W, 20-25 m (Forest and Guinot,<br />
1966).
NUMBER 306 57<br />
Ghana: No specific locality, shore to 15 m (Gauld, 1960).<br />
Accra, 3-8 fm (5-15 m) (Buchanan, 1958).<br />
Congo: Pointe-Noire, beach (Rossignol, 1957). Baie de<br />
Pointe-Noire, beach (Rossignol, 1962).<br />
Angola: Bala de Benguela, beach at low tide (Guinot and<br />
Ribeiro, 1962).<br />
Family LEUCOSIIDAE Samouelle, 1819<br />
LEUCOSIADAE Samouelle, 1819:91 [corrected to Leucosiidae<br />
by Miers, 1879a:671; name 374 on Official List}.<br />
ILIINAE Stimpson, 1871a: 155 [name 372 on Official List].<br />
EBALIINAE Stimpson, 1871a: 159.<br />
MYRODINAE Miers, 1886:297.<br />
OREOPHORINAE Miers, 1886:297.<br />
MYROIDA Alcock, 1896:167.<br />
IPHICULOIDA Alcock, 1896:167.<br />
NURSILIOIDA Alcock, 1896:167.<br />
NURSIOIDA Alcock, 1896:166.<br />
NUCIOIDA Alcock, 1896:167.<br />
CRYPTOCNEMIDAE Stimpson, 1907:161.<br />
PHILYRINAE Rathbun, 1937:151.<br />
EASTERN ATLANTIC GENERA.—Nine, of which<br />
six, Atlantotlos, Ebalia, Ilia, Merocryptus, Philyra, and<br />
Pseudomyra, are represented by tropical species<br />
occurring off <strong>West</strong> Africa. Indo-<strong>West</strong> Pacific species<br />
belonging to three genera have become established<br />
in the eastern Mediterranean, having colonized<br />
that area via the Suez Canal. These genera<br />
are as follows:<br />
Ixa Leach (1815a:334). Type-species: Cancer cylindricus<br />
Fabricius, 1777, by monotypy; gender:<br />
feminine; name 161 on Official List.<br />
Leucosia Weber (1795:92). Type-species: Cancer<br />
craniolaris Linnaeus, 1758, by subsequent designation<br />
by Holthuis, 1959a: 106; gender: feminine;<br />
name 1631 on Official List.<br />
Myra Leach (1817:19, 23). Type-species: Leucosia<br />
fugax Fabricius, 1798, by monotypy; gender:<br />
feminine; name 1635 on Official List.<br />
EASTERN ATLANTIC SPECIES.—Twenty, of which<br />
12 occur in tropical waters. The extralimital species<br />
are as follows.<br />
Ebalia deshayesi Lucas, 1846. Canary Islands,<br />
Madeira (Tiirkay, 1976b), and Mediterranean;<br />
sublittoral, to 100 m (Zariquiey Alvarez, 1968).<br />
Ebalia edwardsii Costa, 1838. Mediterranean;<br />
littoral and sublittoral, to about 200 m (Zariquiey<br />
Alvarez, 1968).<br />
Ebalia granulosa H. Milne Edwards, 1837. Eastern<br />
Atlantic, from England to the Mediterranean;<br />
littoral and sublittoral, to 445 m (Zariquiey Alvarez,<br />
1968).<br />
Ebalia tumefacta (Montagu, 1808). Norway and<br />
Shetland Islands to Spanish Sahara; sublittoral,<br />
to 130 m (Zariquiey Alvarez, 1968; Christiansen,<br />
1969).<br />
Ixa monodi Holthuis and Gottlieb, 1956. An<br />
Indo-<strong>West</strong> Pacific immigrant into the eastern<br />
Mediterranean, Turkey; Red Sea (Holthuis and<br />
Gottlieb, 1956, 1958).<br />
Leucosia signata Paulson, 1875. An Indo-<strong>West</strong><br />
Pacific immigrant into the eastern Mediterranean;<br />
Israel and Egypt (Holthuis and Gottlieb,<br />
1958; Lewinsohn and Holthuis, 1964; Ramadan<br />
and Dowidar, 1976).<br />
Merocryptus boletifer A. Milne Edwards and Bouvier,<br />
1894. Azores, Seine Seamount, Mediterranean;<br />
sublittoral, 100-600 m (Zariquiey Alvarez,<br />
1968).<br />
Myra fugax (Fabricius, 1798). An Indo-<strong>West</strong><br />
Pacific immigrant into the eastern Mediterranean;<br />
Egypt, Israel, and possibly Turkey (Holthuis<br />
and Gottlieb, 1958; Ramadan and Dowidar,<br />
1976).<br />
The status of Ebalia fragifera Miers (1881a:268)<br />
from the Canary Islands, which Monod (1956:<br />
131) considered to be distinct from E. tuberculata,<br />
remains to be determined.<br />
The names used by Monod (1956) have not<br />
been changed, so the species therein are not listed<br />
here.<br />
REMARKS.—Glaessner's (1969:R496) observation<br />
that "this family is commonly divided into<br />
subfamilies which are constituted and defined<br />
differently by different authors (Miers, Alcock,<br />
Ihle, Rathbun, Balss) and are not considered<br />
helpful to paleontologists at the present stage of<br />
our knowledge of the family" is still quite pertinent.<br />
Of the four subfamilies currently recognized<br />
within the Leucosiidae, two, Ebaliinae and Cryptocneminae,<br />
do not offer too many difficulties<br />
and most authors agree in the limits of the two.<br />
The confusion that exists concerning the size and
58<br />
composition of the two other subfamilies, however,<br />
is very great.<br />
The root of all the difficulties in the subfamily<br />
classification of the Leucosiidae is to be found in<br />
a nomenclatural mix-up concerning the type genus<br />
of the family, Leucosia. Until 1897 the name<br />
Leucosia Fabricius, 1798, was used for a genus of<br />
Indo-<strong>West</strong> Pacific crabs, the type of which is<br />
Cancer craniolaris Linnaeus, 1758. In 1897(b), however,<br />
Rathbun showed this usage to be incorrect<br />
as the first type selection for Leucosia Fabricius,<br />
1798, is that by Latreille (1810:97, 422), who<br />
selected Cancer nucleus Linnaeus, 1758, as the type<br />
of the genus. As Cancer nucleus Linnaeus is also the<br />
type-species of the genus Ilia Leach, 1817, the<br />
name Leucosia Fabricius, 1798, had to be substituted<br />
for Ilia Leach, a name until then generally<br />
accepted by carcinologists. For the genus Leucosia<br />
auctt. (non Fabricius, 1798), Rathbun (1897b:<br />
160) proposed the new name Leucosides. There<br />
was considerable opposition to Rathbun's action,<br />
although this action was nomenclaturally entirely<br />
justified; most non-American authors ignored the<br />
changes proposed. In 1959 Holthuis (1959a: 106)<br />
showed that Leucosia Fabricius, 1798, is a junior<br />
homonym of Leucosia Weber, 1795, and thus is<br />
invalid. For Leucosia Weber, 1795, Holthuis selected<br />
as type-species Cancer craniolaris Linnaeus,<br />
1758, thereby making this name a senior synonym<br />
of Leucosides Rathbun, 1897, and Leucosia auctt.<br />
(non Fabricius, 1798). In this way the former,<br />
pre-1897, usage of Leucosia was restored, albeit<br />
with a different author's name. In 1964 the situation<br />
was further consolidated by the International<br />
Commission on Zoological Nomenclature<br />
(1964, in Opinion 712, Bulletin of Zoological Nomenclature,<br />
21(4): 336-351), who placed both<br />
names Leucosia Weber, 1795, and Ilia Leach, 1817,<br />
on the Official List of Generic Names in Zoology.<br />
At the generic level Rathbun's action had<br />
caused hardly any disturbance because (1) non-<br />
American authors had, incorrectly, ignored her<br />
changes, and (2) the genera Ilia Leach, 1817, and<br />
Leucosia Weber, 1795, do not occur in American<br />
waters, the former being restricted to the eastern<br />
Atlantic, the latter to the Indo-<strong>West</strong> Pacific (ex-<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
cept for L. signata which has entered the Mediterranean<br />
via the Suez Canal).<br />
Rathbun's (1937:194) record of "Leucosia planata<br />
(Fabricius)" from Tierra del Fuego rests upon<br />
an uncharacteristic mistake on her part. Fabricius'<br />
species is not a leucosiid but is Halicarcinus<br />
planatus (Fabricius), a species of Hymenosomatidae,<br />
already dealt with by Rathbun (1925:563)<br />
in an earlier work.<br />
At the subfamily level Rathbun's (1897b)<br />
transfer of the name Leucosia from one genus<br />
(Leucosia Weber, 1795) to another (Ilia Leach,<br />
1817) caused much confusion as these two genera<br />
were generally considered to form part of two<br />
different subfamilies. These subfamilies were indicated<br />
by Rathbun (1937:122, 151, and 123,<br />
183) with the names Philyrinae and Leucosiinae,<br />
respectively, while most other authors used the<br />
names Leucosiinae and Iliinae, respectively, for<br />
them. The fact that the name Leucosiinae could<br />
stand for two entirely different groups was not<br />
realized by several authors.<br />
A year before Rathbun (1897b) published her<br />
controversial paper, Alcock (1896) gave an important<br />
classification of the Leucosiidae, which<br />
he divided into two subfamilies: Leucosiinae<br />
(containing 5 subdivisions, named by him Alliances<br />
i-v) and Iliinae (with Alliances I-IV).<br />
Ihle (1918) recognized three subfamilies within<br />
the Leucosiidae: Ebaliinae (with 12 genera, most<br />
of which were placed in Alcock's Alliances I-III of<br />
his Leucosiinae), Iliinae (with 19 genera including<br />
all of those placed by Alcock in his Iliinae, those<br />
of his Alliance iv of Leucosiinae and part of those<br />
of Alliances n and m of the latter subfamily), and<br />
Leucosiinae (with 7 genera including all those of<br />
Alcock's Alliance v of the Leucosiinae and the<br />
genera Cryptocnemus and Carcinaspis which Alcock<br />
had placed (the latter with some doubt) in Alliance<br />
i).<br />
Rathbun (1937) followed Ihle in recognizing<br />
three subfamilies of Leucosiidae, and in his delimitation<br />
of the subfamily Ebaliinae. Rathbun's<br />
Philyrinae contained all the genera that Ihle<br />
assigned to the Leucosiinae plus several that he<br />
considered to belong to the Iliinae (viz., all those
NUMBER 306 59<br />
of Alcock's Leucosiinae Alliance iv, plus Randallia<br />
of Alliance m). Rathbun's Leucosiinae consisted<br />
of the other Iliinae sensu Ihle. As Rathbun dealt<br />
only with the American representatives, it is difficult<br />
to know the exact limits of her subfamilies.<br />
Balss (1957:1612-1615), in his authoritative<br />
treatment of the Decapoda in Bronn's Klassen und<br />
Ordnungen des Tierreichs, which of necessity is<br />
mostly based on study of the literature, evidently<br />
became severely confused. He recognized, like<br />
Rathbun, the two subfamilies Philyrinae and<br />
Leucosiinae, but unlike Rathbun used the generic<br />
names Leucosia and Ilia for the genera that she<br />
named Leucosides and Leucosia, respectively. In the<br />
subfamily Philyrinae Balss placed all the genera<br />
that Ihle had assigned to the Iliinae with the<br />
exception of two, viz., Calhdactylus and Iliacantha,<br />
the only two that Rathbun (1937) mentioned by<br />
name and which she placed in the Leucosiinae.<br />
Balss' Leucosiinae contained all genera placed by<br />
Ihle in that subfamily with the exception of Philyra<br />
(which Rathbun, 1937, had placed in the<br />
Philyrinae) and Pseudophilyra. Balss' definition of<br />
the Philyrinae is based almost exclusively on that<br />
by Rathbun (1937), but his definition of the<br />
Leucosiinae has the first five sentences adapted<br />
from Ihle's (1918:207) account of the Leucosiinae<br />
(sometimes in the identical wording), while the<br />
last sentence (dealing with the third maxilliped)<br />
is taken from Rathbun's (1937:183) definition of<br />
her Leucosiinae. Balss' action was taken in spite<br />
of the fact that Rathbun (1937:151, 183) made<br />
clear that her subfamily Philyrinae corresponds<br />
with all of Ihle's Leucosiinae plus part of his<br />
Iliinae, while her Leucosiinae are formed by the<br />
rest of Ihle's Iliinae. Unfortunately Balss's (1957:<br />
1612-1615) garbled version of the classification<br />
of the Leucosiinae provides the most recent complete<br />
list of all the genera of the family and has<br />
been accepted as the basis for the work of several<br />
subsequent authors (Guinot, 1967a:246-251; Serene,<br />
1968:41-48; Sakai, 1976:76-126). Serene<br />
(1965:11-17) separated the subfamily Cryptocneminae<br />
from the Leucosiinae, placing the genera<br />
Cryptocnemus, Leucisca (= Carcinaspis), Onychomorpha<br />
and Lissomorpha in the former subfamily,<br />
leaving only Leucosia in the latter (probably Callidactylus<br />
and Iliacantha are also left in the Leucosiinae<br />
by Serene, but as these genera do not occur<br />
in the Indo-<strong>West</strong> Pacific region they are not<br />
mentioned by him). For the time being we follow<br />
the system of Ihle (1918), which seems to be more<br />
logical than the one by Rathbun (1937) or than<br />
the garbled version of the latter produced by<br />
Balss (1957). A closer comparative study of the<br />
various genera of Leucosiidae is, however, badly<br />
needed.<br />
Subfamily EBALIINAE Stimpson, 1871<br />
Genus Atlantotlos Doflein, 1904<br />
Atlantotlos Doflein, 1904:49 [type-species: Atlantotlos rhombifer<br />
Doflein, 1904, by monotypy; gender: masculine].<br />
* Atlantotlos rhombifer Doflein, 1904<br />
Atlantotlos rhombifer.—Capart, 1951:57, fig. 16, pi. 2: fig. 1.—<br />
Monod, 1956:134, figs. 162, 163.—Longhurst, 1958:87.—<br />
Gauld, 1960:69.<br />
Atlantotlus rhombifer.—Rossignol, 1962:114 [erroneous spelling]-<br />
Atlantotlos.—Voss, 1966:35.<br />
Merocryptus rhombifer.—Forest and Guinot, 1966:54.<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 68, 70 m, broken shell, 56, 4$ (W).<br />
Ivory Coast: Sta 42, 62-75 m, mud with brown, branched<br />
Foraminifera, let (L). Sta 63, 64 m, sandy mud with shells,<br />
1$ (L).<br />
Nigeria: Sta 239, 73 m, 1$ (L). Sta 241, 59-63 m, mud<br />
and shell, Id, 1$ ov (L). Sta 248, 33 m, 29 (W).<br />
Cameroon: Sta 260, 46 m, 3d, 19, 1 juv (L).<br />
Other Material: Ghana: 05°13'42"N, 03°59'48"W, 50-55<br />
m, dredge, Guinean Trawling Survey, La Rafale, Tr 12, Id<br />
(W).<br />
DESCRIPTION.—Capart, 1951:57.<br />
Figures: Capart, 1951, fig. 16, pi. 2: fig. 1.<br />
Male Pleopod: Capart, 1951, pi. 2: fig. 1 (Gabon);<br />
Monod, 1956, fig. 163 (Senegal).<br />
Color: According to Capart (1951:58), in this<br />
species "Les males et les femelles sont orange rose,<br />
mais les sillons entre les saillies sont bruns avec<br />
des tubercules presque blancs."<br />
MEASUREMENTS.—Our specimens have cara-
60<br />
pace widths of 7 to 16 mm. The carapace width<br />
of the ovigerous female is 15 mm.<br />
REMARKS.—Forest and Guinot (1966) synonymized<br />
Atlantotlos Doflein, 1904 with Merocryptus A.<br />
Milne Edwards, 1873, noting Monod's (1956:<br />
136) comment: "Je ne suis pas certain du tout<br />
que le genre Atlantotlos soit distinct de Merocryptus."<br />
However, the male pleopod of A. rhombifer,<br />
figured in Monod, differs from that of two species<br />
of Merocryptus figured in Guinot and Ribeiro<br />
(1962, fig. 5a, b (M obsoletus), and fig. 6 (M.<br />
boletifer)), in having a subterminal bulbous enlargement<br />
rather than tapering evenly to the<br />
apex. Until all the representatives of these genera<br />
can be studied in more detail, we prefer to retain<br />
Atlantotlos for Doflein's species.<br />
BIOLOGY.—This species occurs in depths between<br />
44 and 115m, but most specimens were<br />
taken in depths between 70 and 100 m. The<br />
Pillsbury specimens were collected on mud with<br />
brown, branched Foraminifera in 62-75 m, sandy<br />
mud with shells in 64 m, and on mud and shell<br />
in 59-63 m. The material reported by Capart<br />
(1951) was taken on brown muddy sand and<br />
coral, brown sandy mud and sandy brown mud,<br />
and on muddy sand or muddy sand and rock in<br />
85 to 100 m. Longhurst (1958) found it on shelly<br />
mud in 106 m, and Rossignol (1962:114) found<br />
it in 115 m, "en bordure des affleurements rocheux."<br />
Forest and Guinot (1966) found the species<br />
on mud, rocks, calcareous algae, sand and<br />
Foraminifera in 51-55 m and on mud and shell<br />
in 73-80 m.<br />
Ovigerous females have been recorded in February,<br />
May, September, and December (Capart,<br />
1951; Pillsbury).<br />
DISTRIBUTION.—<strong>West</strong> Africa, where it has been<br />
recorded from a few localities between Senegal<br />
and Angola, in depths between 44 and 115 m.<br />
Monod (1956) summarized earlier records and<br />
recorded the species from Senegal. Since 1956 it<br />
has been recorded from the following localities.<br />
Senegal: 13°01'N, 17°24'W, 51-55 m (Forest and Guinot,<br />
1966).<br />
Liberia: 05°21.5'N, 09°54.5'W, 73-80 m (Forest and<br />
Guinot, 1966).<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Sierra Leone: No specific locality, 106 m (Longhurst,<br />
1958).<br />
Ghana: Off Accra, 80 m (Gauld, 1960).<br />
Nigeria: 04°56'N, 05°00'E, to 04°54'N, 05°05'E, 73 m<br />
(Voss, 1966).<br />
Congo: W of Pointe-Noire, 115 m (Rossignol, 1962).<br />
Genus Ebalia Leach, 1817<br />
Ebalta Leach, 1817, in 1815-1875, pi. 25 [type-species: Ebalia<br />
bryerii Leach, 1817, a subjective junior synonym of Cancer<br />
tumefactus Montagu, 1808, by subsequent designation by<br />
H. Milne Edwards, 1837, in 1836-1844, pi. 24: fig. 3 (as<br />
Ebalia brayerii); gender: feminine; name 145 on Official<br />
List, type-species there given as Cancer tuberosus Pennant,<br />
1777, in error].<br />
Phlyxia Bell, 1855:303 [type-species: Phlyxia crassipes Bell,<br />
1855, by selection by Bell, 1855:304; gender: feminine].<br />
Bellidilia Kinahan, 1856:115, 117, 128 [type-species: Bellidilia<br />
undecimspinosa Kinahan, 1856, by present designation; gender:<br />
feminine].<br />
* Ebalia affinis Miers, 1881<br />
Ebalia atlantica.—Capart, 1951:54.<br />
Ebalia affinis.—Monod, 1956:117, figs. 134-144.—Longhurst,<br />
1958:87.—Gauld, 1960:69.—Forest and Guinot, 1966:53.<br />
SYNONYM.—Ebalia atlantica A. Milne Edwards<br />
and Bouvier, 1898.<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 68, 70 m, broken shell, 3c? (W).<br />
Ivory Coast: Sta 47, 37 m, bottom with Jullienella, 3c5, 1$<br />
(L).<br />
Ghana: Sta 17, 48 m, fine sand and green mud, Ic5 (W).<br />
Sta 23, 42 m, foliate brown to orange bryozoans, 1
NUMBER 306 61<br />
BIOLOGY.—This species has been recorded from<br />
depths between 4 and 140 m, but more than 85%<br />
of the records are between 4 and 45 m. Capart<br />
(1951) reported it on brown muddy sand in 100<br />
m. Longhurst (1958) found it on muddy sand,<br />
shelly sand, and shelly mud in depths between 8<br />
and 140 m. Forest and Guinot (1966) reported it<br />
from mud or shelly mud in 18-30 m, mud, calcareous<br />
algae, and shell in 31 m, rocks and coral<br />
in 3-10 m, sand, algae, and calcareous algae in<br />
8-30 m, and mud and calcareous algae in 4-5 m.<br />
The Pillsbury specimens were taken on fine sand<br />
and green mud in 48 m, on bottom with bryozoans<br />
in 35-37 and 42 m, on bottom with Jullienella<br />
in 37 m, and on broken shell in 70 m.<br />
Apparently the species prefers soft bottom mixed<br />
with shell or other hard substances.<br />
Ovigerous females have been collected in January,<br />
February, March, April, May, and June<br />
(Monod, 1956; Forest and Guinot, 1966).<br />
DISTRIBUTION.—Off <strong>West</strong> Africa, from localities<br />
between Senegal and Angola; it also has been<br />
reported from the Seine Seamount north of Madeira.<br />
Monod (1956) summarized earlier records<br />
and reported material from Senegal, Guinea,<br />
Sierra Leone, and Ghana. Since 1956 the species<br />
has been reported from the following localities.<br />
Guinea: 09°40'N, 14°05'W, 18 m, andO9°36'N, 13°57'W,<br />
18-30 m (Forest and Guinot, 1966).<br />
Sierra Leone: No specific locality, 8-140 m (Longhurst,<br />
1958).<br />
Ghana: Accra, 32 m (Gauld, 1960).<br />
Principe: O1°37'N, 07°22'E, 30 m, and 01°38'25"N,<br />
07°22'05"E, 31 m (Forest and Guinot, 1966).<br />
Sao Tome: Praia de Santa Catarina, W coast, 3-10 m;<br />
00°25'15"N, 06°43'05"E, 8-30 m; Baia de Ana de Chaves,<br />
5 m; and in front of Ponta Diogo Nunes, 4 m (Forest and<br />
Guinot, 1966).<br />
Ebalia cranchii Leach, 1817<br />
Ebalia cranchi. —Monod, 1956:122, figs. 145, 146 [Senegal;<br />
references].—Zariquiey Alvarez, 1968:329, figs. 108a,b,<br />
lllb,d, lllAa, lllCb [Spain; references].—Tiirkay,<br />
1976a:25 [listed], 37, fig. 19 [Morocco].<br />
Ebalia cranchii.—Christiansen, 1969:31, fig. 11, map 5 [Scandinavia].<br />
DISTRIBUTION.—Eastern Atlantic, from Norway<br />
to Senegal, Mediterranean, in depths between 20<br />
and 550 m (Zariquiey Alvarez, 1968).<br />
Ebalia nux A. Milne Edwards, 1883<br />
Ebalia nux A. Milne Edwards, 1883, pi. 5.—Monod, 1956:<br />
121 [references].—Zariquiey Alvarez, 1968:328, figs. 108c,<br />
111 Ab [Spain; references].—Turkay, 1976a:25 [listed], 37,<br />
fig. 18 [Morocco].<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Morocco: Off Cap de Mazagan, 33°38'N,<br />
08°45'W, 420 m, Agassiz trawl, 28 Mar 1976, Onversaagd Sta<br />
150, 1
62 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
SYNONYM.—Lithadia barnardi Stebbing, 1920.<br />
MATERIAL EXAMINED.—Pillsbuty Material: Ghana:<br />
Sta 22, 51 m, rough bottom, 29 (L). Sta 23, 42 m, foliate<br />
brown to orange bryozoans, Ic5 (L). Sta 24, 35-37 m, dark<br />
red bryozoans, 7
NUMBER 306 63<br />
Burger (1902, 1904) described a gynandromorph<br />
specimen of Jasus frontalis (H. Milne Edwards),<br />
and Hay (1905) mentioned the case of such a<br />
specimen in the freshwater crayfish Orconectes propinquus<br />
(Girard). According to Balss (1944:634,<br />
635), apart from a case in Lucifer, no gynandromorphs<br />
have been reported in Decapoda other<br />
than the Macrura Reptantia. However, a gynandromorph<br />
of the Mediterranean crab, Brachynotus<br />
gemmellari (Rizza), was reported by Froglia<br />
and Manning (1978:700, fig. 5). In Homarus and<br />
Jasus the body of the gynandromorph usually is<br />
skewed because of the different longitudinal proportions<br />
of males and females.<br />
Nothing irregular was found in the shape of<br />
the carapace or the thoracic appendages of this<br />
specimen of E. tuberculata, which in this case may<br />
be an indication that in this species the sexes are<br />
not different in size.<br />
Both Capart (1951:56) and Guinot (1967a:247,<br />
footnote) suggest that material from South Africa<br />
referred to this species by Barnard (1950) probably<br />
belongs to a distinct species. Barnard (1955:<br />
4) proposed Ebalia pondoensis for South <strong>African</strong><br />
specimens previously identified with E. tuberculata.<br />
COMMENSALS AND PARASITES.—The carapace of<br />
the gynandromorph is covered dorsally and ventrally<br />
by Bryozoa, but the appendages, sternum,<br />
and abdomen are entirely free from this growth.<br />
In the space enclosed by the abdomen and the<br />
thoracic sternum a number of sausage-shaped<br />
organisms, resembling Rhizocephala of the genus<br />
Thompsonia, can be seen. One such organism is<br />
fastened to the male half of the thoracic sternite<br />
of the fifth pereiopod, whereas on the female half<br />
of the abdomen seven similar organisms are present,<br />
two attached to a pleopod, the other five to<br />
the abdominal sternites.<br />
BIOLOGY.—The depths at which this species<br />
has been taken range from 10-12 m to 135-250<br />
m. Of the known depth records, more than 85%<br />
are from depths of less than 75 m, the optimal<br />
depth apparently being between 30 and 50 m.<br />
Little is known of the habitat preferences of the<br />
species. The Pillsbury specimens were taken on<br />
rough bottom in 51 m, on bottom with bryozoans<br />
in 35-37 and 42 m, and on bottom with nodular<br />
coralline algae in 51-55 m. The species was reported<br />
from mud, sand, and compacted sand<br />
[sable construit] in 65-75 m and on mud with<br />
Area in 32 m by Forest and Guinot (1966).<br />
Apparently the species spawns all year. Ovigerous<br />
females have been recorded in January,<br />
March, April, May, and September through December<br />
(Monod, 1956; Pillsbury).<br />
DISTRIBUTION.—Off <strong>West</strong> Africa, where it occurs<br />
from localities between Morocco and Angola<br />
and possibly South Africa. Monod (1956), who<br />
recorded material from Morocco, the Cape Verde<br />
Islands, Senegal, Ghana, and Angola, summarized<br />
the earlier literature. Since 1956 this species<br />
has been recorded from the following localities.<br />
Senegal: 12°55.5'N, 17°33'W, 65-75 m (Forest and<br />
Guinot, 1966).<br />
Ghana: Accra, 37-51 m (Gauld, 1960).<br />
Nigeria: Off the mouths of the Niger River, 04°03'N,<br />
06°12'E, 32 m (Forest and Guinot, 1966).<br />
This species has not been recorded previously<br />
from Annobon.<br />
Ebalia tuberosa (Pennant, 1777)<br />
Ebalia tuberosa.—Lebour, 1954:236.—Monod, 1956:124, figs.<br />
150, 151 [England, Algeria, Bonifacio, Ilhas Desertas,<br />
Canary Islands, Mauritania; references].—Zariquiey Alvarez,<br />
1968:326, figs. 109d, HOa-c, Ilia [Spain; references].—Christiansen,<br />
1969:27, fig. 9, map 3 [Scandinavia].—<br />
Tiirkay, 1976a:25 [listed], 37, fig. 20 [Morocco];<br />
1976b:61 [listed], 63 [Madeira, Ilhas Desertas].<br />
SYNONYMS.—Ebalia pennantii Leach, 1817; Ebalia<br />
insignis Lucas, 1849; PEbalia madeirensis Stimpson,<br />
1858.<br />
REMARKS.—This species is included in this list<br />
of tropical species on the basis of the single juvenile<br />
female reported by Monod (1956:1Z6) from<br />
the Bane d'Arguin, Mauritania, 21°51'N,<br />
19° 48'W. Other specimens from the same station<br />
off the Bane d'Arguin were referred to E. tuberculata<br />
by A. Milne Edwards and Bouvier (1900:50).<br />
Its occurrence northward, off Spanish Sahara<br />
and Morocco, is not questioned. Christiansen<br />
(1969) did not accept Monod's record, noting
64<br />
that the species occurred southward to Spanish<br />
Sahara. Lebour (1954:236) reported on some larval<br />
and postlarval specimens of Ebaha from the<br />
Benguela Current that she believed might be<br />
identified with this species.<br />
DISTRIBUTION.—Eastern Atlantic, from Norway<br />
and the Hebrides southward to Mauritania (?),<br />
including the Azores and the Canary Islands,<br />
Mediterranean; sublittoral to about 138 m (Zariquiey<br />
Alvarez, 1968; Christiansen, 1969).<br />
Genus Merocryptus A. Milne Edwards, 1873<br />
Merocryptus A. Milne Edwards, 1873a: 78 [type-species: Merocryptus<br />
lambriformis A. Milne Edwards, 1873, by monotypy;<br />
gender: masculine; name 166 on Official List].<br />
Merocryptus obsoletus A. Milne Edwards and<br />
Bouvier, 1898<br />
Merocryptus obsoletus.— Monod, 1956:132, figs. 161, 161 bis<br />
[Morocco; Senegal].—Guinot and Ribeiro, 1962:28, figs.<br />
5a, b, pi. 1: figs. 1, 2, 4 [Cape Verde Islands, Senegal,<br />
Angola].<br />
DISTRIBUTION.—Eastern Atlantic, from Morocco,<br />
the Cape Verde Islands, Senegal, and Angola,<br />
in depths between 75 and 132 m.<br />
Subfamily ILIINAE Stimpson, 1871<br />
Genus Ilia Leach, 1817<br />
Leucosia Fabricius, 1798:313, 349 [an invalid junior homonym<br />
of Leucosia Weber, 1795 (Decapoda); type-species:<br />
Cancer nucleus Linnaeus, 1758, by subsequent designation<br />
by Latreille, 1810:97, 422; gender: feminine; name 1732<br />
on Official Index].<br />
Ilia Leach, 1817:19, 24 [type-species: Cancer nucleus Linnaeus,<br />
1758, by monotypy; gender: feminine; name 1628 on<br />
Official List].<br />
Thaumasta Gistel, 1848:ix [substitute name for Leucosia Fabricius,<br />
1798; type-species: Cancer nucleus Linnaeus, 1758;<br />
gender: feminine].<br />
Ilia nucleus (Linnaeus, 1758)<br />
Ilia nucleus.—Monod, 1956:139 [references].—Guinot and<br />
Ribeiro, 1962:30 [Cape Verde Islands].—Zariquiey Alvarez,<br />
1968:322, figs. 1 lc, 94f [Spain; references].<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
PIlia nucleus spinosa.— Tiirkay, 1975a:71 [listed], 72 [Spanish<br />
Sahara]. [Not Ilia spinosa Miers, 1881.]<br />
SYNONYMS.—Cancer orbicularis Olivi, 1792; Leucosia<br />
leachii Risso, 1822; Ilia laevigala Risso, 1827;<br />
Ilia rugulosa Risso, 1827; Ilia parvicauda Costa,<br />
1853.<br />
REMARKS.—Tiirkay's (1975a) record of Ilia<br />
from off Cabo Blanco, Spanish Sahara, may be<br />
referable to this species rather than to /. spinosa.<br />
Monod (1933b) recorded /. nucleus from the same<br />
locality, and this apparently is the southernmost<br />
limit of the species along the <strong>African</strong> mainland.<br />
Ilia spinosa is not known with certainty to occur<br />
north of Mauritania.<br />
DISTRIBUTION.—Eastern Atlantic, from the<br />
Cape Verde Islands (where it occurs together with<br />
/. spinosa), Spanish Sahara, and from the Mediterranean,<br />
in depths between 4 and 162 m (Zariquiey<br />
Alvarez, 1968).<br />
* Ilia spinosa Miers, 1881<br />
PIlia nucleus.—Bouvier, 1911:226 [not Ilia nucleus (Linnaeus,<br />
1758)].<br />
Leucosia spinosa.—Capart, 1951:52, fig. 15.<br />
Ilia spinosa.—Monod, 1956:136, 632, figs. 164-166.—Buchanan,<br />
1958:28, 54.—Longhurst, 1958:87.—Lebour,<br />
1959:133, 135, 136, 137, fig. 21 [larvae].—Gauld, 1960:<br />
69.—Rossignol, 1962:114.—Guinot and Ribeiro, 1962:30,<br />
pi. 4: fig. 1.—Crosnier, 1964:38.—Ribeiro, 1964:4.—Forest<br />
and Guinot, 1966:55.—Crosnier, 1967:323.—LeLoeuff<br />
andlntes, 1968:31, table 1.<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 68, 70 m, broken shell, 1 juv (L).<br />
Ivory Coast: Sta 42, 62-75 m, mud with brown, branched<br />
Foraminifera, 1(5, 1 juv (W). Sta 46, 38-42 m, mud bottom<br />
with dense Julltenella, 26, 19 (W). Sta 48, 22 m, 1(5 (W). Sta<br />
62, 46 m, brown, branching and foliate Foraminifera, 3
NUMBER 306 65<br />
mud, stones, calcareous algae, sand, Foraminifera, 15 May<br />
1956, Calypso Sta 3, 1$ ov (W).<br />
Cameroon: 02°39'N, 09°40'E, 50-65 m, mud, 22 Aug<br />
1963, A. Crosnier, 29 (1 ov) (W).<br />
DESCRIPTION.—Capart, 1951:52.<br />
Figures: Monod, 1956, figs. 164-166.<br />
Male Pleopod: Monod, 1956, figs. 165, 166 (Senegal).<br />
Color: Capart (1951:53) noted that in this species<br />
"Le carapace est orange clair, les epines et<br />
appendices presque blancs."<br />
MEASUREMENTS.—Our specimens have carapace<br />
lengths of 5 to 17 mm; the carapace lengths<br />
of ovigerous females are 10 to 16 mm.<br />
REMARKS.—In none of our specimens are the<br />
posterolateral spines of the carapace so well developed<br />
as shown for a specimen from Angola by<br />
Guinot and Ribeiro (1962, pi. 4: fig. 1).<br />
It seems likely that Tiirkay's (1975a) record of<br />
Ilia nucleus spinosa from Spanish Sahara is based<br />
on material of/, nucleus (Linnaeus) rather than /.<br />
spinosa. For that reason we have placed it in the<br />
synonymy of the latter species.<br />
Guinot and Ribeiro (1962) noted that one of<br />
their specimens from the Cape Verde Islands was<br />
similar to /. nucleus in having relatively short<br />
posterolateral spines on the carapace. Apparently<br />
both species occur in the Cape Verde Islands.<br />
Bouvier (1911:226) reported /. nucleus from Baie<br />
de l'Ouest, Mauritania, and noted "avec passage<br />
a la spinosa, Miers."<br />
BIOLOGY.—This species lives in relatively shallow<br />
water, in depths between 6 and 132 m. Of 49<br />
depth records in Monod (1956), 1 was at 132 m,<br />
1 at 96 m, 1 at 75 m, and 46 were at 50 m or less.<br />
Apparently, like many of the leucosiids, it prefers<br />
soft bottoms with harder (larger?) substances in<br />
it. Sourie (1954b) found it on coarse shelly sand,<br />
bottom with Area and Pyura, in 10-12 m in the<br />
Baie de Dakar, and Buchanan (1958) took it in<br />
the silty sand community with Jullienella off<br />
Ghana. Crosnier (1964) reported that it was<br />
found on mud or sandy mud in cold water off<br />
Cameroon. Forest and Guinot (1966) reported it<br />
from a variety of substrates in depths between 18<br />
and 90-100 m. Most of the Pillsbury specimens<br />
were taken on bottom with shell, bryozoans, or<br />
Foraminifera.<br />
Ovigerous females have been collected in May,<br />
June, July, August, October, November, and December<br />
(Capart, 1951; Monod, 1956; Forest and<br />
Guinot, 1966; Crosnier, 1967; Pillsbury).<br />
DISTRIBUTION.—Off <strong>West</strong> Africa, from the Canary<br />
Islands, the Cape Verde Islands, and Mauritania<br />
southward to Angola, in depths between 4<br />
and 132 m, generally in less than 50 m. Monod<br />
(1956) reported material from Mauritania, Senegal,<br />
Guinea, Sierra Leone, Ghana, and Principe.<br />
In addition, the species has been recorded from<br />
the following localities.<br />
Cape Verde Islands: Bafa de Porto Grande, Sao Vicente,<br />
8 m (Guinot and Ribeiro, 1962; Ribeiro, 1964).<br />
Mauritania: Baie de l'Ouest (Bouvier, 1911).<br />
Senegal: 13°01'N, 17°24'W, 51-55 m, and 12°55.5'N,<br />
17°33'W, 65-75 m (Forest and Guinot, 1966).<br />
Guinea-Bissau: 10°19'N, 16°34'W, 18 m (Forest and<br />
Guinot, 1966).<br />
Guinea: 09°36'N, 13°57'W, 18-30 m (Forest and Guinot,<br />
1966).<br />
Sierra Leone: No specific locality, in 8-32 m (Longhurst,<br />
1958).<br />
Ivory Coast: Off Jacqueville and Grand-Bassam, 40-<br />
100 m (Le Loeuff and Intes, 1968).<br />
Ghana: 04°40'N, 02°08'W to 04°39'N, 02°05'W, 50 m;<br />
04°36.5'N, 01°31'W, 50 m; 04°37'N, 00°50'W, 90-100 m<br />
(all Forest and Guinot, 1966). Off Accra (Buchanan, 1958);<br />
in 20-55 m (Gauld, 1960).<br />
Cameroon: No specific locality (Crosnier, 1964).<br />
Principe: 01°38'25"N, 07°22'05"E, 31 m (Forest and<br />
Guinot, 1966).<br />
Annobon: 01°25'30"S, 05°39'E, 52 m (Crosnier, 1967).<br />
Gabon: 00°40'S, 08°46'25"E, 18 m, and 00°38'20"S,<br />
08°48'30"E, 35 m (Forest and Guinot, 1966).—W of Mayumba,<br />
20 m (Rossignol, 1962).<br />
Congo: W of Pointe-Noire, 20-30 m (Rossignol, 1962).<br />
Angola: Baia Farta, Benguela, 40 m (Guinot and Ribeiro,<br />
1962).<br />
Lebour (1959) recorded larvae of this species<br />
from the following localities: Guinea, 10°22'N,<br />
16°34'W; Sierra Leone, 08°22'N, 14°08'W;<br />
Ghana, 05°44'N, 01°02'E; Cameroon, 04°01'N,<br />
07°23'E; and Angola, 07°35'S, 12°46'E.<br />
Subfamily LEUCOSIINAE Samouelle, 1819<br />
Genus Philyra Leach, 1817<br />
Philyra Leach, 1817:18, 22 [type-species: Leucosia globulosus
66<br />
Bosc, 1802, a subjective junior synonym of Cancer globus<br />
Fabricius, 1775, by subsequent designation by H. Milne<br />
Edwards, 1837, in 1836-1844, pi. 24: fig. 4 (as Philyra<br />
globulosa); gender: feminine; name 1642 on Official List.]<br />
Philyra cristata Miers, 1881<br />
Philyra cristata.—Monod, 1956:144, figs. 177-183 [Senegal,<br />
Guinea, Sierra Leone; references].—Longhurst, 1958:87<br />
[Sierra Leone].—Rossignol, 1962:115 [Congo].—Forest<br />
and Guinot, 1966:56 [Guinea; Sao Tome].<br />
DISTRIBUTION.—<strong>West</strong> Africa, from Senegal to<br />
the Congo, in depths between 4 and 25 m.<br />
* Philyra lacvidorsalis Miers, 1881<br />
Philyra laevidorsalis.—Capart, 1951:47, fig. 13.—Monod,<br />
1956:141, figs. 169-176.—Rossignol, 1957:77.—Longhurst,<br />
1958:87.—Buchanan, 1958:20.—Gauld, 1960:<br />
69.—Rossignol, 1962:115.—Forest and Guinot, 1966:56.<br />
MATERIAL EXAMINED.—Pillsbury Material: Ivory<br />
Coast: Sta 47, 37 m, bottom with Jullienella, \6 (W).<br />
Other Material: Liberia: Off St. Paul River, Monrovia,<br />
22-29 m, trawl, 4 Nov 1953, G. C. Miller, \6 (W).<br />
Ivory Coast: Off Sassandra, 11 m, 3 Apr 1964, Guinean<br />
Trawling Survey, Tr 22, Sta 1, 46\ 69 (L).<br />
DESCRIPTION.—Capart, 1951:47.<br />
Figures. Monod, 1956, figs. 169-176.<br />
Male Pleopod: Monod, 1956, figs. 173-176 (Senegal).<br />
Color: Rossignol (1957) noted that this species<br />
is a uniform gray beige, with the ventral surface<br />
lighter. Capart (1951:47) reported that this species<br />
has "couleur uniforme gris brunatre, plus<br />
claire sur la face inferieure; telson blanc ivoire."<br />
MEASUREMENTS.—Our specimens have carapace<br />
lengths of 6 to 11 mm.<br />
BIOLOGY.—Philyra laevidorsalis is a coastal species,<br />
generally found in shallow water, in depths<br />
between 4 and 18-30 m. Longhurst (1958) found<br />
it in 9-86 m off Sierra Leone, but all other depth<br />
records in the literature are from 20-25 m or less.<br />
Sourie (1954b) found it on coarse shelly sand,<br />
bottom with Area and Pyura, in the Baie de Dakar,<br />
and Buchanan (1958) characterized it as a member<br />
of the active epifauna, inshore fine sand<br />
community, in 3-8 fm (5-15 m) off Ghana. Off<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Sierra Leone, Longhurst (1958) found it on shelly<br />
sand. Forest and Guinot (1966) reported it from<br />
mud in 18-30 m, shells in 20-25 m, calcareous<br />
algae in 10-12 m, and calcareous algae, sand and<br />
mud or shells in 6 to 11 m.<br />
Ovigerous females have been collected in April<br />
and May (Monod, 1956; Forest and Guinot,<br />
1966).<br />
DISTRIBUTION.—<strong>West</strong> coast of Africa, from Cap<br />
Blanc, Mauritania, southward to Angola, including<br />
the Cape Verde Islands and Principe, in<br />
shallow water, from a depth of 4 m to about 30<br />
m. Monod (1956), who summarized the earlier<br />
literature, reported material from Senegal and<br />
Ghana. Since 1956 it has been recorded from the<br />
following localities.<br />
Guinea: 09°36'N, 13°57'W, 18-30 m (Forest and Guinot,<br />
1966).<br />
Sierra Leone: No specific locality, 9-86 m (Longhurst,<br />
1958).<br />
Ivory Coast: 05°03'N, 05°25'W, 20-25 m (Forest and<br />
Guinot, 1966).<br />
Ghana: Accra, 3-8 fm (5-15 m) (Buchanan, 1958). Off<br />
Accra and Takoradi, 15-20 m (Gauld, 1960).<br />
Principe: Between Ponta da Mina and Ilheu Santana,<br />
10-12 m; in front of (Cais de) Santana, 11 m; and between<br />
Ponta da Mina and Ponta Novo Destino, 6 m (all Forest<br />
and Guinot, 1966).<br />
Congo: Pointe-Noire, beach (Rossignol, 1957). Baie de<br />
Pointe-Noire, 7-15 m (Rossignol, 1962).<br />
Genus Pseudomyra Capart, 1951<br />
Pseudomyra Capart, 1951:48 [type-species: Pseudomyra mbizi<br />
Capart, 1951, by original designation and monotypy;<br />
gender: feminine].<br />
* Pseudomyra mbizi Capart, 1951<br />
Pseudomyra mbizi Capart, 1951:49, fig. 14, pi. 2: fig. 24.—<br />
Monod, 1956:140, figs. 167, 168.—Rossignol, 1962:115.—<br />
Guinot and Ribeiro, 1962:30.—Crosnier, 1964:35.—Forest<br />
and Guinot, 1966:56.—Voss, 1966:35.—Le Loeuff and<br />
Intes, 1968:40.—Maurin, 1968b:491, 492.—Crosnier,<br />
1970:1215, 1216.<br />
Pseudomyra.—Voss, 1966:33, 36.<br />
Pseudomyra m'bizi.—Le Loeuff and Intes, 1968, table 1 [erroneous<br />
spelling]; 1969:66.<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:
NUMBER 306 67<br />
Sta 68, 70 m, broken shell, 196", 32$ (23 ov), 7 juv (W). Sta<br />
69, 29 m, coral or rock, lcj (W).<br />
Ivory Coast: Sta 42, 62-75 m, mud with brown, branched<br />
Foraminifera, 174c5, 9, juv (L,W). Sta 45, 73-97 m, 96\ 3$<br />
(W). Sta 49, 73-77 m, lie?, 39, 7juv (L). Sta 50, 128-192 m,<br />
16, 29 (W). Sta 59, 55-64 m, mud with dense, branched<br />
Foraminifera, 5ct, 49 (W). Sta 60, 79-82 m, coral or rock, 3
68<br />
the eastern Atlantic. Guinot (1976) provided a<br />
recent review of the family.<br />
Family ATELECYCLIDAE Ortmann, 1893<br />
ATELECYCLIDAE Ortmann, 1893a:27 [name 369 on Official<br />
List].<br />
CHEIRAGONIDAE Ortmann, 1893b:413, 419.<br />
EASTERN ATLANTIC GENERA.—One, Atelecyclus,<br />
represented in the tropical fauna of <strong>West</strong> Africa.<br />
EASTERN ATLANTIC SPECIES.—Two, both occurring<br />
in Europe as well as off tropical <strong>West</strong> Africa.<br />
Monod (1956) included accounts of both species,<br />
as follows, but had material of only one:<br />
Name in Monod Current Name<br />
Atelecyclus septemdentatus Atelecyclus rotundatus<br />
Atelecyclus undecimdentatus Atelecyclus undecimdentatus<br />
REMARKS.—Neither species was taken by the<br />
Pillsbury.<br />
Genus Atelecyclus Leach, 1814<br />
Atelecyclus Leach, 1814:430 [type-species: Cancer {Hippa) septemdentatus<br />
Montagu, 1813, a subjective junior synonym of<br />
Cancer rotundatus Olivi, 1792, by monotypy; gender: masculine;<br />
name 1608 on Official List].<br />
Atelecyclus rotundatus (Olivi, 1792)<br />
Cancer rotundatus Olivi, 1792, pi. 2: fig. 2.<br />
Atelecyclus rotundatus.— Sourie, 1954b: 150.—Forest, 1958:472,<br />
fig. 2 [references].—Peres, 1964:20.—Forest and Guinot,<br />
1966:57.—Maurin, 1968a:30.—Zariquiey Alvarez, 1968:<br />
342, figs. Id, 112b [Spain; references].—Christiansen,<br />
1969:37, fig. 13, map 7 [Scandinavia].—Turkay, 1976a:25<br />
[listed], 37, fig. 21 [Portugal].<br />
Atelecyclus septemdentatus.—Bouvier, 1911:236.—Monod,<br />
1956:148.—Longhurst, 1958:87.<br />
Atelecyclus.— Maurin, 1968a, fig. 13.<br />
SYNONYMS.—Cancer septemdentatus Montagu,<br />
1813; A telecyclus heterodon Leach, 1815.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Gtrommo Material: Gabon: Sta 211, 100 m, 1$ (W).<br />
DESCRIPTION.—Christiansen, 1969:37.<br />
Figure: Christiansen, 1969, fig. 13.<br />
MEASUREMENTS.—Our specimen, a non-oviger-<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
ous female, has a carapace length and a carapace<br />
width of 15 mm.<br />
REMARKS.—Our specimen agrees well with the<br />
accounts of this species given by Christiansen<br />
(1969) and Forest (1958). The latter author established<br />
the identity of A. rotundatus (Olivi, 1792)<br />
with A. septemdentatus (Montagu, 1813).<br />
A comparison of our young specimen with<br />
material of this species from the Mediterranean<br />
shows that the <strong>West</strong> <strong>African</strong> specimen has a<br />
considerably rougher carapace; no other differences<br />
were observed.<br />
BIOLOGY.—Little is known about the habitat<br />
requirements of this species, which, as Forest<br />
(1958) noted, lives from shallow water to a depth<br />
of 200-300 m; Forest questioned the validity of<br />
some early records from very deep water. Off<br />
Scandinavia, Christiansen (1969) found it on<br />
sand and soft bottom, often with gravel and small<br />
stones, in depths between 15-40 and 190-324 m.<br />
Longhurst (1958) collected it on shelly sand in<br />
121-160 m. Peres (1964) recorded it from reddish<br />
gravel with shell debris in 210 m off the Bane de<br />
Spartel, and Maurin (1968a) reported it from<br />
muddy sand and sandy detritus in 10-30 m. It<br />
was collected on mud, rocks, calcareous algae,<br />
sand, and Foraminifera by the Calypso (Forest<br />
and Guinot, 1966).<br />
DISTRIBUTION.—Eastern Atlantic, from Scandinavia<br />
and the Hebrides southward to South<br />
Africa, including the Mediterranean, and, off the<br />
<strong>African</strong> Coast, a few localities between Morocco<br />
and Gabon, as well as the Cape Verde Islands;<br />
littoral to about 300 m. Monod (1956) summarized<br />
the literature but recorded no material of<br />
this species. In addition, the species has been<br />
recorded from the following localities.<br />
Morocco: Bane de Spartel, 35°54'N, 06°14'W, 210 m<br />
(Peres, 1964). Between Cap Rhir (as Cap Ghir) and Cap<br />
Draa (as Cap Noun), 10-30 m (Maurin, 1968a).<br />
Mauritania: Baie de l'Ouest (Bouvier, 1911).<br />
Senegal: Baie de Dakar, 0-17 m (Sourie, 1954b). 13°01'N,<br />
17°24'W, 51-55 m (Forest and Guinot, 1966).<br />
Sierra Leone: No specific locality, 121-160 m (Longhurst,<br />
1958).<br />
Although Gabon is well within the known
NUMBER 306 69<br />
range of the species, it is the southernmost locality<br />
on the tropical mainland.<br />
Atelecyclus undecimdentatus (Herbst, 1783)<br />
Atelecyclus cruentatus.—Capart, 1951:136, pi. 2: fig. 7 [Spanish<br />
Sahara].<br />
Atelecyclus undecimdentatus.—Monod, 1956:148, figs. 184-186<br />
[Mauritania, Senegal, Gambia, Gabon; references].—Forest,<br />
1958:472, fig. 1 [references].—Longhurst, 1958:87<br />
[Sierra Leone].—Forest and Gantes, 1960:350 [Morocco].<br />
—Zariquiey Alvarez, 1968:342, fig. 112d [Spain; references].—Maurin,<br />
1968b:486, 489, fig. 4 [Mauritania].—<br />
Bas, Arias, and Guerra, 1976, table 3 [Spanish Sahara].<br />
SYNONYMS.—Atelecyclus cruentatus Desmarest,<br />
1825; Atelecyclus omiodon Risso, 1827.<br />
DISTRIBUTION.—Eastern Atlantic, from France<br />
to Gabon, Mediterranean, from shore to about<br />
30 m.<br />
Family THIIDAE Dana, 1852<br />
THIIDAE Dana, 1852a: 120 [name 361 on Official List, dated<br />
1862 in error].<br />
EASTERN ATLANTIC GENERA.—One, Thia, represented<br />
in the fauna of tropical <strong>West</strong> Africa.<br />
EASTERN ATLANTIC SPECIES.—One (possibly<br />
two), occurring off tropical <strong>West</strong> Africa. The<br />
species reported as Thia residua by Monod (1956)<br />
is now known as Thia scutellata.<br />
REMARKS.—This family was not represented in<br />
the Pillsbury collections.<br />
Genus Thia Leach, 1815<br />
Thia Leach, 1815a:312 [type-species: Thiapolila Leach, 1815,<br />
a subjective junior synonym of Hippa scutellatus Fabricius,<br />
1793, by monotypy; gender: feminine; name 1577 on<br />
Official List].<br />
Thia scutellata (Fabricius, 1793)<br />
Hippa scutellata Fabricius, 1793:474.<br />
Thia residua.—Monod, 1956:153, figs. 186 bis, 186 ter [Sierra<br />
Leone; references].—Longhurst, 1958:87 [Sierra Leone].<br />
Thia scutellata.—Zariquiey Alvarez, 1968:343, fig. 11 If<br />
[Spain; references].—Christiansen, 1969:40, fig. 14, map<br />
8 [Scandinavia].<br />
Thia sp. aff. residua.—Forest and Guinot, 1966:57, fig. 3 [Sao<br />
Tome].<br />
SYNONYMS.—Cancer residuus Herbst, 1799; Thia<br />
polita Leach, 1815; Thia blainvillii Risso, 1822.<br />
REMARKS.—Forest and Guinot (1966) pointed<br />
out several differences between Monod's and their<br />
specimens from <strong>West</strong> Africa and European specimens<br />
and suggested that a direct comparison of<br />
material from the two areas might result in the<br />
recognition of a small, distinct <strong>West</strong> <strong>African</strong> species.<br />
DISTRIBUTION.—Eastern Atlantic, from the<br />
British Isles and Sweden southward to Portugal,<br />
Mediterranean, and off Sierra Leone and Sao<br />
Tome in the Gulf of Guinea; usually in depths<br />
between 10 and 25 m.<br />
Family CANCRIDAE Latreille, 1803<br />
Cancerides Latreille, 18O3b:35O [corrected to Cancridae by<br />
MacLeay, 1838:59].<br />
TRICHOCERIDAE Dana, 1852a: 120.<br />
EASTERN ATLANTIC GENERA.—One, not represented<br />
in the tropical fauna, is Cancer Linnaeus,<br />
1758:625. Type-species: Cancer pagurus Linnaeus,<br />
1758, by subsequent designation by Latreille,<br />
1810:422; gender: masculine; name 491 on Official<br />
List.<br />
EASTERN ATLANTIC SPECIES.—Two, both extralimital:<br />
Cancer bellianus Johnson, 1861. Shetland Islands<br />
and Iceland southward to Canary Islands, Madeira<br />
(Tiirkay, 1976b), and Spanish Sahara<br />
(Maurin, 1968b); sublittoral, to 620 m (Zariquiey<br />
Alvarez, 1968; Christiansen, 1969).<br />
Cancer pagurus Linnaeus, 1758. Northern Norway<br />
to the Mediterranean; sublittoral, between<br />
6 and 100 m (Zariquiey Alvarez, 1968; Christiansen,<br />
1969).<br />
Family PIRIMELIDAE Alcock, 1899<br />
Pirimelides A. Milne Edwards, 1862a:41; 1865:181, 204.<br />
PIRIMELINAE Alcock, 1899a:5, 95. [Elevated to Pirimelidae<br />
by Bouvier, 1940:217.]
70<br />
PERIMELIDAE Monod, 1956:147, 157. [Unjustified emendation.]<br />
EASTERN ATLANTIC GENERA.—Two, Pirimela<br />
and Sirpus, both represented in the fauna of tropical<br />
<strong>West</strong> Africa.<br />
EASTERN ATLANTIC SPECIES.—Four, three of<br />
which occur in tropical waters. The extralimital<br />
species, Sirpus zariquieyi Gordon, 1953, occurs in<br />
the Mediterranean (Lewinsohn and Holthuis,<br />
1964; Zariquiey Alvarez, 1968).<br />
REMARKS.—Monod (1956) recorded two species<br />
from <strong>West</strong> Africa, as listed below, and we<br />
add a previously undescribed species of Sirpus (S.<br />
gordonae), the only representative of the family<br />
collected by the Pillsbury:<br />
Name in Monod Current "Name<br />
Perimela denticulata Pirimela denticulata<br />
Sirpus monodi Sirpus monodi<br />
Genus Pirimela Leach, 1816<br />
Pirimela Leach, 1816, in 1815-1875, pi. 3 [type-species: Cancer<br />
denticulatus Montagu, 1808, by monotypy; gender: feminine;<br />
name 181 on Official List].<br />
Perimela Agassiz, 1846:280, 293 [unjustified emendation of<br />
Pirimela Leach, 1816; type-species: Cancer denticulatus Montagu,<br />
1808; gender: feminine].<br />
Pirimela denticulata (Montagu, 1808)<br />
Pirimela denticulata.—Bouvier, 1911:226 [PMauritania; listed].<br />
—Zariquiey Alvarez, 1968:350, figs. 7a, lid, 112a, 113a<br />
[Spain; references].—Christiansen, 1969:46, fig. 17, map<br />
11 [Scandinavia].—Tiirkay, 1976b:61 [listed], 64 [Madeira].<br />
Perimela denticulata.—Capart, 1951:137 [Spanish Sahara].—<br />
Monod, 1956:157, figs. 187-190 [Mauritania, Senegal,<br />
Gabon(?)].<br />
SYNONYM.—Pirimela princeps Hope, 1851.<br />
DISTRIBUTION.—Eastern Atlantic, from Norway<br />
to Senegal, Mediterranean, usually in shallow<br />
water, intertidal to 40-50 m. There is one questionable<br />
record from Gabon (Monod, 1956);<br />
Bouvier (1940) recorded material to a depth of<br />
200 m.<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Genus Sirpus Gordon, 1953<br />
Sirpus Gordon, 1953a:304 [type-species: Sirpus zariquieyi Gordon,<br />
1953, by original designation and monotypy; gender:<br />
masculine].<br />
Sirpus monodi Gordon, 1953<br />
Sirpus monodi.— Monod, 1956:159, figs. 191-193 [Mauritania,<br />
Senegal; references].—Rossignol, 1962:115 [Congo].—<br />
Monod, 1963, fig. 33 [no locality],<br />
Zirpus Monodi.—Monod, 1963:124 [erroneous spelling].<br />
DISTRIBUTION.—Off <strong>West</strong> Africa, from Mauritania,<br />
Senegal, and the Congo, intertidal to a<br />
depth of 12 m.<br />
* Sirpus gordonae, new species<br />
FIGURE 15<br />
MATERIAL EXAMINED.—Pillsbury Material: Annobon:<br />
Sta 271, shore, 3c5 (includes holotype), 39 (L,W).<br />
DESCRIPTION.—Carapace (Figure I5a,b) about<br />
as long as wide, greatest width behind postorbital<br />
tooth, at level of first anterolateral tooth. Front<br />
(Figure I5a,b) 3-lobed rather than 3-toothed as<br />
in Sirpus monodi Gordon and S. zariquieyi Gordon,<br />
median lobe bluntly triangular, extending farthest<br />
anteriorly, submedian lobes shorter,<br />
rounded triangular in shape, pointed or with<br />
small, sharp, almost granuliform apex. Anterior<br />
supraorbital angle triangularly pointed or<br />
rounded, with small granuliform apex. Dorsal<br />
orbital margin straight, sloping posteriorly, terminating<br />
in distinct incision. Posterior orbital<br />
margin, between incision and postorbital tooth,<br />
rounded or angular. Postorbital tooth large,<br />
sharply pointed, curved anteriorly. Lateral margin<br />
(Figure I5a,b), posterior to postorbital tooth,<br />
with 3 spiniform teeth, anteriormost larger than<br />
postorbital, with margins, particularly anterior,<br />
crenulate. Posterior 2 lateral teeth short, small,<br />
contrasting greatly with anterior tooth, terminating<br />
in sharp apices, often dorsally directed or<br />
twisted. Surface of carapace strongly areolated,<br />
in this resembling other Sirpus species, with elevations<br />
as follows: 2 protogastric, 1 mesogastric,
NUMBER 306 71<br />
FIGURE 15.—Sirpus gordonae, new species, paratypes. Female, cl 2.8 mm, Pillsbury Sta 271: a,<br />
carapace. Male, cl 2.8 mm, Pillsbury Sta 271: b, carapace; c, abdomen; d, first pleopod; e, apex<br />
of first pleopod;/, second pleood.<br />
1 on each branchial region, and 2 submedian on<br />
cardiac region. No elevation present at base of<br />
posterior lateral tooth, but small spinule or tooth<br />
there on dorsal surface of carapace in some specimens.<br />
Posterior part of carapace rugose, with<br />
short transverse striae, as in other species of genus.<br />
Lower border of orbit terminating in blunt<br />
tooth. Antepenultimate segment of antennular<br />
peduncle, forming part of orbital floor, terminating<br />
in distinct anterior spine and spinules. Antennal<br />
peduncle extending beyond front.<br />
Male abdomen (Figure 15
72<br />
TYPE-LOCALITY.—Annobon Island, 01°25'S,<br />
05°38'E, Gulf of Guinea.<br />
DISPOSITION OF TYPES.—The holotype (Crust.<br />
D. 31543), a male specimen with carapace length<br />
and width 2.6 mm, is in the Rijksmuseum van<br />
Natuurlijke Historie, Leiden. Two paratypes, a<br />
male and a female, also are in that collection and<br />
a male and two female paratypes (USNM<br />
169534) are in the National Museum of Natural<br />
History, <strong>Smithsonian</strong> <strong>Institution</strong>, Washington,<br />
D. C.<br />
ETYMOLOGY.—It is a great pleasure to dedicate<br />
this new species to Dr. Isabella Gordon, British<br />
Museum (Natural History), who was the first to<br />
make the existence of the genus Sirpus known to<br />
the world and who exhaustively treated the first<br />
two species assigned to it.<br />
DISTRIBUTION.—Our material of Sirpus gordonae<br />
was obtained at Annobon during a fish poison<br />
station at a sandy beach in a rocky cove. Sirpus<br />
zariquieyi is a Mediterranean species, known from<br />
scattered localities between Spain and Israel; it<br />
has been found in depths between 1 to about 40<br />
m. Sirpus monodi is known from NW Africa, from<br />
Mauritania and Senegal, and has been taken in<br />
depths between 0 and 12 m. This represents the<br />
first find of the genus in the southern hemisphere.<br />
Family CORYSTIDAE Samouelle, 1819<br />
CORYSTIDAE Samouelle, 1819:82 [name 357 on Official List).<br />
NAUTILOCORYSTIDAE Ortmann, 1893a:26, 28.<br />
EURYALIDAE Rathbun, 1930:10.<br />
EASTERN ATLANTIC GENERA.—Two, only one of<br />
which, Nautilocorystes, is represented in the tropical<br />
fauna. The other genus is Corystes Bosc (1802:65).<br />
Type-species: Hippa dentata Fabricius, 1793, by<br />
monotypy; gender: masculine; name 1571 on Official<br />
List.<br />
EASTERN ATLANTIC SPECIES.—Two, the first of<br />
which is the only species of the family occurring<br />
in tropical waters:<br />
Nautilocorystes ocellatus, was first discovered there<br />
after Monod's account of the tropical species;<br />
Monod remarked (1956:155) that it occurred as<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
far north as 17°S. No representatives of the family<br />
were taken by the Pillsbury.<br />
Corystes cassivelaunus (Pennant, 1777). Southern<br />
Norway and Sweden southward to Gibraltar and<br />
the Mediterranean, in depths between 7 and 90<br />
m; both Zariquiey Alvarez (1968) and Christiansen<br />
(1969) gave accounts of this species.<br />
Genus Nautilocorystes H. Milne Edwards, 1837<br />
Dicera de Haan, 1833:14 [invalid junior homonym of Dicera<br />
Germar, 1817 (Hymenoptera); type-species: Corystes (Dicera)<br />
octodentata de Haan, 1833, a subjective junior synonym<br />
of Corystes ocellatus Gray, 1831, by monotypy; gender:<br />
feminine].<br />
Nautilocorystes H. Milne Edwards,, 1837:149 [type-species:<br />
Nautilocorystes ocellatus H. Milne Edwards, 1837, an invalid<br />
junior homonym of Corystes ocellatus Gray, 1831, by<br />
monotypy; gender: masculine].<br />
Alyptes Gistel, 1848:ix [substitute name for Dicera de Haan,<br />
1833; type-species: Corystes (Dicera) octodentata de Haan,<br />
1833, a subjective junior synonym of Corystes ocellatus Gray,<br />
1831; gender: masculine].<br />
Nautilocorystes ocellatus (Gray, 1831)<br />
Corystes ocellata Gray, 1831:39.<br />
Nautilocorystes ocellata.—Barnard, 1950:303, figs. 57a-c [South<br />
Africa].—Monod, 1956:155 [Great Fish Bay (= Baia dos<br />
Tigres), Angola].—Rossignol, 1962:115 [listed].—Crosnier,<br />
1967:324 [Congo].<br />
Nautilocorystes ocellatus.—Capart, 1951:110, fig. 38 [Waivis<br />
Bay, Southwest Africa].<br />
SYNONYMS.—Corystes {Dicera) octodentata de<br />
Haan, 1833; Nautilocorystes ocellatus H. Milne Edwards,<br />
1837.<br />
DISTRIBUTION.—South Africa and South-<strong>West</strong><br />
Africa northward to Angola and off the Congo,<br />
in depths between 18 and 82 m.<br />
Family BYTHOGRAEIDAE Williams, 1980<br />
BYTHOGRAEIDAE Williams, 1980:444.<br />
This family, comprising a single genus and<br />
species from the Galapagos Rift in the eastern<br />
Pacific, is not represented in the study area.<br />
Family PORTUNIDAE Rafinesque, 1815<br />
PORTUNIDIA Rafinesque, 1815:97 [corrected to Portunidae<br />
by Samouelle, 1819:83; name 69 on Official List].<br />
MEGALOPIDAE Haworth, 1825:184.
NUMBER 306 73<br />
CARCINIDAE MacLeay, 1838:59.<br />
LUPINAE Dana, 1851b: 129.<br />
ARENAEINAE Dana, 1851b: 129.<br />
PLATYONYCHIDAE Dana, 1851b: 130.<br />
PODOPHTHALMIDAE Dana, 1851b: 130.<br />
Neptuniden Nauck, 1880:65.<br />
THALAMITINAE Miers, 1886:170, 193.<br />
CAPHYRINAE Miers, 1886:170.<br />
Cronius ruber<br />
Neptunus vocans<br />
Neptunus validus<br />
Neptunus inaequalis<br />
Callinectes gladiator<br />
Callinectes marginatus<br />
Callinectes latimanus<br />
Cronius ruber*<br />
Portunus vocans<br />
Portunus validus*<br />
Portunus inaequalis *<br />
Callinectes pallidus *<br />
Callinectes marginatus *<br />
Callinectes amnicola *<br />
POLYBIIDAE Ortmann, 1893a:66, 68.<br />
The extralimital species are as follows:<br />
CARUPIDAE Ortmann, 1893a:66, 68.<br />
Bathynectes longipes (Risso, 1816). England to<br />
LISSOCARCINIDAE Ortmann, 1893a:67, 87.<br />
LUPOCYCLOIDA Alcock, 1899a:22.<br />
PORTUMNINAE Ortmann, 1899:1170.<br />
GONIOCAPHYRINAE Borradaile, 1900:577.<br />
XAIVIDAE Berg, 1900:224.<br />
Portugal, Madeira, Mediterranean; sublittoral,<br />
20-90 m (Zariquiey Alvarez, 1968; Tiirkay,<br />
1976b).<br />
Bathynectes maravigna (Prestandrea, 1839). See<br />
CATOPTRINAE Borradaile, 1902a: 200.<br />
page 76.<br />
LIOCARCININAE Rathbun, 1930:18.<br />
MACROPIPINAE Stephenson and Campbell, 1960:75, 76, 88.<br />
Callinectes sapidus Rathbun, 1896. A. Milne Edwards<br />
and Bouvier (1900:71, pi. 4: fig. 5) reported<br />
EASTERN ATLANTIC GENERA.—Twelve, of which<br />
three specimens (2 6, 1 9) of "Callinectes diacanthus,<br />
nine, Bathynectes, Callinectes, Carcinus, Cronius, Lio-<br />
Latr., var. africanus, A. M.-Edw." from Porto da<br />
carcinus, Macropipus, Portunus, Thalamita, and Xaiva,<br />
Praia (as Praya), Cape Verde Islands. The colored<br />
are represented by tropical species. Three genera<br />
figure that they published shows definitely Calli-<br />
do not occur in the tropical region:<br />
nectes sapidus, having only two frontal teeth. As<br />
Charybdis de Haan (1833:3, 10). Type-species:<br />
Monod (1956:204) pointed out, the figure is noth-<br />
Cancer sexdentatus Herbst, 1783, a subjective junior<br />
ing but a colored copy of the figure that A. Milne<br />
synonym of Cancer feriatus Linnaeus, 1758, by Edwards (1861:425, pi. 30: fig. 1) published of<br />
subsequent designation by Glaessner (1929:113); "Neptunus diacanthus. Individu male de gran-<br />
gender: feminine; name 1616 on Official List. deur naturelle, rapporte des cotes de l'Amerique<br />
Polybius Leach (1820, in 1815-1875, pi. 9b: figs.<br />
septentrionale. (Variete a front quadrilobe)." As<br />
1-4). Type-species: Polybius henslowii Leach, 1820, to the identity of the three specimens taken by<br />
by monotypy; gender: masculine; name 184 on the Talisman, Monod (1956:204) stated: "J'ai vu<br />
Official List.<br />
un des 6 d'africanus (M.P.): c'est un C. marginatus."<br />
Portumnus Leach (1814:391, 429-430). Type- This identification is confirmed by Williams<br />
species: Cancer latipes Pennant, 1777, by mono- (1974:729), who likewise examined the male spectypy;<br />
gender: masculine; name 185 on Official imen in the collection of the Paris museum. The<br />
List.<br />
female specimen of the Talisman set now forms<br />
EASTERN ATLANTIC SPECIES.—36, of which 16 part of the collection of the Museum of Compar-<br />
occur in tropical waters. The following species ative Zoology, Harvard University (no. 6530) and<br />
were recorded by Monod (1956):<br />
was likewise examined by Williams (1974:729); it<br />
also is C. marginatus. The third specimen of the lot,<br />
Name in Monod Current Name<br />
a male (USNM 23950), is now in the collection<br />
Carcinus maenas Carcinus mamas<br />
of the National Museum of Natural History,<br />
Xaiva biguttata Xaiva biguttata<br />
Xaiva mcleayi Xaiva mcleayi<br />
<strong>Smithsonian</strong> <strong>Institution</strong>, Washington, D.C.; it<br />
Portunus arcuatus Liocarcinus arcuatus<br />
was examined by us, and like the two other<br />
Portunus corrugatus Liocarcinus corrugatus<br />
specimens belongs to C. marginatus. A. Milne Ed-<br />
Portunus tuberculatus<br />
Bathynectes superbus<br />
Thalamita africana<br />
Macropipus rugosus *<br />
Bathynectes piperitus,<br />
new species*<br />
Thalamita poissonii<br />
wards and Bouvier (1900:71) thus did not correctly<br />
report C. sapidus from <strong>West</strong> Africa.<br />
A second dubious record of C. sapidus from
74<br />
<strong>West</strong> Africa is the one by Gruvel (1912, pi. 2: fig.<br />
1), who in an account of the edible Crustacea of<br />
<strong>West</strong> Africa figured as Callinectes africanus a specimen<br />
that clearly represents C. sapidus. The explanation<br />
of Gruvel's figure does not give any information<br />
about the locality or origin of the specimen<br />
figured. As Callinectes sapidus otherwise has<br />
not been reported from <strong>West</strong> Africa, notwithstanding<br />
intensive collecting, we agree with<br />
Monod (1956:204) who thinks that "on est bien<br />
oblige de penser- qu'ici encore le specimen represente<br />
est un sapidus (americain)."<br />
Although so far there has not been a single<br />
reliable record of Callinectes sapidus from <strong>West</strong><br />
Africa, and the species certainly is not native<br />
there, the possibility exists that it will be accidentally<br />
introduced in <strong>West</strong> <strong>African</strong> waters as it was<br />
in Europe, the Mediterranean, and Japan; for an<br />
enumeration of the European and eastern Mediterranean<br />
localities of the species see Holthuis<br />
(1969a:34, fig. 1) and Christiansen (1969:72, fig.<br />
29, map 23). Since 1969, introduced specimens<br />
have been reported from the North Sea off the<br />
English coast (Dr. R. W. Ingle, in litt.), Normandy,<br />
France (Maury, 1975:25), possibly from<br />
Nice (Tiirkay, 1971:129), the central Adriatic Sea<br />
coast of Italy (Froglia, 1972:48), Strait of Messina<br />
(Cavaliere and Berdar, 1977), and Japan (Dr. T.<br />
Sakai, in litt.). In view of what is said in the<br />
previous paragraphs, it is unlikely that Rathbun's<br />
(1921:384) suggestion that the European species<br />
came from a <strong>West</strong> <strong>African</strong> population is correct.<br />
Carcinus aestuarii Nardo, 1847. Mediterranean,<br />
also in Canary Islands, possibly in Atlantic adjacent<br />
to Mediterranean and possibly in portions<br />
of the Suez Canal; intertidal (Zariquiey Alvarez,<br />
1968). Until now this species was known as Carcinus<br />
mediterraneus Czerniavsky, 1884. Nardo's<br />
(1847b) account, however, based on material<br />
from the lagoon at Venice, clearly provides the<br />
oldest available name for the Mediterranean species<br />
of Carcinus.<br />
Charybdis helleri (A. Milne Edwards, 1867). Eastern<br />
Mediterranean, Egypt and Israel (from the<br />
Red Sea); sublittoral (Holthuis and Gottlieb,<br />
1958; Ramadan and Dowidar, 1976).<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Charybdis longicollis Leene, 1938. Eastern Mediterranean,<br />
Egypt to Turkey (from the Red Sea);<br />
sublittoral (Holthuis, 1961; Lewinsohn and Holthuis,<br />
1964; Ramadan and Dowidar, 1976).<br />
Liocarcinus bolivari (Zariquiey Alvarez, 1948).<br />
Mediterranean, 8-60 m (Zariquiey Alvarez,<br />
1968).<br />
Liocarcinus depurator (Linnaeus, 1758). Norway<br />
to Spanish Sahara, Mediterranean; in depths to<br />
450 m, usually shallower than 100 m (Zariquiey<br />
Alvarez, 1968; Christiansen, 1969).<br />
Liocarcinus holsatus (Fabricius, 1798). Hebrides<br />
southward to Portugal, Canary Islands, possibly<br />
also Cap Blanc, Mauritania (Monod, 1956, recorded<br />
a juvenile 8 X 10 mm from there); in<br />
depths to 350 m, generally in less than 100 m<br />
(Zariquiey Alvarez, 1968; Christiansen, 1969).<br />
Liocarcinus maculatus (Risso, 1827). Mediterranean<br />
records of L. pusillus are referable to this<br />
species, according to a study by Manning and C.<br />
Froglia, Laboratorio di Tecnologia della Pesca,<br />
Ancona, to be published in the proceedings of II<br />
Colloquium Crustacea Decapoda Mediterranea<br />
held in Ancona, Italy, in May 1979.<br />
Liocarcinus marmoreus (Leach, 1814). Southern<br />
North Sea and British Isles southward to Spain,<br />
Azores, and Madeira; in shallow water to about<br />
85 m (Zariquiey Alvarez, 1968; Christiansen,<br />
1969; Tiirkay, 1976b).<br />
Liocarcinus puber (Linnaeus, 1767). Norway to<br />
Spanish Sahara, Mediterranean; littoral to 70 m<br />
(Zariquiey Alvarez, 1968; Christiansen, 1969).<br />
Liocarcinus pusillus (Leach, 1815). Norway to<br />
Portugal, possibly on northwest <strong>African</strong> coast (see<br />
Christiansen, 1969, for comments); in depths<br />
from 6 to 200 m, usually less than 50 m (Zariquiey<br />
Alvarez, 1968).<br />
Liocarcinus vernalis (Risso, 1816). Mediterranean,<br />
and possibly (doubtfully) off Cabo Blanco,<br />
Spanish Sahara (Monod, 1956); shallow water<br />
(Zariquiey Alvarez, 1968).<br />
Liocarcinus zariquieyi Gordon, 1968. England to<br />
Canary Islands, Mediterranean; 5 to 30 m (Zariquiey<br />
Alvarez, 1968).<br />
Macropipus tuberculatus (Roux, 1830). Norway to<br />
the Azores, Morocco, Mediterranean; 20-30 to
NUMBER 306 75<br />
834 m (Zariquiey Alvarez, 1968; Christiansen,<br />
1969).<br />
Polybius henslowii Leach, 1820. North Sea and<br />
British Isles southward to Morocco, Mediterranean;<br />
pelagic (Zariquiey Alvarez, 1968; Christiansen,<br />
1969).<br />
Portumnus latipes (Pennant, 1777). North Sea<br />
and British Isles to northern Morocco (there is<br />
one doubtful record from Mauritania), Mediterranean;<br />
intertidal and subtidal to a depth of 28<br />
m (Zariquiey Alvarez, 1968; Christiansen, 1969).<br />
Portumnus pestai Forest, 1967. Mediterranean;<br />
shallow water (Forest, 1967).<br />
Portunus pelagicus (Linnaeus, 1758). Mediterranean<br />
(from the Red Sea, Egypt to Turkey, Italy);<br />
sublittoral (Holthuis and Gottlieb, 1958; Ramadan<br />
and Dowidar, 1976).<br />
Portunus sayi (Gibbes, 1850). Canary Islands (see<br />
P. hastatus, p. 101); Cap Spartel, Morocco and<br />
Balearic Islands, from drifting Sargassum (Bouvier,<br />
1922; Zariquiey Alvarez, 1968). Bouvier (1940)<br />
does not refer to his earlier record of this species<br />
from the Balearic Islands, so there is some question<br />
as to its authenticity. There are no other<br />
records of this species from the Mediterranean.<br />
Subfamily CARCININAE MacLeay, 1838<br />
Genus Carcinus Leach, 1814<br />
Ligia Weber, 1795:92 [type-species: Cancer granarius Herbst,<br />
1783, a subjective junior synonym of Cancer maenas Linnaeus,<br />
1758, by monotypy; gender: feminine; name suppressed<br />
by the International Commission on Zoological<br />
Nomenclature in Opinion 330, 1955, and placed on the<br />
Official Index as name 207].<br />
Carcinus Leach, 1814:390 [type-species: Cancer maenas Linnaeus,<br />
1758, by monotypy; gender: masculine; name 798<br />
on Official List].<br />
Megalopa Leach, 1814:431 [type-species: Cancer granarius<br />
Herbst, 1783, a subjective junior synonym of Cancer maenas<br />
Linnaeus, 1758, by present selection; gender: feminine].<br />
Macropa Latreille, 1822:9 [type-species: Megalopa montagui<br />
Leach, 1817, an objective synonym of Cancer rhomboidalis<br />
Montagu, 1804, a subjective junior synonym of Cancer<br />
maenas Linnaeus, 1758, by monotypy; gender: feminine].<br />
Megalops H. Milne Edwards, 1837:260 [erroneous spelling of<br />
Megalopa Leach, 1814].<br />
Sympractor Gistel, 1848:ix [replacement name for Megalopa<br />
Leach, 1814 (as Megalopus); type-species: Cancer granarius<br />
Herbst, 1783, a subjective junior synonym of Cancer maenas<br />
Linnaeus, 1758; gender: masculine].<br />
Carcinides Rathbun, 1897b: 164 [replacement name for Carcinus<br />
Leach, 1814; type-species: Cancer maenas Linnaeus,<br />
1758; gender: masculine; name 209 on Official Index].<br />
Carcinus maenas (Linnaeus, 1758)<br />
Carcinus maenas.—Monod, 1956:165, fig. 194 [Mauritania;<br />
references].—Forest and Games, 1960:351 [Morocco].—<br />
Guinot, 1967a:253 [Indian Ocean localities; listed].—<br />
Monod, 1967:178 [no locality].—Zariquiey Alvarez, 1968:<br />
354, figs. 115a, c [Spain; references].—Christiansen, 1969:<br />
49, fig. 18, map 12 [Scandinavia].<br />
SYNONYMS.—Cancer viridis Herbst, 1783; Cancer<br />
granarius Herbst, 1783; Cancer pygmaeus Fabricius,<br />
1787; Cancer rhomboidalis Montagu, 1804; Megalopa<br />
montagui Leach, 1817; Portunus menoides Rafinesque,<br />
1817; Cancer granulatus Say, 1817.<br />
DISTRIBUTION.—Eastern Atlantic, from Norway,<br />
N of 70° N latitude, S to Mauritania, about<br />
20° S latitude; western Atlantic; eastern Pacific;<br />
Australia. In the Mediterranean it is replaced by<br />
the allied C. aestuarii Nardo, 1847 (= C. mediterraneus<br />
Czerniavsky, 1884) (see Bacescu, 1967, and<br />
Zariquiey Alvarez, 1968, for differences between<br />
the two species).<br />
Genus Xaiva MacLeay, 1838<br />
Xaiva MacLeay, 1838:62 [type-species: Xaiva pulchella Mac-<br />
Leay, 1838 (? = Portunus biguttatus Risso, 1816), by monotypy;<br />
gender: feminine; name 1649 on Official List].<br />
Portumnoides Bohn, 1901:270, 271 [type-species: Portumnoides<br />
garstangi Bohn, 1901, a subjective junior synonym of Portunus<br />
biguttatus Risso, 1816, by monotypy; gender: masculine].<br />
Xaiva biguttata (Risso, 1816)<br />
Xaiva biguttata.—Monod, 1956:168, fig. 195 [Cape Verde<br />
Islands].—Guinot, 1967a:253 [Indian Ocean; listed].—<br />
Zariquiey Alvarez, 1968:359, figs. 8d, 116c, 122b [Spain;<br />
references].—Kensley, 1970:182 [South-<strong>West</strong> Africa].—<br />
PenrithandKensley, 1970b:248,261 [South-<strong>West</strong> Africa].<br />
SYNONYMS.—Platyonichus nasutus Latreille, 1828;<br />
Portumnoides garstangi Bohn, 1901.
76<br />
REMARKS.—Material of this species from the<br />
northern and southern parts of its reported range<br />
(England to Cape Verde Islands versus South-<br />
<strong>West</strong> Africa and South Africa) should be studied<br />
to determine whether or not two species might be<br />
recognized. If the southern form proves to be<br />
distinct, the name Xaiva pulchella MacLeay, 1838,<br />
is available.<br />
DISTRIBUTION.—Eastern Atlantic, from England<br />
southward to the Cape Verde Islands,<br />
South-<strong>West</strong> Africa, South Africa, and the Mediterranean,<br />
in shallow water.<br />
Xaiva mcleayi (Barnard, 1947)<br />
Portumnus mcleayi.—Capart, 1951:114, fig. 40, pi. 2: fig. 9<br />
[Angola].—Rossignol, 1957:123 [key].<br />
Xaiva mcleayt.—Monod, 1956:169, 632, figs. 196-201 [Mauritania,<br />
Senegal, Sierra Leone; references].—Longhurst,<br />
1958:87 [Sierra Leone].—Guinot and Ribeiro, 1962:31<br />
[Angola].—Forest and Guinot, 1966:59 [Principe, Sao<br />
Tome].—Guinot, 1967a:253 [Indian Ocean; listed].—<br />
Crosnier, 1967:324 [Dahomey, Annobon].<br />
?Xaiva biguttata.—Gauld, 1960:69 [Ghana] [not Xaiva biguttata<br />
(Risso, 1816)].<br />
REMARKS.—We suspect that Gauld's record<br />
from Ghana of X. biguttata, a species not known<br />
to occur south of the Cape Verde Islands on the<br />
tropical <strong>West</strong> <strong>African</strong> Coast, may have been<br />
based on this species, now known to occur at least<br />
as far north as Mauritania on the <strong>West</strong> <strong>African</strong><br />
coast (Monod, 1956).<br />
DISTRIBUTION.—Eastern Atlantic, from scattered<br />
localities between Mauritania and Angola,<br />
South Africa, in depths between 8-30 and 73 m.<br />
Subfamily POLYBIINAE Ortmann, 1893<br />
Genus Bathynectes Stimpson, 1871<br />
Bathynectes Stimpson, 1871a: 145 [type-species: Bathynectes longispina<br />
Stimpson, 1871, by subsequent designation by<br />
Rathbun, 1930:27; gender: masculine; name 127 on Official<br />
List].<br />
Thranites Bovallius, 1876:60, 61 [type-species: Thranites velox<br />
Bovallius, 1876, a subjective junior synonym of Portunus<br />
maravtgna Prestandrea, 1839, by monotypy; gender: masculine].<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Bathynectes maravigna (Prestandrea, 1839),<br />
new combination<br />
Portunus Maravigna Prestandrea, 1839:132.<br />
Portunus superbus Costa, in Costa and Costa, 1838-1871: 19,<br />
pi. 5 [color].<br />
Bathynectes.—Filhol, 1885a:56.—Maurin, 1968a, fig. 29;<br />
1968b, figs. 1,4.<br />
Bathynectes superba.—Forest and Gantes, 1960:351 [Morocco].—Peres,<br />
1964:20, 26, 29 [Morocco].<br />
Bathynectes superbus.—Maurin, 1968a: 19, 45, 50, 64 [Spain,<br />
Spanish Sahara, Mauritania]; 1968b:482, 484, 489, 491,<br />
fig. 6 [Spanish Sahara, Mauritania].—Zariquiey Alvarez,<br />
1968:382, fig. 127g [Spain; references].—Christiansen,<br />
1969:70, fig. 28, map 22 [Scandinavia].—Tiirkay, 1976b:<br />
61 [listed], 64 [Madeira].—Lewis and Haefner, 1978:164<br />
[part; Spain].<br />
SYNONYM.—Thranites velox Bovallius, 1876.<br />
MATERIAL EXAMINED.—France: Bane de la Chapelle,<br />
off Brittany, 47°58'N, 08°00'W, 9 Sep 1921, L. Fage, 16\ 1$<br />
(MP).<br />
Spain: NE of Santander, (43°28'N, 03°48'W), 564 m,<br />
sand and shells, 6 Jul 1882, Travailleur, 6 juv (MP, W). Off<br />
western Galicia, 43°41.2'N, 08°57.6'W, 995 m, rocky bottom,<br />
8 Aug 1967, Thalassa Sta T.491, 1 juv (MP).<br />
Madeira: SE of Madeira, 32°43'N, 16°43'W, 420 m, 16<br />
(L).<br />
Morocco: Fosse de Rabat (approx. 34°N, 07°W), 300-<br />
400 m, N. Pigeault, 1
NUMBER 306 77<br />
delle articolazioni dei piedi camminatori. . . . Gli occhi sono m, 3 juv (W). Sta 220, 300 m, 1 carapace (W). Sta 247, 400<br />
sferici nerastri. [The anterolateral teeth] sono tutti del colore m, 1 9(W).<br />
del torace con l'estremita color caffe cotto. . . . Tutte le spine Undaunted Material: Angola: Sta 111, ca. 366 m, 2 6*<br />
dei piedi-mani [= chelipeds] sono bianche colla estremita (includes holotype), 39 (1 ov) (L).<br />
color caffe cotto; pero quella del lato interno del carpo a Other Material: Cape Verde Islands: 16°53'N, 27°30'W to<br />
una banda bianca; quindi una rosso-corallo chiara e la 16°54'N, 27°30'W of Paris [= 25° 10'W of Greenwich], 410estremita<br />
come le altre. [The fingers are] verso l'estremita 460 m, sand and gravel, 29 Jul 1883, Talisman Sta 110, 16<br />
brunastre.<br />
(MP). 15°14'N, 23°03'45"W, 628 m, muddy sand, fish trap,<br />
In listing material of this species that does not<br />
occur off tropical <strong>West</strong> Africa, we have departed<br />
14-15 Aug 1901, Princesse Alice Sta 1189, 19 (MP).<br />
Senegal: NW of Pointedes Almadies (14°45'N, 17°32'W),<br />
250-300 m, 21 Oct 1952, Ghard Tre'ca, 26, 19 (MP). Off<br />
from our format. However, the material listed Pointe des Almadies, ca. 300 m, 16 Oct 1952, Cremoux, leg.,<br />
above was used to compare this species with the Gerard Tre'ca, 19 (MP).<br />
new species from <strong>West</strong> Africa described below. DESCRIPTION—Front (Figures 16, 17a) with 4<br />
Material from Spanish Sahara and Mauritania rounded teeth, inner 2 about half as wide as<br />
identified with this species by Maurin (1968a,b) outer, extending about as far forward as outer.<br />
might well be referable to B. piperitus, new species. Latter broadly rounded, more semicircular than<br />
This cannot be determined without examining triangular. Inner orbital teeth low, rectangular,<br />
his material.<br />
wide, with blunt apex and short dorsal carina.<br />
DISTRIBUTION.—Eastern Atlantic, from Norway Upper orbital margin with 2 distinct narrow,<br />
and the Faroes southward to NW Morocco, pos- open fissures. Exorbital tooth triangular, blunt,<br />
sibly to Spanish Sahara and Mauritania, Medi- especially in larger specimens. Lower orbital marterranean;<br />
in moderately deep water, from 100 to gin with small, triangular, often blunt, narrow<br />
1455 m.<br />
tooth next to exorbital tooth. Third tooth present<br />
medially, wider, blunter than second, separated<br />
*Bathynectes piperitus, new species from second by U-shaped incision. Inner, lower<br />
FIGURES 16, 17<br />
orbital tooth triangular, with narrowly rounded<br />
apex, latter occasionally curved inward. First 3<br />
Bathynectes superba.—A. Milne Edwards and Bouvier, 1899, anterolateral teeth of carapace, excluding exor-<br />
pi. 2: figs. 16-18 [part, not p. 25, pi. 2: figs. 1-15, bital tooth, sharply triangular, curved anteriorly,<br />
19-24]; 1900:65 [part].—Bouvier, 1922:59 [part, Cape<br />
Verde Islands only].—Monod, 1933b:510 [listed].—Ca- third smallest. Fourth (= last) tooth large, spinepart,<br />
1951:121, fig. 42. [Not Portunus superbus Costa, 1853.] like, curved anteriorly, especially apically. Dorsal<br />
Bathynectes superbus.—Monod, 1956:183, figs. 210-212.— surface of carapace with 4 transverse ridges: (1)<br />
Longhurst, 1958:87.—Guinot and Ribeiro, 1962:45.— indistinct postfrontal ridge, widely interrupted in<br />
Forest, 1963:628.—Monod, 1967, pi. 15: fig. 4 [no records].—Maurin,<br />
1968b:492.—Intes and Le Loeuff, 1976:<br />
103.—Lewis and Haefner, 1978:164, pi. la [part]. [Not<br />
Portunus superbus Costa, 1853.].<br />
Bathynectes suberbus.—Gauld, 1960:69 [erroneous spelling].<br />
Bathynectes.—Voss, 1966:19, 25.—Maurin, 1968a, fig. 29;<br />
1968b, fig. 9 [part, Senegal only].<br />
midline; (2) mesogastric ridge, divided by median<br />
MATERIAL EXAMINED.—Pilbbury Material: Liberia:<br />
Sta 73, 311-366 m, 13 6, 4 9 ov (L, W).<br />
Ivory Coast: Sta 44, 403-586 m, hard dark gray mud,<br />
5 juv (L). Sta 51, 329-494 m, 31 6\ 16 9 (1 ov), 13 juv (L,<br />
W).<br />
Geronimo Material: Gabon: Sta 179, 293 m, 1 9 (W). Sta<br />
191, 300 m, 2 6 (W). Sta 198, 300 m, 3 6 (W). Sta 199, 400<br />
m, 1 6, 1 9 (W). Sta 203, 200 m, 3 6\ 2 9 (W). Sta 206, 455-<br />
610 m, 1 9, 1 juv (W). Sta 213, 300 m, 1 9 (W). Sta 214, 546<br />
FIGURE 16.—Bathynectes piperitus, new species (from Capart,<br />
1951, fig. 42).
78<br />
FIGURE 17.—Bathynectes piperitus, new species: a, dorsal view;<br />
b, c, first pleopod of male and apex; d, second pleopod of<br />
male. (All from Monod, 1956, figs. 210-212.)<br />
interruption, each half again divided into 2 short,<br />
slightly convex ridges; (3) long lateral spines connected<br />
by third transverse ridge, sinuous and<br />
tuberculate; third ridge convex anteriorly, with<br />
posteriorly directed indentations (incurvations) at<br />
branchiocardiac grooves; and (4) short transverse<br />
carina placed between ends of branchiocardiac<br />
grooves on posterior half of carapace. Carapace<br />
finely and closely pubescent.<br />
Mouthparts not significantly different from<br />
those of Bathynectes longispina Stimpson.<br />
First pereiopods unequal, larger chela higher<br />
and heavier than smaller, with teeth on cutting<br />
edges of fingers larger, blunter. Otherwise chelipeds<br />
very similar. Fingers about as long as palm.<br />
Dactylus with usual 5 ridges, dorsalmost tuberculated<br />
basally. Upper ridge on inner surface of<br />
dactylus divided into 2 branches proximally,<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
branches united distally; upper branch with<br />
blunt tubercle. In larger chela, basal tooth of<br />
cutting edge of dactylus large, swollen, directed<br />
somewhat posteriorly. Opposite tooth of fixed<br />
finger molar-like, with 2 small teeth proximally<br />
at base. Fixed finger, like dactylus, with laterally<br />
compressed teeth, usually alternately large and<br />
small. In smaller chela all teeth on cutting edges<br />
of compressed type. Fixed finger also with 5<br />
ridges. Upper carina of palm with large, subdistal<br />
sharp tooth, lacking spinule adjacent to tooth;<br />
carina tuberculate posterior to tooth. Upper outer<br />
carina of palm with 6 large, triangular, blunt<br />
teeth, several smaller tuberculiform denticles<br />
present between larger teeth. Carina below upper,<br />
outer carina low, often partly absent, with row of<br />
small granules, latter often inconspicuous or absent,<br />
especially in middle of carina. Next lower<br />
carina strong, high and sharp, granular, terminating<br />
before reaching base of fingers; row of<br />
granules below distal end of this carina extending<br />
to first tooth of fixed finger. Lower part of palm<br />
with carina extending onto fixed fingers, more<br />
distinct proximally than distally, ornamented<br />
with numerous granules. All of lower surface of<br />
palm covered by transverse rows of granules;<br />
anteriorly this area terminates on ventral carina<br />
of fixed finger. Inner surface of palm with high,<br />
distinct longitudinal carina terminating somewhat<br />
above base of upper carina of inner surface<br />
of fixed finger. Carpus with very large, sharply<br />
pointed tooth on inner margin, anterior margin<br />
of tooth with 1 to 3, rarely 4, spinules, remainder<br />
of margin smooth or granular. Upper surface of<br />
carpus with 2 spines placed in line with upper<br />
articulation of palm, inconspicuous row of tubercles<br />
extending from posterior spine towards inner<br />
tooth, with more distinct row of tubercles extending<br />
obliquely proximally. Some tubercles scattered<br />
over remainder of upper surface of carpus.<br />
Outer anterior angle of carpus with sharp tooth.<br />
Anterior margin of carpus, between sharp tooth<br />
and upper articulation with propodus, smooth or<br />
evenly granular. Merus slender, more than twice<br />
as long as high, with strong spiniform tooth on<br />
anterior third of upper margin, second tooth
NUMBER 306 79<br />
somewhat before middle of inner, lower margin.<br />
Remainder of merus smooth or slightly tuberculate.<br />
Walking legs (pereiopods 2 to 4) very slender.<br />
Dactylus narrow, evenly curved, upper and lower<br />
surfaces with narrow, longitudinal groove over<br />
entire length; lateral surfaces with broad groove,<br />
distally divided in two by longitudinal carina.<br />
Propodus as long as dactylus, measured dorsally,<br />
and more than 5 to more than 6 times as long as<br />
high. Upper and lower margins with deep longitudinal<br />
groove, flanked by 2 sharp carinae. Outer<br />
surfaces with 2 longitudinal carinae separating 3<br />
grooves, middle groove particularly wide; upper<br />
carina placed close to outer carina of upper margin.<br />
On inner surface of propodus upper carina<br />
absent or very reduced, almost entirely covered<br />
by pubescence, thus only lower carina visible on<br />
surface. Carpus 2/3 to 4/7 length of propodus,<br />
with blunt longitudinal dorsal ridge, flanked on<br />
either side by distinct pubescent groove. Merus 2<br />
to 3V3 times as long as carpus and 4-5 times as<br />
long as wide, upper part with transverse subdistal<br />
groove, occasionally flanked posteriorly by blunt<br />
tubercle. Upper surface of merus sometimes<br />
slightly tuberculate. Dactylus of last pereiopod 3<br />
times as long as high, oval, with lower margin<br />
straight for most of length, merging with distal<br />
tooth in straight line. Upper margin regularly<br />
curved, merging with distal tooth in concave line.<br />
Propodus about 2/3 as long as dactylus. Carpus<br />
less than half as long as dactylus. Merus about as<br />
long as propodus and 2 l k times as long as high.<br />
Male abdomen triangular, third to fifth somites<br />
fused, sixth and seventh somites free, latter triangular,<br />
apex broadly rounded, appearing<br />
broader than that of B. longispina.<br />
Male gonopods similar to those of B. longispina,<br />
with apex less slender, less strongly curved laterally,<br />
as illustrated by Monod (1956, fig. 211)<br />
(Figure \lb-d).<br />
Color: The first color note published of this<br />
species is the one by Capart (1951:122): "Couleur<br />
rouge orange, les epines generalement plus<br />
rouges; les pattes marquees de zones rouges et<br />
orange alternees." Monod (1956:185) described<br />
the color as follows: "gris a rouge-brique, parfois<br />
rose vif; chelipedes en partie rouges, un anneau<br />
rouge sur les merus, carpe, propode et base du<br />
dactyle des pattes." The following color description<br />
is made after notes and photographs taken<br />
of specimens of Pillsbury Station 73, immediately<br />
after they came aboard: The carapace is orangered<br />
with a white spot at the front, one such spot<br />
just behind either of the large lateral teeth, one<br />
in each posterolateral angle, and one in the middle<br />
of the posterior margin. The apex of the<br />
lateral spine is very dark, almost black. The chelipeds<br />
are orange with the main spines and teeth<br />
white. Also the distal part of the palm and the<br />
fingers are white, except for an orange spot at the<br />
base of the dactylus. The tips of the fingers and<br />
a large part of the cutting edges are of a dark,<br />
almost black color. The walking legs have a broad<br />
orange band over the distal part of the merus,<br />
one over the carpus and one over the proximal<br />
half of the propodus, sometimes the proximal<br />
part of the dactylus also is orange. All the rest of<br />
the leg is white, viz., the base of the merus, all the<br />
articulations, the distal part of the propodus and<br />
the entire dactylus or the larger part of it. In the<br />
fifth leg the orange color is somewhat more extensive<br />
than in the other legs, here the basal part of<br />
the merus is white, from there on the leg is orange<br />
up to the middle of the propodus or slightly<br />
beyond (with the exception of small white articulations),<br />
and also the basal part of the dactylus<br />
is orange, the distal part white. The lower surfaces<br />
of body and chelipeds are white, that of the<br />
second to fifth pereiopods is of the same color as<br />
the dorsal surface. The eggs are carmine.<br />
MEASUREMENTS.—The specimens examined<br />
had the carapace width from 15 to 86 mm (inclusive<br />
of the lateral spine); the largest specimen is<br />
a male. The ovigerous females have carapace<br />
widths between 55 and 72 mm; the largest of the<br />
females is ovigerous. In the literature, male specimens<br />
with carapace lengths from 17 to 57 mm<br />
and carapace widths 32 to 80 mm (including the<br />
lateral spines) are recorded; for the non-ovigerous<br />
females these values are 22 to 49 mm and 38 to<br />
68 mm, respectively, and for the ovigerous fe-
80<br />
males 40 to 54 mm and 62 to 78 mm. The<br />
diameter of the eggs is 0.42 mm (Monod, 1956).<br />
REMARKS.—Most authors have considered all<br />
Atlantic Bathynectes, other than Bathynectes longipes<br />
(Risso, 1816), to belong to a single species, B.<br />
superbus (Costa, 1853), the type-locality for which<br />
is Naples, Italy. As stated (p. 76), that species<br />
should be known as Bathynectes maravigna (Prestan-<br />
drea, 1839), type-locality "Mare di Messina,"<br />
Sicily, Italy. However, a comparison of the present<br />
<strong>West</strong> <strong>African</strong> material with <strong>West</strong> Atlantic<br />
specimens and with material from the Mediterranean,<br />
European, and NW <strong>African</strong> waters,<br />
showed constant differences between the three<br />
groups, which leads to the conclusion that three<br />
species are involved. For the <strong>West</strong> Atlantic species<br />
the name Bathynectes longispina Stimpson, 1871, is<br />
available (B. brevispina Stimpson, 1871, is a synonym).<br />
The name Bathynectes maravigna (Prestandrea,<br />
1839) has to be used for the species from<br />
the Mediterranean and NE Atlantic. A new<br />
name, Bathynectes piperitus, is proposed here for the<br />
<strong>West</strong> <strong>African</strong> species. Bathynectes piperitus differs<br />
from B. maravigna in the following respects:<br />
1. The frontal teeth of the carapace in B.<br />
piperitus are always blunt, even in the large specimens,<br />
and the median teeth are always distinctly<br />
narrower than the outer. In B. maravigna the large<br />
specimens have the frontal teeth sharply pointed<br />
and triangular, while the median teeth are hardly<br />
narrower than the outer. In juveniles of B. maravigna<br />
the situation is much like in the large B.<br />
piperitus.<br />
2. The lateral spine of the carapace in B. maravigna<br />
is long and straight, in B. piperitus it is<br />
much shorter than in B. maravigna and is curved<br />
forward in the adults. Although in juveniles of B.<br />
piperitus the lateral spines are straighter and relatively<br />
longer than in the adults, they are still<br />
shorter than in the corresponding stages of B.<br />
maravigna.<br />
3. The postfrontal ridges in B. maravigna as a<br />
rule are more distinct than in B. piperitus, and<br />
each of the two mesogastric carinae is mostly<br />
entire and not divided in two short ridges.<br />
4. The last somite of the male abdomen is<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
wider and flatter in B. piperitus, being relatively<br />
longer and narrower, with a convex upper surface,<br />
in B. maravigna. The fourth and fifth abdom-<br />
inal somites in males of B. piperitus each show a<br />
distinct transverse carina, which are absent or<br />
very vague in B. maravigna.<br />
5. The outer half of the anterior margin of the<br />
carpus of the cheliped (between the outer spine<br />
and the upper articulation with the palm) in B.<br />
piperitus usually is straight or finely granular; in<br />
B. maravigna it carries some spinules.<br />
6. The inner surface of the propodus of the<br />
second pereiopod in B. maravigna shows two distinct<br />
longitudinal carinae separating three longitudinal<br />
pubescent grooves. In B. piperitus the upper<br />
of these carinae is absent or hardly indicated,<br />
so that the lower carina is separated from the<br />
upper margin of the propodus by a pubescent<br />
area that is much wider than the area separating<br />
it from the lower margin.<br />
7. The merus of the fifth pereiopod is longer<br />
and more slender in B. maravigna than in B.<br />
piperitus. The dactylus of this leg in B. maravigna<br />
is slightly more slender than in B. piperitus and<br />
has the lower margin somewhat convex and joining<br />
the distal tooth under a concave curve.<br />
8. The coloration of living B. maravigna is quite<br />
different from that of B. piperitus. The colored<br />
plate of B. maravigna published by A. Milne Edwards<br />
and Bouvier (1899, pi. 2: fig. 19) shows a<br />
uniformly orange red crab. Costa's figure (in<br />
Costa and Costa, 1838-1871, pi. 7) of the species<br />
has the carapace irregularly mottled with reddish<br />
brown and yellowish, the pereiopods 1 to 4 are<br />
uniformly reddish brown, only the dactyli of the<br />
walking legs and a few spines being yellowish; the<br />
fifth pereiopods are shown entirely yellowish. A<br />
male specimen of B. maravigna (cl 61 mm, cb 73<br />
mm without lateral spines, 96 mm with lateral<br />
spines) from east of Madeira (11-12 Mar 1976,<br />
L) was examined rather soon after capture, when<br />
the colors were still visible and a color photograph<br />
of the living specimen was made. This specimen<br />
showed the following color pattern: The animal<br />
is dark brownish orange dorsally. The margins of<br />
the frontal teeth are white. A very short median
NUMBER 306 81<br />
line extends from between the bases of the median<br />
frontal teeth backward. The tips of the three short<br />
anterolateral spines of the carapace (not the outer<br />
orbital angle) have the tips white. One to three<br />
small white dots are seen on the middle of the<br />
large lateral spine. A few very small white spots<br />
are placed in the cervical groove. A large lateral<br />
white spot stands at each side of the carapace<br />
about halfway between the lateral spine and the<br />
posterior margin of the carapace. A narrow white<br />
line extends along the posterior carina and a<br />
small median spot is placed somewhat before this<br />
line. The chelipeds have dorsally the same color<br />
as the carapace, with a small white spot near the<br />
tips of many of the spines. The tips of the fingers<br />
and the teeth on their cutting edges are black.<br />
Also the tips of various of the spines are black.<br />
The lower surface of the chelipeds is lighter than<br />
the upper. The following legs are of the same<br />
brownish orange color as the carapace. The second<br />
and third legs have the basal part slightly<br />
lighter than the rest. There may be some small<br />
white spots at the articulations and at the tips of<br />
the second to fourth pereiopods, but otherwise<br />
the legs are of a uniform brownish orange color.<br />
The dactyli of the fifth leg show two very small<br />
spots: one just before the tip, the other near the<br />
middle of the lower margin, otherwise the leg is<br />
of a uniform dark brownish orange color. At first<br />
view B. maravigna seems to be uniformly brownish<br />
orange, and the white spots are rather small and<br />
inconspicuous. They evidently have been overlooked<br />
by the artist of A. Milne Edwards and<br />
Bouvier's colored plate. Costa's specimen may<br />
have been made after a specimen that was preserved<br />
or dead for a considerable time. The most<br />
important color difference between B. maravigna<br />
and B. piperitus seems to be the less striking color<br />
pattern of the carapace and the absence of white<br />
and orange bands on the pereiopods in the former<br />
species.<br />
9. The range of B. maravigna is in the N Atlantic<br />
Ocean between Iceland and Norway south to the<br />
Mediterranean (as far as Greece), Madeira, the<br />
Azores, and NW Morocco. We have examined<br />
specimens of this species from several localities<br />
between France and Morocco, as noted above.<br />
Bathynectes piperitus is a <strong>West</strong> <strong>African</strong> species,<br />
known from the Cape Verde Islands and Senegal<br />
to Angola.<br />
The differences between B. piperitus and B.<br />
longispina are as follows:<br />
1. In B. longispina the outer frontal teeth are<br />
more acute and are more triangular in shape; the<br />
inner teeth are narrowly triangular.<br />
2. The anterolateral teeth of B. longispina are<br />
longer, more robust, and relatively narrower than<br />
in B. piperitus. The length of the second tooth<br />
after the exorbital angle is greater than half the<br />
distance between it and the first tooth; in B.<br />
piperitus it is narrower than half that distance.<br />
3. The lateral spine of the carapace is straight<br />
in B. longispina and is longer than in B. piperitus.<br />
4. The transverse ridge that connects the lateral<br />
spines of the carapace in B. longispina is<br />
practically straight and does not show the posterior<br />
incurvations at the level of the branchiocardiac<br />
grooves.<br />
5. The two mesogastric ridges are each divided<br />
in two smaller ridges like in B. piperitus, but<br />
they are usually shorter, less distinct (sometimes<br />
hardly visible), and lie in one line.<br />
6. The last somite of the male abdomen in B.<br />
longispina is more triangular than in B. piperitus,<br />
and has the distal margin less broadly rounded.<br />
7. The fourth and fifth abdominal somites in<br />
B. longispina do not show a sharp transverse carina.<br />
8. The first male gonopod of B. longispina is<br />
more slender distally and more strongly curved<br />
than in B. piperitus.<br />
9. In B. longispina the chelae are relatively<br />
shorter and higher: the height of the palm (inclusive<br />
of the dorsal spine) is more than its dorsal<br />
length. In B. piperitus the dorsal length of the<br />
palm is definitely greater than the height.<br />
10. In B. longispina the subdistal tooth on the<br />
upper margin of the palm is much higher, being<br />
large, compressed, almost wing-like, reaching<br />
practically to the base of the dactylus. Before this<br />
large tooth a small but distinct spine is present<br />
on the dorsal margin. In B. piperitus the dorsal
82<br />
tooth is much shorter and less wide and does not<br />
reach the base of the dactylus; the dorsal margin<br />
bears no distal spine.<br />
11. The inner surface of the propodus of the<br />
second pereiopod in B. longispina shows a single<br />
deep longitudinal groove in the middle, and the<br />
two other grooves are inconspicuous. The legs<br />
have hardly any pubescence. Also the upper<br />
groove on the inner surface of the carpus is very<br />
obscure.<br />
12. The fifth legs in B. longispina have the<br />
merus more slender and the propodus less slender<br />
than in B. piperitus.<br />
13. Also in the color there are marked differences<br />
between the two species. In B. longispina the<br />
carapace is reddish because all or practically all<br />
the tubercles are of that color; the lateral spine is<br />
dark red in the larger distal part, the anterolateral<br />
teeth are whitish with a red tip or with a red ring.<br />
The front is pale and the orbital margin is often<br />
darker red than the surroundings. A median<br />
white line extends from the transverse ridge of<br />
the carapace forwards reaching to the postfrontal<br />
ridges. The mesogastric ridges, at least the inner<br />
two, are white and form a more or less distinct<br />
cross with the median white line. The chelipeds<br />
are pink; the large teeth on the inner margin of<br />
the carpus and the upper margin of the palm are<br />
white at the base and dark red distally. The<br />
dactylus is red with a white base and a light spot<br />
in the middle of the dorsal margin; the tip of the<br />
finger and practically the entire cutting edge (in<br />
the large chela) or the distal part of it (in the<br />
smaller) are black. The fixed finger has the distal<br />
part and the lower surface red, the tip black. The<br />
legs are pink because of the presence of scattered<br />
red spots; the articulations are pale or white. Very<br />
striking is the presence of a solid deep red color<br />
on the propodus and dactylus of the second to<br />
fourth leg; this color, which persists long in alcohol-preserved<br />
specimens, occupies practically the<br />
entire segments with the exception of a very<br />
narrow area near the articulation. The carpus<br />
sometimes shows a narrow red band over its<br />
middle. In the fifth leg the dactylus, but not the<br />
propodus, has this same strikingly deep red, al-<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
most lacquer-like, color. The presence of these<br />
deep red bands on the legs distinguishes this<br />
species immediately from the others belonging to<br />
this genus. The lower surface of the body is white.<br />
TYPE-LOCALITY.—Angola, 10°36'S, 13° 12%<br />
ca 366 m {Undaunted Sta 111).<br />
DISPOSITION OF TYPES.—The holotype (Crust.<br />
D.31505) is a male, carapace width 79 mm, in the<br />
collection of the Rijksmuseum van Natuurlijke<br />
Historie, Leiden. The remainder of the specimens<br />
examined are paratypes, which have been deposited<br />
in the Rijksmuseum and in the Division of<br />
Crustacea, National Museum of Natural History,<br />
<strong>Smithsonian</strong> <strong>Institution</strong>.<br />
ETYMOLOGY.—The Latin name piperitus is given<br />
in allusion to the bright red and white color,<br />
resembling that of a peppermint cane.<br />
BIOLOGY.—The species has been reported from<br />
depths between 200 and 628 m, and has been<br />
taken most often between 300 and 450 m. The<br />
bottom on which it was found was described as<br />
mud (Guinot and Ribeiro, 1962; Forest, 1963),<br />
hard, dark gray mud (Voss, 1966), muddy sand<br />
(Bouvier, 1922), mud and sand, brown mud and<br />
sand, green mud and sand (Capart, 1951), sand<br />
and gravel (A. Milne Edwards and Bouvier, 1899,<br />
1900), corals (Guinot and Ribeiro, 1962), green<br />
muddy sand (Maurin, 1968b). A. Milne Edwards<br />
and Bouvier (1900) mentioned the bottom at<br />
Talisman Sta 113 as "Sable, roche," but in the<br />
official list of stations of the expedition it is<br />
indicated as "Sable. Gravier," exactly as in Sta<br />
110 of the same expedition.<br />
Ovigerous females of this species have been<br />
collected in February (Monod, 1956), March,<br />
April, and May (Capart, 1951), May and June<br />
(p. 77). Several specimens carry Lepadidae on the<br />
dorsal surface of the carapace and on the legs. A<br />
male from Geronimo Sta 198 carries a number of<br />
hydroid colonies on the merus of the last pereiopod.<br />
DISTRIBUTION.—Bathynectes piperitus is a <strong>West</strong><br />
<strong>African</strong> species, known from the Cape Verde<br />
Islands and the coast of Africa from Senegal to<br />
Angola. Records of B. superbus (= B. maravigna)<br />
from Spanish Sahara and Mauritania given by
NUMBER 306 83<br />
Maurin (1968a,b) and of Bathynectes given by<br />
Filhol (1885a) may be referable to this species.<br />
Records in the literature include the following:<br />
Cape Verde Islands: No specific locality (Filhol, 1885a).<br />
16°53'N, 27°3O'W of Paris [= 25°10'W of Greenwich] to<br />
16°54'N, 27°30'W of Paris [= 25°10'W of Greenwich], 410-<br />
460 m (A. Milne Edwards and Bouvier, 1899, 1900). 16°-<br />
52'N, 27°30'W of Paris [= 25°10'W of Greenwich] to 16°-<br />
52'N, 27°32'W of Paris [= 25°12'W of Greenwich], 550-760<br />
m (A. Milne Edwards and Bouvier, 1900). 15°14'N, 23°-<br />
03'45"W, 628 m (Bouvier, 1922).<br />
Senegal: Off Saint-Louis, 300 m (Maurin, 1968b). Fosse<br />
de Kayar, 85-660 m (Monod, 1956), 300-350 to 600 m<br />
(Maurin, 1968b). Off Pointe des Almadies, 150-245 m, 250-<br />
300 m, ±300 m, and 270-500 m (Monod, 1956).<br />
Guinea-Bissau: 10°05'N, 17°00'W, 320-360 m (Capart,<br />
1951).<br />
Guinea: 09°05'N, 15°10'W, 310-380 m (Monod, 1956).<br />
Liberia: 04°40'N, 09°20'W, 311-366 m (Voss, 1966;<br />
Lewis and Haefner, 1978).<br />
Ivory Coast: No specific locality (Intes and Le Loeuff,<br />
1976). 05°05'N, 04°00'W to 05°04'N, 04°02'W, 403-586 m<br />
(Voss, 1966). 04°32.5'N, 06°31'W, 300-455 m, and 04°54'N,<br />
03°23'W, 380-400 m (Forest, 1963).<br />
Ghana: 04°39'N, 02°46'W, 300-400 m (Forest, 1963). Off<br />
Accra, 200 m (Longhurst, 1958; Gauld, 1960).<br />
Gabon: Off Port-Gentil, 00°15'S, 08°47'E, 290-390 m<br />
(Capart, 1951). 0O°02.2'S, 08°50.2'E, 293 m (cited as Liberia);<br />
01°26.4'S, 08°24'E, 400 m (cited as 01°6.4'S, 08°-<br />
44'E, 396 m); 01°28'S, 08°24.5'E, 300 m (as 297 m); 02°01'S,<br />
08°50.5'E, 200 m; 02°31'S, 08°51'E, 300 m (as 288 m);<br />
04°38.4'S, ll°01.2'E, 400 m (all Lewis and Haefner, 1978).<br />
Cabinda: 05°23'S, 11°32'E, 290-350 m, and 05°39'S,<br />
11°25'E, 470 m (Capart, 1951).<br />
Angola: Off Ponta da Moita Seca, 06°08'S, 11°24'E, 350-<br />
380 m, and 06°23'S, 11°29'E, 400-430 m; off Ambrizete,<br />
07°16'S, 12°02'E, 380-420 m; off Ponta do Morro (as Cap<br />
Morro), 10°45'S, 13°O7'E, 400-500 m; off Egito [Praia],<br />
11°53'S, 13°28'E, 300 m, 11°53'S, 13°20'E, 480-510 m, and<br />
11°53'S, 12°23'E, 400-500 m (all Capart, 1951). Off Benguela,<br />
405-505 m; off Baia dos Tigres, 320-400 m (Guinot<br />
and Ribeiro, 1962).<br />
Genus Uocarcinus Stimpson, 1871<br />
Liocarcinus Stimpson, 1871a: 145 [type-species: Portunus holsatus<br />
Fabricius, 1798, by original designation; gender: masculine].<br />
DEFINITION.—Carapace broader than long,<br />
with 5 anterolateral teeth, lacking iridescent<br />
patches on surface. Front with 3 lobes, 7-10 irreg-<br />
ular teeth or tubercles, or entire. Dorsal margin<br />
of orbit with 2 closed incisions, ventral margin<br />
(Figure 18a) with narrow, V-shaped, incision.<br />
Basal antennal article touching front, scarcely or<br />
not at all movable. Chelipeds unequal, generally<br />
shorter than pereiopods; merus without distal<br />
ventral tooth; carpus with strong inner tooth,<br />
generally lacking outer tooth; dactylus (Figure<br />
18c) usually with 3 dorsal ridges. Propodus of<br />
second to fourth pereiopods each with 1 ventral<br />
ridge (Figure 18*,g). Third to fifth abdominal<br />
somites fused in male.<br />
REMARKS.—It is with some hesitation that we<br />
separate Liocarcinus, including most of the European<br />
species assigned to Macropipus for the last 20<br />
years or so, from Macropipus; but as Guinot (1961:<br />
7) pointed out, the species of Macropipus sensu<br />
stricto, restricted to three species, share certain<br />
features that serve to distinguish them readily<br />
from those species here assigned to Liocarcinus.<br />
Indeed, 50 years ago Palmer (1927, fig. 1) came<br />
to the conclusion that "Portunus" tuberculatus [the<br />
type-species of Macropipus] occupied an isolated<br />
position from the remainder of the British species<br />
of "Portunus" [here assigned to Liocarcinus]. Liocarcinus<br />
has not been used by European zoologists<br />
but was used by Bennett (1964:65) for L. corrugatus<br />
from New Zealand.<br />
Liocarcinus differs from Macropipus as follows:<br />
(1) The ventral margin of the orbit has one<br />
narrow, V-shaped incision (Figure 18a); in Macropipus<br />
(Figure 18b) the ventral margin is open and<br />
deeply U-shaped (see also Guinot and Ribeiro,<br />
1962, fig. 8). (2) Liocarcinus lacks iridescent patches<br />
on the carapace and pereiopods. (3) The ornamentation<br />
of the cheliped is different: Liocarcinus<br />
lacks a distal ventral spine on the merus, generally<br />
lacks a well-developed outer tooth on the carpus<br />
(such as in L. marmoreus and L. corrugatus), and<br />
usually has three rather than two longitudinal<br />
ridges on the dactylus (there may be only two in<br />
the highly ornamented L. corrugatus). (4) The<br />
propodi of the second to fourth pereiopods are<br />
ornamented with a single ventral ridge in Liocarcinus<br />
(Figure I8g), two distinct ridges in Macropipus<br />
(Figure \8h). (5) The walking legs are distinctly
84<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 18.—Comparison of morphological features. Liocarcinus holsatus (Fabricius), male, cb<br />
36.5 mm, England: a, orbit, ventral view; c, chela; e, propodus of second pereiopod, lateral<br />
view; g, propodus of second pereiopod, ventral view. Macropipus rugosus (Doflein), male, cb 36.7<br />
mm, Guinea-Bissau: b, orbit, ventral view; d, chela; /, propodus of second pereiopod, lateral<br />
view; h, propodus of second pereiopod, ventral view.<br />
longer in Macropipus (compare figure 26 (Macropipus<br />
tuberculatus) with figures 20-25 (Liocarcinus<br />
spp.) in Christiansen, 1969).<br />
Liocarcinus, Macropipus, and Bathynectes form a<br />
progression from generally shallow-water, shortlegged<br />
species to deep-water, long-legged species.<br />
One species of Liocarcinus, L. bolivari (Zariquiey<br />
Alvarez, 1968) apparently is endemic in the Mediterranean.<br />
Two species, L. holsatus (Fabricius,<br />
1798) and L. marmoreus (Leach, 1814), are not<br />
known to occur in the Mediterranean; records of<br />
these species in the latter area probably are referable<br />
to L. vernalis (Risso, 1816).<br />
As Christiansen (1969:60) pointed out, there<br />
has been some confusion regarding identity of<br />
Liocarcinus spp. off the northwest <strong>African</strong> coast,<br />
due to similarities between Xaiva mcleayi and L.<br />
pusillus.<br />
Eleven species of Liocarcinus occur in European-<br />
Mediterranean waters; Zariquiey Alvarez (1968:<br />
367) provided a key to the species then known.<br />
Liocarcinus arcuatus (Leach, 1814),<br />
new combination<br />
Portunus arcuatus.— Capart, 1951:115 [Mauritania].—Monod,<br />
1956:173 [references].<br />
Macropipus arcuatus.—Zariquiey Alvarez, 1968:369, figs.<br />
116d-h, 120a, 122c, 123a [Spain; references].—Christiansen,<br />
1969:57, fig. 21, map 15 [Scandinavia].<br />
SYNONYMS.—Portunus emarginatus Leach, 1814;<br />
Portunus guttatus Risso, 1816; Portunus rondeleti<br />
Risso, 1816; Portunus infractus Otto, 1828.<br />
DISTRIBUTION.—Eastern Atlantic from the<br />
North Sea and the British Isles to Mauritania,<br />
Mediterranean, in depths to 108 m, usually between<br />
10 and 50 m.<br />
Liocarcinus corrugatus (Pennant, 1777),<br />
new combination<br />
Portunus corrugatus.—Bouvier, 1911:226 [Mauritania].—Capart,<br />
1951:120 [Spanish Sahara].—Monod, 1956:174, fig.<br />
202 [Senegal, Guinea, Gabon, Angola; references].—<br />
Maurin, 1968b:484 [Spanish Sahara].<br />
Macropipus corrugatus.—Guinot and Ribeiro, 1962:31 [Cape<br />
Verde Islands, Angola].—Ribeiro, 1964:5 [Cape Verde<br />
Islands].—Forest and Guinot, 1966:59 [Senegal].—<br />
Maurin, 1968b:484 [Spanish Sahara].—Zariquiey Alvarez,<br />
1968:372, figs. 13e, 118c-e, 120c, 122d, 123e, 124c<br />
[Spain; references].—Bas, Arias, and Guerra, 1976, table<br />
3 [Spanish Sahara].—Tiirkay, 1976b:61 [listed], 66 [Madeira].
NUMBER 306 85<br />
SYNONYM.—Portunus leachi Risso, 1827.<br />
DISTRIBUTION.—Eastern Atlantic, from England<br />
to Senegal, Angola, including the Azores,<br />
Canary Islands and Cape Verde Islands, Mediterranean;<br />
also Indo-<strong>West</strong> Pacific (Stephenson,<br />
1972:23); littoral to 60 m.<br />
Genus Macropipus Prestandrea, 1833<br />
Macropipus Prestandrea, 1833:5 [type-species: Portunus macropipus<br />
Prestandrea, 1833, a subjective junior synonym of<br />
Portunus tuberculatus Roux, 1830, by tautonymy; gender:<br />
masculine; name 987 on Official List].<br />
Elliptodactylus Doflein, 1904:93 [type-species: Elliptodactylus<br />
rugosus Doflein, 1904, by monotypy; gender: masculine].<br />
DEFINITION.—Carapace broader than long,<br />
with 5 anterolateral teeth, iridescent patches present<br />
on surface and on pereiopods. Front with 3<br />
lobes. Dorsal margin of orbit with 2 closed incisions,<br />
ventral margin (Figure 186) deeply Ushaped.<br />
Basal antennal article touching front,<br />
usually freely movable. Chelipeds unequal,<br />
shorter than pereiopods 2 to 5; merus with distal<br />
ventral tooth; carpus with strong inner tooth and<br />
smaller outer tooth; dactylus (Figure ISd) with 2<br />
dorsal ridges. Propodus of second to fourth pereiopods<br />
each with 2 ventral ridges (Figure I8f,h).<br />
Third to fifth abdominal somites fused in male.<br />
REMARKS.—As noted above (p. 83), we are<br />
extremely reluctant to separate the genus Liocarcinus,<br />
from Macropipus sensu stricto. However, as<br />
Guinot (1961:7) pointed out in her account of the<br />
three species of Macropipus, "Ce trois especes ont<br />
en commun [in contrast with other species then<br />
assigned to Macropipus] certains caracteres, en particulier<br />
l'orntmentation de la face dorsale de la<br />
carapace et des appendices, la disposition des<br />
epines sur le chelipede, la forme du maxillipede<br />
externe, la presence de zones iridescentes nacrees<br />
sur la carapace et les appendices." These features,<br />
and those discussed under Liocarcinus, warrant the<br />
recognition of two genera: Liocarcinus, including<br />
11 relatively short-legged, shallow water species,<br />
and Macropipus, including 3 relatively longlegged,<br />
deep-water species from the Mediterranean<br />
and <strong>West</strong> Africa.<br />
Macropipus austral is Guinot, 1961<br />
Elliptodactylus rugosus.—Barnard, 1950:817 [not Elliptodactylus<br />
rugosus Doflein, 1904 = Macropipus rugosus].<br />
Portunus tuberculatus.—Capart, 1951:117, fig. 41 [part, specimens<br />
from Angola (Sta A.S. 110) and South-<strong>West</strong> Africa<br />
(Sta A.S. 108) only].—Barnard, 1954:123 [listed]; 1955:3<br />
[listed]. [Not Portunus tuberculatus Roux, 1830 = Macropipus<br />
tuberculatus.]<br />
Macropipus australis Guinot, 1961:5, figs. 1, 2, 5, pi. 1, pi. 2:<br />
fig. 1— Crosnier, 1970:1216.<br />
Macropipus sp.—Guinot and Ribeiro, 1962:44, 45, figs. 13-<br />
15, 24a,b, pi. 2: fig. 3, pi. 3: fig. 3.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Undaunted Material: Angola: Sta 94, 90 m, \6 (L). Sta 95,<br />
126 m, 56, 6? (W). Sta 96, 162 m, 1$ (L). Sta 103, 90 m,<br />
20(5, 6? (L). Sta 104, 126 m, 16, 1? (L). Sta 105, 155 m, 36<br />
(L).<br />
South-<strong>West</strong> Africa: Sta 106, 225 m, 1$ (L).<br />
DESCRIPTION.—Capart, 1951:117; Guinot,<br />
1961:5.<br />
Figures: Capart, 1951, fig. 41; Guinot, 1961,<br />
figs. 2, 5, pi. 1, pi. 2: fig. 1; Guinot and Ribeiro,<br />
1962, figs. 13-15, 24a,b, pi. 2: fig. 3, pi. 3: fig. 3.<br />
Male Pleopod: Guinot and Ribeiro, 1962, fig.<br />
24a,b (South-<strong>West</strong> Africa).<br />
Color: "Brick-red, under surface greyish; bright<br />
iridescent patches as follows: on antero-lateral<br />
margin between each pair of teeth, and a larger<br />
crescentic patch behind the last lateral tooth, in<br />
the smooth grooves between the regions of the<br />
carapace, anterior surface of the palp of mxp. 3,<br />
on the smooth distal portion of upper surface of<br />
arm of cheliped, the spine and other smooth areas<br />
on wrist, the smooth areas on upper surface of<br />
hand, and in the groove on upper margin of<br />
finger" (Barnard, 1950:818).<br />
BIOLOGY.—Macropipus australis is a sublittoral<br />
species living on the outer shelf and upper slope<br />
in depths between 90 and 238 m. Of ten depth<br />
records available, two are from less than 100 m<br />
(both 90 m), five are from between 100 and<br />
200 m (110, 126, 126, 155, and 162 m), and three<br />
are from depths greater than 200 m (220, 225,<br />
and 238 m). The only information on the nature<br />
of bottom on which the species has been found is<br />
given by Capart (1951); both of the lots he re-
86<br />
ported were taken on muddy sand in 110 and<br />
220 m. Ovigerous females have not been collected.<br />
DISTRIBUTION.—Off <strong>West</strong> Africa, from the<br />
coasts of Angola and South-<strong>West</strong> Africa, in<br />
depths between 90 and 238 m. Records in the<br />
literature include:<br />
Angola: 16°27'S, 11°35'E, 90 m (Crosnier, 1970). 18 mi<br />
[29 km] WSW of Baia dos Tigres, 16°36'S, 11°27'E, 110 m<br />
(Capart, 1951; Guinot, 1961). 16°37'S, 11°22'E, 126 m;<br />
16°41'S, 11°21'E, 162 m; 17°06'S, 11°35'E, 90 m; 17°09'S,<br />
ll°30'E, 126 m; 17°13'S, 11°27'E, 155 m (all Crosnier,<br />
1970).<br />
South-<strong>West</strong> Africa: 17°18'S, 11°24'E, 225 m (Crosnier,<br />
1970). 52 mi [84 km] SxW Fort Rock Point, 19°52'S, 12°-<br />
20'E, 220 m (Capart, 1951; Guinot, 1961). 26°36'S, 14°37'E,<br />
130 fm (238 m) (Barnard, 1950; Guinot, 1961; Guinot and<br />
Ribeiro, 1962).<br />
* Macropipus rugosus (Doflein, 1904)<br />
Portunus sp.—Leach, 1818, in 1817-1818:413.—Monod,<br />
1970:66.<br />
Elliptodoctylus rugosus Doflein, 1904:94, figs. 7, 8, pi. 30: figs.<br />
1-3, pi. 32: fig. 7.<br />
Portunus tuberculatus.—Capart, 1951:117 [part, not fig. 41, not<br />
specimens from A.S. 108, 110].—Monod, 1956:180, 632,<br />
figs. 207-209.—Rossignol, 1957:80, 123 [key].—Longhurst,<br />
1958:87.—Rossignol, 1962:115.—Maurin, 1968b:<br />
484, fig. 5.—Le Loeuff and Intes, 1968:44, table 1, figs.<br />
49, 63. [Not Portunus tuberculatus Roux, 1830.]<br />
Macropipus rugosus.—Guinot, 1961:2, 9, figs. 1, 4, 7, pi. 2: fig.<br />
3 [synonymy].—Guinot and Ribeiro, 1962:32-34, 35, figs.<br />
7-9, 17, 19, 20, 22a,b, pi. 2: fig. 1, pi. 3: fig. 1.—Crosnier,<br />
1964:34.—Forest and Guinot, 1966:60.—Le Loeuff and<br />
Intes, 1969:63, 65.—Turkay, 1976a:25 [listed], 37.<br />
Macropipus.—Voss, 1966:27.<br />
Portunus.—Maurin, 1968b, fig. 4.<br />
Portunus (Macropipus) tuberculatus.—Maurin, 1968b:486, 489,<br />
491 [not Portunus tuberculatus Roux, 1830].<br />
Macropipus tuberculatus.—Maurin, 1968b:491.—Bas, Arias<br />
and Guerra, 1976, table 3. [Not Portunus tuberculatus Roux,<br />
1830.]<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 68, 70 m, broken shell, 3 juv (W). Sta 83, 156-220 m, 2?<br />
ov (L).<br />
Ivory Coast: Sta 42, 62-75 m, mud with brown, branched<br />
Foraminifera, 1 juv (W). Sta 64, 68 m, 1 carapace (L). Sta<br />
65, 46-49 m, 19 ov (dry) (L).<br />
Ghana: Sta 23, 42 m, foliate brown to orange bryozoans,<br />
2$ (L). Sta 24, 35-37 m, dark red bryozoans, 1$ (W).<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Nigeria: Sta 241, 59-63 m, mud and shell, 56*, 69, 9 juv<br />
(W).<br />
Cameroon: Sta 259, 59 m, mud and broken shell, 96, 9?<br />
(L). Sta 260, 46 m, 16*, 19 (L).<br />
Gerommo Material: Gabon: Sta 202, 100 m, 366*, 30? (W).<br />
Sta 211, 100 m, 36*, 7° (W). Sta 213, 300 m, 16", 2$ (W). Sta<br />
228, 300 m, 16*, 29 (W).<br />
Other Material: Guinea-Bissau: 10°19'N, 16°34'W, 174 m,<br />
mud, sand, 16 May 1956, Calypso Sta 5, 16* (W).<br />
Liberia: Off Monrovia, 400 m, 27 Apr 1964, Guinean<br />
Trawling Survey 14/8, 1 specimen (L).<br />
Gulf of Guinea: Between Ghana and Sierra Leone, Feb<br />
and Apr 1964, Guinean Trawling Survey, 16* (L).<br />
DESCRIPTION.—Guinot, 1961, table 1 (comparison<br />
with M. tuberculatus (Roux, 1830) and M.<br />
australis Guinot, 1961; Guinot and Ribeiro, 1962:<br />
37-42 (comparison with M. tuberculatus).<br />
Figures: Monod, 1956, figs. 207-209.<br />
Male Pleopod: Monod, 1956, fig. 209 (Senegal);<br />
Guinot and Ribeiro, 1962, fig. 22a,b (Guinea-<br />
Bissau).<br />
Color: According to Capart (1951:118), this species<br />
is "gris-jaune avec taches roses, prenant un<br />
aspect nacre apres fixation." His color notes may<br />
have been based on M. australis, for part of his<br />
material is referable to that species (see p. 85).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 8 to 39 mm; ovigerous females<br />
have carapace widths of 32 to 39 mm.<br />
BIOLOGY.—Macropipus rugosus inhabits moderate<br />
depths and seems to prefer depths of less than<br />
100 m; the shallowest record is of 5 m and the<br />
deepest is 400 m. 78% of the records for which<br />
depth is given are from depths of less than 100 m,<br />
and, in those depths, records are equally divided<br />
between those in 50 m or less and those in 51-<br />
100 m. All but two of the Pillsbury specimens came<br />
from depths of less than 100 m and the species<br />
was found on bottoms with bryozoans, Foraminifera,<br />
broken shell, and mud and shell or mud<br />
and broken shell. Guinot (1961:10) noted that<br />
"Son habitat le plus commun semble etre le sable<br />
ou la vase." Forest and Guinot (1966) reported it<br />
from mud, rocks and calcareous algae, mud, mud<br />
and sand or broken shell, mud and calcareous<br />
algae and shells, sand, and algae and calcareous<br />
algae. Maurin (1968b) found it on mud in 40-
NUMBER 306 87<br />
60 m, on detrital mud in 50-90 and 200 m, on Mauritania: Bane d'Arguin, 40-60 and 90-100 m; off<br />
fine detrital muddy sand in 200-400 m, and on Tamzak (as Tamxat), 200-400 m (Maurin, 1968b).<br />
Senegal: Off Saint-Louis, 35-40 m; off Mboro, 35-40 m<br />
fluid mud with fine sand or mud and very muddy<br />
(Maurin, 1968b). 13°01'N, 17°24'W, 51-55 m, and<br />
fine sand in 35-40 m.<br />
12°55.5'N, 17°33'W, 65-75 m (Guinot and Ribeiro, 1962;<br />
Le Loeuffand Intes (1968:44) studied this spe- Forest and Guinot, 1966). SE of lie de la Madeleine, 48 m;<br />
cies (as Portunus tuberculatus) off the Ivory Coast S border, Fosse de Kayar, 300 m; off Camberene, 50 m (all<br />
and commented:<br />
Guinot and Ribeiro, 1962).<br />
Guinea-Bissau: 10°32'N, 16°53.5'W, 174 m, and 10°19'N,<br />
La zone des 50 m constitue l'habitat privilegie de l'espece 16°34'W, 60-73 m (Guinot and Ribeiro, 1962; Forest and<br />
qui accompagne cependant les eaux froides jusqu'a 35 m Guinot, 1966).<br />
quand elles remontent le long du plateau continental en Sierra Leone: No specific locality, 15-236 m (Longhurst,<br />
Mai, puis Aout-Septembre-Octobre et meme Janvier-Fev- 1958).<br />
rier. En fait P. tuberculatus vit dans des eaux bien determinees Liberia: 04°59'N, 09°37'W, to O4°57.5'N, 09°33'W, 156-<br />
(22°C-16°5C de temperature, 35,55%o a 35,75%o de salinite) 220 m (Voss, 1966).<br />
qui, en Cote d'lvoire correspondent a l'eau subtropicale. Ivory Coast: No specific locality (Le Loeuff and Intes,<br />
C'est sans doute, parmi tous les organismes benthiques lit- 1969). Off Grand-Lahou, off Grand-Bassam, 35-100 m (Le<br />
toraux de cette etude, l'espece la plus stenotherme et la plus Loeuff and Intes, 1968).<br />
infeodee a une masse d'eau.<br />
Cameroon: No specific locality, in more than 50 m (Crosnier,<br />
1964).<br />
Principe: 01°38'25"N, 07°22'05"E, 31 m (Guinot and<br />
Ribeiro, 1962; Forest and Guinot, 1966).<br />
Sao Tome: 00°25'40"N, 06°40'10"E, 50 m (Guinot and<br />
Ribeiro, 1962; Forest and Guinot, 1966).<br />
Gabon: 00°38'25"S, 08°46'E, 5 m; 00°38'20"S, 08°48'-<br />
30"E, 35 m (Guinot and Ribeiro, 1962; Forest and Guinot,<br />
1966). W of Pointe Panga, 70 m (Rossignol, 1962).<br />
Congo: Off Pointe-Noire (Rossignol, 1957).<br />
Zaire: 06°16'S, 12°07'E, 50 m; 06°28'S, 12°05'03 w E<br />
(Guinot, 1961).<br />
Angola: 10°S, 13°3O'E, 60-70 m (Guinot, 1961). Bai'a<br />
Farta, Benguela, 90 m; Baia da Caota, Benguela, 13 m (both<br />
Guinot and Ribeiro, 1962).<br />
Tvirkay's (1976a) record of this species from off<br />
Morocco seems questionable to us, not only because<br />
it is from so far north (33°37.5'N) but also<br />
because of the relatively great depth, 952-<br />
1038 m, at which his material is supposed to have<br />
been collected. That depth is more than twice as<br />
great as the deepest record otherwise observed for<br />
this species.<br />
Ovigerous females have been collected in June<br />
and September (Monod, 1956; Forest and<br />
Guinot, 1966; Pillsbury).<br />
DISTRIBUTION.—Off <strong>West</strong> Africa, from Mauritania<br />
to Angola, in moderate depths, from 5 to<br />
400 m. Monod (1956) identified this species with<br />
the Mediterranean M. tuberculatus (Roux) and<br />
reported material from Mauritania, Senegal,<br />
Sierra Leone, and Guinea. Guinot (1961) differentiated<br />
earlier records and provided synonymies<br />
for each of the three nominal species: M. australis,<br />
M. rugosus, and M. tuberculatus. Subsequent records<br />
include:<br />
<strong>West</strong> Africa: Gulf of Guinea (Leach, 1818, in 1817-1818;<br />
Monod, 1970).<br />
Morocco: 33°37.5'N, 09°02.2'W, 952-1038 m (?) (Turkay,<br />
1976a).<br />
Spanish Sahara: No specific locality (Guinot and Ribeiro,<br />
1962). Between Cabo Barbas and Cabo Blanco, 59-90 m;<br />
between Cabo Corbeiro and Cabo Blanco, 200 m (Maurin,<br />
1968b). 25°37.5'N, 14°52'W to 25°39.2'N, 14°51.5'W, 72-<br />
78 m, and 24°06.3'N, 16°20.2'W to 24°06'N, 16°24.8'W,<br />
61-68 m (Bas, Arias, and Guerra, 1976).<br />
Subfamily PORTUNINAE Rafinesque, 1815<br />
REMARKS.—This subfamily is characterized by<br />
the presence of a strong tooth on the upper, outer<br />
surface of the palm of the cheliped near the upper<br />
articulation of the carpus.<br />
Genus Callinectes Stimpson, 1860<br />
Callinectes Stimpson, 1860:220 [type-species: Callinectes sapidus<br />
Rathbun, 1896, by designation under the plenary powers<br />
of the International Commission on Zoological Nomenclature,<br />
Opinion 712 in Bulletin of Zoological Nomenclature,<br />
21(5):337, 1964; gender: masculine; name 1613 on Official<br />
List].<br />
REMARKS.—The genus Callinectes has been reviewed<br />
recently in detail by Williams (1974); not<br />
all of his distribution records are duplicated here.
88<br />
Key to <strong>West</strong> <strong>African</strong> Species of Callinectes*<br />
(modified from Williams, 1974)<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
1. Granules of carapace (Figure 19a) anterior to epibranchial ridges larger<br />
and placed wider apart than those posterior to ridges. Epibranchial<br />
ridges without distinct inflection in middle, almost straight. Submedian<br />
pair of frontal teeth well developed, usually half or more than half as<br />
long as lateral pair (measuring from base of lateral notch between teeth).<br />
First abdominal somite laterally ending in triangular, rather blunt point,<br />
not sharply pointed nor curved upward. Tips of male gonopods reaching<br />
beyond suture between thoracic sternite vi and mesially expanded<br />
sternite vn C. amnicola, new combination<br />
Granulation of carapace not different anterior and posterior to epibranchial<br />
ridges. Frontal teeth decidedly unequal in size, submedian pair no more<br />
than half as long as the lateral pair. Tips of male gonopods falling well<br />
short of suture between thoracic sternite vi and mesially expanded<br />
sternite VII 2<br />
2. Carapace (Figure 196) coarsely granulated. Epibranchial ridges with a<br />
distinct deflection in the middle. First abdominal somite ending laterally<br />
in a triangular point, which is neither sharply drawn out nor curved up.<br />
Male gonopods well separated from each other, not touching or<br />
crossed C. marginatus<br />
Carapace (Figure 19
NUMBER 306 89<br />
Other Material: Senegal: No specific data, 1 juv
90<br />
species, distinguishing it from other <strong>African</strong> species<br />
of the genus.<br />
An excellent account of the species has been<br />
given by Williams (1974:775).<br />
Figures: Rathbun, 1921, fig. 4, pi. 15: fig. 2, pi.<br />
21, pi. 22: fig. 1; Capart, 1951, fig. 47; Monod,<br />
1956, figs. 240-243; Williams, 1974, figs. 15, 19c,<br />
20p, 23a, 27.<br />
Male Pleopod: Rathbun, 1921, fig. 4e (Zaire);<br />
Williams, 1974, figs. 19c, 20p (Congo).<br />
Color: Williams (1974:776) summarized some<br />
earlier observations on color and noted that "a<br />
mottled olive coloration persists at least as long<br />
as 20 yr in some preserved specimens." Capart<br />
(1951:133) noted that in his material the color<br />
was "uniforme bleuatre (exemplaires fixes, orange<br />
clair)." Gauld (1960:69) observed that off Accra<br />
material of this species is "olive-brown with pale<br />
legs." De Rochebrune's (1883) color descriptions<br />
of the adult and juvenile are cited on p. 91.<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 15 to 126.5 mm; the ovigerous<br />
female has a carapace width of 120 mm.<br />
REMARKS.—In the recent literature this species<br />
has usually been given the name Callinectes latimanns<br />
Rathbun, 1897, but there is an older name<br />
(or actually two older names) available for it. De<br />
Rochebrune (1883:168-171) described three new<br />
species of swimming crabs from Senegal: Neptunus<br />
amnicola (p. 168), N. edwardsi (p. 169), and N.<br />
pallidus (p. 170). The identity of these species has<br />
puzzled many authors. Miers (1886:175) listed<br />
them in the subgenus Amphitrite de Haan, 1833<br />
[= Monomia Gistel, 1848] of the genus Neptunus de<br />
Haan, 1833 [= Portunus Weber, 1795], without<br />
any comment other than that amnicola perhaps<br />
should be placed in the subgenus Neptunus. Rathbun<br />
(1900a: 290) placed amnicola with some doubt<br />
in the synonymy of Callinectes bocourti A. Milne<br />
Edwards, and treated edwardsi (p. 289) and pallidus<br />
(p. 290) as good species of Portunus. Later<br />
Rathbun (1921:398) reassigned the <strong>West</strong> <strong>African</strong><br />
specimens that she at first had thought to be C.<br />
bocourti to C. latimanus, but at that occasion she<br />
did not deal with amnicola. Monod (1956:215)<br />
listed all three species of De Rochebrune as "spe-<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
cies inquirendae" under Callinectes and stated that<br />
he had not been able to locate De Rochebrune's<br />
type material in the Paris Museum. So far as we<br />
know all later authors ignored De Rochebrune's<br />
species.<br />
A study of De Rochebrune's descriptions makes<br />
clear that Monod was right in considering all to<br />
belong to the genus Callinectes. The species N.<br />
amnicola was described from a fullgrown specimen<br />
(cb 141 mm, cl 76 mm), the other two species<br />
after juveniles, which makes their identification<br />
especially difficult. Neptunus amnicola was described<br />
by De Rochebrune (1883:168-169) as<br />
follows (only the French description is cited and<br />
the abbreviated Latin diagnosis is omitted).<br />
Carapace beaucoup plus large que haute, faiblcment<br />
bombee, fortement granuleuse, a granulations plus grosses<br />
et plus espacees dans la region anterieure; lignes epigastriques<br />
et hypogastriques bien marquees, les epibranchiales<br />
peu indiquees; bords latero-anterieurs plus courts que les<br />
latero-posterieurs; premiere dent de forme trapezoidale, obtuse<br />
au sommet; les suivantes epaisses, brusquement terminees<br />
en pointe courte et aigue; la neuvieme tres longue,<br />
etroite, aceree, faiblement incurvee en dehors; front decoupe<br />
en six dents, les deux medianes petites et obtuses, les mitoyennes<br />
plus longues, subaigues, les externes tres larges courtes<br />
arrondies au sommet; apophyse epistomienne grosse, atteignant<br />
le niveau du front; pattes anterieures tres robustes,<br />
celles de droite plus volumineuses que celles de gauche; bras<br />
arme a la partie anterieure de trois dents en forme d'epine<br />
de Rosier, la premiere faible, les deux suivantes tres fortes;<br />
partie posterieure du bras aplatie, carenee, a carene obtusement<br />
denticulee et terminee en haut par une epine courte;<br />
avant-bras tricarene en dessous et en cote, chaque carene<br />
terminee par une dent obtuse; mains quadrangulaires, a<br />
angles fortement granuleux, armees en dessous d'une epine<br />
courte, en cote de deux tubercules et a Particulation de<br />
l'avant-bras, d'une dent triangulaire robuste et aigue.<br />
Teinte generale d'un beau bleu d'outre-mer nuance de<br />
rose, deux taches arrondies, rouges, de chaque cote et a la<br />
base de la carapace.<br />
This description agrees quite well with the<br />
present species, especially the very characteristic<br />
granulation of the carapace shows that Neptunus<br />
amnicola is a synonym of Callinectes latimanus. De<br />
Rochebrune's remark that the chelae are "armees<br />
en dessous d'une epine courte" evidently is a<br />
misprint for "armees en dessus. ..." The fact<br />
that Callinectes latimanus is a common species in
NUMBER 306 91<br />
fresh and brackish waters of Senegal makes the<br />
identification even more likely. Rathbun's<br />
(1900a) tentative identification of De Rochebrune's<br />
Neptunus amnicola with what she at that<br />
time considered C. bocourti A. Milne Edwards thus<br />
proves to have been correct. The epithet amnicola<br />
De Rochebrune, 1883, being older than latimanus<br />
Rathbun, 1897, has priority and thus must be<br />
used for the species.<br />
De Rochebrune's second species, Neptunus edwardsi,<br />
is based on a very small specimen (cb<br />
26 mm, cl 14 mm), much smaller than N. amnicola<br />
which measured 141 mm in carapace width and<br />
76 mm in carapace length. De Rochebrune's<br />
(1883:169-170) description of N. edwardsi runs as<br />
follows.<br />
Carapace plus large que haute, aplatie, subhexagonale,<br />
tres faiblement granuleuse a la partie anterieure; lignes<br />
epigastriques et hypogastriques a peine indiquees, lignes<br />
epibranchiales fortement accusees; bords latero-anterieurs<br />
plus longs que les latero-posterieurs; premiere dent droite<br />
epaisse, les suivantes tres obtuses a bords libres tres finement<br />
creneles; la neuvieme aigue un peu incurvee en haut, relativement<br />
courte; front decoupe en six dents, les deux<br />
medianes tres petites, les mitoyennes et les externes egales et<br />
obtuses; apophyse epistomienne courte, ne depassant pas le<br />
front; pattes anterieures faibles; bras arme sur le bord anterieur<br />
de trois epines, et de deux autres epines plus faibles<br />
a l'extremite articulaire; avant-bras portant sur la face externe<br />
une crete saillante limitee en dehors par une epine<br />
courte et accompagnee en dedans d'un tubercule obtus;<br />
mains anguleuses, a cretes longitudinales assez saillantes<br />
faiblement rugueuses, et armees de deux petites epines en<br />
dessus de l'articulation du pouce.<br />
Teinte generate d'un roux canelle marbre de bleuatre et<br />
de blanc.<br />
The description of Neptunus edwardsi fits juvenile<br />
specimens of Callinectes latimanus very well. In<br />
specimens of the size of that of Rochebrune's the<br />
carapace is smooth with a few granules in the<br />
area before the epibranchial ridge. The lateral<br />
spine is short, being only somewhat longer than<br />
the last anterolateral tooth, which gives the carapace<br />
indeed a "subhexagonal" shape. The lateral<br />
frontal and inner orbital teeth are rounded<br />
and similar, while the epistomial spine does not<br />
show in dorsal view. In juvenile Callinectes gladiator<br />
the lateral spines are well developed and long. It<br />
seems quite safe to consider Neptunus edwardsi<br />
De Rochebrune a synonym of N. amnicola De<br />
Rochebrune, being based only on a juvenile specimen<br />
of that species.<br />
De Rochebrune's third species, Neptunus pallidus,<br />
proves to belong to the species that is generally<br />
known as Callinectes gladiator, and as De<br />
Rochebrune's name is older it has to be known as<br />
Callinectes pallidus (De Rochebrune) (p. 95).<br />
The material that Irvine (1947:297) referred to<br />
Callinectes latimanus proves to belong to Portunus<br />
validus (p. 103); his figure 202, however, represents<br />
Callinectes pallidus (p. 95).<br />
The specimens recorded by Herklots (1851),<br />
De Man (1883), and Biittikofer (1890) are in the<br />
collections of the Rijksmuseum van Natuurlijke<br />
Historie in Leiden. Herklots' specimen, from Elmina<br />
(as St. George del Mina), Gold Coast<br />
(Ghana), is a male C. amnicola (Crust. D. 376). De<br />
Man's material included a specimen of C. amnicola<br />
from the Congo (Crust. D. 1863), and Biittikofer's<br />
material included a C. amnicola from Grand Cape<br />
Mount (Crust. D. 1873), as well as from Lake<br />
Piso (as Fisherman's Lake), Liberia (Crust. D.<br />
1864). Monod (1956:214) had listed the records<br />
by Herklots and De Man as unidentifiable.<br />
There is a syntype of C. latimanus Rathbun<br />
(USNM 19877), a female 52 X 114 mm, from<br />
Lagos, in the collection of the National Museum<br />
of Natural History, <strong>Smithsonian</strong> <strong>Institution</strong>.<br />
Pauly (1957:75) noted that in the Sakumo<br />
lagoon, Ghana, this species was common enough<br />
to support a subsistence trap fishery.<br />
BIOLOGY.—As pointed out by Williams (1974)<br />
and earlier authors, this species is primarily estuarine,<br />
although it is not restricted to estuaries.<br />
De Rochebrune (1883:171) commented on the<br />
"habitat exclusivement fluviatile" of this species,<br />
and said that it is common in the Senegal River<br />
and disappears when the water becomes salt, to<br />
reappear when it turns again completely fresh.<br />
Actually the species can stand brackish water and<br />
Monod (1956:214) classified it as occurring in<br />
marine, brackish, and freshwater habitats. Generally<br />
it is considered to be more characteristic of<br />
estuaries than C. pallidus, but in the Pillsbury
92<br />
collections from Lagos harbor, a brackish to fluviatile<br />
habitat, the latter species (55 specimens)<br />
far outnumbers C. amnicola (2 specimens).<br />
Le Loeuff and Intes (1968:44) made the following<br />
observations on this species:<br />
C'est un element important de la faune des milieux<br />
lagunaires de l'ouest Africain. Nous l'avons cependant<br />
recolte a 9 reprises en mer dans la zone littorale au-dessus de<br />
30 metres. Les individus etaient tous des femelles de grande<br />
taille, ovigeres a une exception pres. Ces collectes ont eu lieu<br />
de juillet a aout, puis de decembre a fevrier. Les deux sorties<br />
de decembre 1967 a Grand-Bassam ont permis encore de<br />
ramener onze femelles ovigeres. C. latimanus migre done en<br />
mer pour pondre; et il y aurait deux saisons de ponte, l'une<br />
en saison froide, apres la periode des fortes precipitations de<br />
Mai et Juin, l'autre au moment des petits upwellings, apres<br />
les crues des fleuves de Septembre et Octobre.<br />
In a subsequent paper these same authors commented<br />
(1969:63):<br />
Le Grabe de lagune Callinectes latimanus Rathbun vient<br />
egalement pondre en mer; des femelles de grande taille, en<br />
grande majorite ovigeres ont ete recoltees a plusieurs reprises<br />
sans qu'il soit possible encore d'affirmer que la sortie en mer<br />
et la ponte ont lieu a certaines periodes definies ou tout au<br />
long de l'annee.<br />
Ovigerous females have been collected in January,<br />
February, March, May, June, July, August,<br />
October, and December (Monod, 1956; Ribeiro,<br />
1964; Le Loeuff and Intes, 1968; Williams, 1974;<br />
Pilbbury).<br />
DISTRIBUTION.—Callinectes amnicola is an inshore,<br />
estuarine species, occurring off <strong>West</strong> Africa<br />
from Baie de Saint-Jean (19°27'N, 16°22'W),<br />
Mauritania to at least as far south as a lagoon N<br />
of Luanda (08°47'S, 13°16'E), Angola. Williams<br />
(1974) summarized earlier records; distributional<br />
records from the literature not in Williams include<br />
the following:<br />
<strong>West</strong> Africa: No specific locality (Monod, 1967).<br />
Cape Verde Islands: Baia [Porto] de Sal Rei, Boavista<br />
(Guinot and Ribeiro, 1962; Ribeiro, 1964). Porto da Praia,<br />
SaoTiago (Ribeiro, 1964).<br />
Senegal: Senegal River, Faleme River [?], Bakoy [?] [= ?<br />
Bako, 14°00'N, 15° 16^, or Bakor River 12°55'N, 14°-<br />
M'W], Bating [?] [= Baling ?] "et tous les Marigots tributaires,"<br />
Casamance River (all De Rochebrune, 1883).<br />
PresquTIe du Cap Vert; Rosso; Dakar (Sourie, 1954a).<br />
Dakar; Saloum (Sourie, 1957).<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Gambia: Gambia River (13°28'N, 16°34'W) (De Rochebrune,<br />
1883).<br />
Sierra Leone: Sierra Leone River (Longhurst, 1957).<br />
Liberia: No specific locality (Biittikofer, 1890; Johnston,<br />
1906).<br />
Ivory Coast: Off Sassandra, off Grand-Lahou, off Grand-<br />
Bassam, 15-30 m (Le Loeuff and Intes, 1968).<br />
Ghana: Off Accra, in shallow water (Gauld, 1960). Sakumo<br />
lagoon (Pauly, 1975).<br />
Nigeria: Lagos (Bruce-Chwatt and Fitz-John, 1951). Elechi<br />
Creek, Port Harcourt, 04°47'15"N, 06°58'45"E (Powell,<br />
1979).<br />
Cameroon: No specific locality, 0-30 m (Crosnier, 1964).<br />
* Callinectes marginatus<br />
(A. Milne Edwards, 1861)<br />
FIGURES 19b, 20a,b<br />
Neptunus diacanthus.—Pechiiel-Loesche, 1882:287.—Osorio,<br />
1892:199. [Not Portunus diacantha Latreille, 1825 = Callinectes<br />
sapidus Rathbun, 1896].<br />
PCallinectes diacanthus.—Gruvel, 1913:78, 106 [this species or<br />
C. amnicola?].<br />
PCallinectes diacanthus var. africanus.—Gruvel, 1913:168 [listed]<br />
[this species or C. amnicola ?].<br />
Callinectes marginatus.—Gruvel, 1913:168 [listed].—Capart,<br />
1951:134, fig. 48.—Monod, 1956:208, figs. 238, 239.—<br />
Rossignol, 1957:82.—Buchanan, 1958:20, 23.—Gauld,<br />
1960:69.—Guinot and Ribeiro, 1962:48.—Rossignol,<br />
1962:116.—Ribeiro, 1964:6.—Forest and Guinot, 1966:<br />
65.—Monod, 1967:180, pi. 15: fig. 6 [no material].—Bott,<br />
1968:169.—Williams, 1974:722 [part, not figs. 3, 18a,b,<br />
20a, 22b, 27] [review of genus].—Hartmann-Schroder and<br />
Hartmann, 1974:19.<br />
Callinectes.—Bayer, 1966:102.<br />
SYNONYM.—Callinectes diacanthus var. africanus A.<br />
Milne Edwards, 1879.<br />
MATERIAL EXAMINED.—Pillsbury Material: Nigeria:<br />
Sta 1, Lagos harbor, washed ashore, 1 carapace (L).<br />
Fernando Poo: Sta 257, shore, 26, 1 juv (L).<br />
Annobon: Sta 281, shore, 26, 2 juv (W).<br />
Other Material: Cape Verde Islands, Porto Grande, Sao<br />
Vicente, U.S. Eclipse Expedition, 1 juv (W). Porto da Praia,<br />
Sao Tiago (as La Praya, Santiago), Jul 1883, Talisman, 16*<br />
(W).<br />
Ghana: Mouth of Hwini River, Takoradi, 27 Jul 1961,<br />
Bane and Richards, Id (W). Virgins Pool, Takoradi, 27 Jul<br />
1961, Bane and Richards, 16" (W).<br />
Zaire: Banana, mouth of Congo River, American Museum<br />
Congo Expedition 1909-1915, Aug 1915, H. Lang, 56",<br />
49 (W). Data same, Jul-Aug 1915, 1? (W).
NUMBER 306 93<br />
Angola: Luanda (as St. Paul de Loanda), U.S. Eclipse<br />
Expedition, 26 (W).<br />
DESCRIPTION.—Carapace (Figure \9b) somewhat<br />
elevated, generally flatter than in C. amnicola,<br />
width slightly more (2.1 to 2.3 times) than<br />
twice as great as length (lateral spines included).<br />
Granulation dense, uniform, granules anterior to<br />
epibranchial ridge as dense and large as those<br />
posterior to ridge; granules finer than in C. amnicola<br />
but distinct. Epibranchial ridge between cervical<br />
groove and lateral spine of carapace not<br />
straight, with strong inflection near midlength.<br />
Length of metagastric area equal to or shorter<br />
than posterior width, latter half anterior width.<br />
Front with submedian teeth very narrow, short,<br />
much shorter than half length outer teeth; latter<br />
bluntly triangular, extending somewhat beyond<br />
more widely triangular inner orbital angles. Epistomial<br />
spine distinct in dorsal view, extending<br />
about as far forward as outer frontal teeth. Anterolateral<br />
margins of carapace more strongly<br />
convex than in C. amnicola. Anterior teeth triangular,<br />
curved somewhat forward, blunt; posterior<br />
teeth also triangular, more sharply pointed. Lateral<br />
spine stout, about twice as long as preceding<br />
tooth.<br />
Carinae on cheliped of adults distinct, granular.<br />
Chelae of adults slender, occasionally somewhat<br />
swollen in basal part of fingers, less strongly<br />
so than in C. amnicola. Basal tooth of dactylus of<br />
large chela slightly enlarged.<br />
First somite of abdomen in both sexes terminating<br />
laterally in triangular or blunt point, neither<br />
prolonged nor turned upward.<br />
First gonopods of male short, failing by far to<br />
reach end of sternite of third pereiopod. Distal<br />
part of gonopods somewhat curved externally,<br />
curve gradual, gonopods generally not touching<br />
(Figure 20a,b).<br />
Figures: Rathbun, 1921, fig. 2, pi. 19: fig. 1, pi.<br />
20: fig. 1; Capart, 1951, fig. 48; Monod, 1956,<br />
figs. 238, 239.<br />
Male Pleopod: Rathbun, 1921, fig. 2d (Zaire).<br />
Color: Capart (1951:135) noted that "coloration<br />
apres fixation non uniforme, des marbrures plus<br />
foncees sur la carapace et sur P.I." Rossignol<br />
(1957:82) remarked that the species can best be<br />
distinguished from C. amnicola by its color, namely<br />
"carapace marbree."<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 15 to 97 mm.<br />
REMARKS.—Since Rathbun (1896:358, 359) referred<br />
<strong>African</strong> specimens of this species to the<br />
East American Callinectes larvatus, the specific<br />
identity of <strong>African</strong> and American specimens of<br />
the two forms have not been questioned. After<br />
having examined the types of Callinectes marginatus<br />
(A. Milne Edwards, 1861) from Gabon, Callinectes<br />
larvatus Ordway, 1863, from Key <strong>West</strong>, the Tortugas,<br />
the Bahamas and Haiti, and Callinectes<br />
diacanthus africanus A. Milne Edwards, 1879, from<br />
the Cape Verde Islands, Rathbun (1897a: 149)<br />
synonymized the three species and used the oldest<br />
available name, marginatus, for it. Rathbun's views<br />
on the taxonomy and nomenclature of the species<br />
have since been followed by most authors, including<br />
Williams (1974) in his fundamental revision<br />
of the genus Callinectes.<br />
However, a comparison of our <strong>African</strong> material<br />
with American specimens identified with C. marginatus<br />
showed a number of seemingly constant<br />
differences, which in our opinion necessitates the<br />
recognition of two distinct species. The American<br />
species should be given the name Callinectes larvatus<br />
Ordway, 1863; the well-known name Callinectes<br />
marginatus (A. Milne Edwards, 1861) is retained<br />
for the <strong>African</strong> species. The differences<br />
between these species are the following:<br />
1. In C. marginatus (Figure 196) the granulation<br />
of the carapace before the epibranchial ridges is<br />
as dense as that behind these ridges, the granules<br />
there are of the same size as those in the posterior<br />
part of the carapace. In C. larvatus the granules<br />
before the epibranchial ridges are larger and<br />
placed much wider apart than those behind the<br />
ridges.<br />
2. The cervical groove behind the orbits is<br />
much deeper in C. larvatus than in C. marginatus.<br />
3. The ventral inner orbital angle in C. marginatus<br />
is narrower than in C. larvatus.<br />
4. The male gonopods are short in both species,<br />
but in C. marginatus (Figure 20a,b) they are more
94<br />
slender and the distal part that is curved outward<br />
is relatively longer and less abruptly bent; the<br />
two gonopods as a rule do not touch. In C. larvatus<br />
(Figure 20c,d) the distal outward curved part of<br />
the gonopod is shorter and more abruptly bent;<br />
the two gonopods touch each other in the midline<br />
of the body.<br />
BIOLOGY.—Monod (1956:210) indicated that<br />
this species is a shallow water form that prefers a<br />
sandy or sandy mud bottom, and is found in<br />
marine habitats. The latter in contrast to C.<br />
amnicola, which is known from fresh, brackish, and<br />
salt water, and C. pallidus, which Monod reported<br />
from salt and brackish waters. Buchanan (1958:<br />
20, 23) indicated C. marginatus as a characteristic<br />
epifaunistic species of the inshore fine sand community.<br />
Rossignol (1962:116) described the species<br />
as a "forme littorale (eaux saumatres, lagunes),"<br />
but possibly based this on the literature<br />
in which more than one species is confused under<br />
the name C. marginatus. Gauld (1960) mentioned<br />
the species to be caught intertidally. Guinot and<br />
Ribeiro (1962:48, 49) listed an abundant material,<br />
mostly from sandy beaches, either taken by<br />
hand or by beach seines; the species was mentioned<br />
once by them from the hull of a ship, and<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
once from under rocks near the shore; there is no<br />
indication that any of this material is estuarine.<br />
Ribeiro (1964:6) reported the species from "zona<br />
intercotidal." Forest and Guinot (1966:65) mentioned<br />
the species from a sand beach (0-9 m) and<br />
from intertidal rocks. <strong>West</strong> <strong>African</strong> material of<br />
Callinectes marginatus was listed by Williams (1974)<br />
from "rock pool" and "beach." The Pillsbury specimens<br />
were collected from sandy beaches (Sta 1<br />
and 281) and from the seashore (Sta 257). All<br />
these data confirm Monod's statement that the<br />
species is marine rather than estuarine. The data<br />
are too few, however, to definitely fix the habitat<br />
of the species; specimens have been reported from<br />
the mouth of the Congo River (Rathbun, 1921;<br />
and p. 92), and from the mouth of the Hwini<br />
River (p. 92), localities that might be estuarine.<br />
Off <strong>West</strong> Africa ovigerous females have been<br />
collected in February, May, June, September,<br />
October, and December (Guinot and Ribeiro,<br />
1962; Ribeiro, 1964; Forest and Guinot, 1966;<br />
Williams, 1974).<br />
DISTRIBUTION.—The species is known from the<br />
Cape Verde Islands and Port Etienne, Mauritania<br />
to central Angola. Monod (1956) summarized<br />
earlier records; distribution records from the<br />
literature not in Monod include the following.<br />
FIGURE 20.—Position and shape of gonopods: a,b, Callinectes marginatus (A. Milne Edwards),<br />
male, cb 90 mm, Congo; c,d, Callinectes larvatus Ordway, male, cb 116 mm, Jamaica.
NUMBER 306 95<br />
<strong>West</strong> Africa: No specific locality (Monod, 1967).<br />
Cape Verde Islands: Baia das Gatas, Sao Vicente (Guinot<br />
and Ribeiro, 1962; Ribeiro, 1964). Porto da Praia, SaoTiago<br />
(Guinot and Ribeiro, 1962; Ribeiro, 1964; Williams, 1974).<br />
Senegal: Dakar (Williams, 1974).<br />
Gambia: Near Gunjur (Williams, 1974).<br />
Guinea: Fotoba, 6 miles [10 km] W of Conakry (Williams,<br />
1974).<br />
Sierra Leone: Murray Town (Williams, 1974).<br />
Ghana: Off Accra (Buchanan, 1958). Apam (Gauld,<br />
1960).<br />
Togo: No specific locality (Gruvel, 1913).<br />
Dahomey: No specific locality (Gruvel, 1913).<br />
Nigeria: Lagos harbor (Williams, 1974).<br />
Fernando Poo: No specific locality (Williams, 1974).<br />
Principe: No specific locality (Forest and Guinot, 1966).<br />
Sao Tome: No specific locality (Forest and Guinot, 1966;<br />
Williams, 1974). Iogoiogo (Osorio, 1892; Forest and Guinot,<br />
1966). Morro Peixe (Forest and Guinot, 1966; Williams,<br />
1974).<br />
Annobon: 01°24'S, O5°37'E, shore (Bayer, 1966).<br />
Gabon: No specific locality (Williams, 1974; syntypes).<br />
Congo: No specific locality (Williams, 1974). Loango<br />
(Pechiiel-Loesche, 1882). Djeno, lagoon (Rossignol, 1957).<br />
Near Pointe-Noire (Rossignol, 1962; Williams, 1974).<br />
Zaire: Mouth of Congo River near Banana (Williams,<br />
1974).<br />
Angola: No specific locality (Forest and Guinot, 1966).<br />
Luanda (Forest and Guinot, 1966; Bott, 1968; Williams,<br />
1974). Baia do Lobito (Guinot and Ribeiro, 1962; Williams,<br />
1974). Praia da Rocha, Benguela (Guinot and Ribeiro,<br />
1962). Between Cacuaco and Lobito-Benguela (Hartmann-<br />
Schroder and Hartmann, 1974).<br />
* Callinectes pallidus (De Rochebrune, 1883),<br />
new combination<br />
FIGURE 19C<br />
Neptunus pallidus De Rochebrune, 1883:170.—Miers, 1886:<br />
175.—Balss, 1921:59 [listed].—Monod, 1956:215.<br />
Neptunus diacanthus.—De Man, 1883:150 [part].—Biittikofer,<br />
1890:466, 487 [part]. [Not Portunus diacantha Latreille, 1825<br />
= Callinectes sapidus Rathbun, 1896].<br />
Portunus pallidus.—Rathbun, 1900a:290.<br />
Callanectes Sp.?.—Irvine, 1932, fig. 13.<br />
Callinectes Gladiator.—Irvine, 1932:15.<br />
Callinectes lalimanus.—Irvine, 1947, fig. 202 [not C. latimanus<br />
Rathbun, 1897 = C. amnicola (De Rochebrune, 1883)].<br />
CallinecUs gladiator.—Irvine, 1947:298.—Frade, 1950:11,<br />
26.—Bruce-Chwatt and Fitz-John, 1951:117.—Capart,<br />
1951:130, fig. 46.—Monod, 1956:205, figs. 236, 237.—<br />
Rossignol, 1957:82.—Sourie, 1957:13, 51.—Longhurst,<br />
1958:87.—Gauld, 1960:69.—Guinot and Ribeiro, 1962:<br />
48.—Rossignol, 1962:116.—Crosnier, 1964:32.—Forest<br />
and Guinot, 1966:64.—Monod, 1967:180, pi. 15: fig. 3 [no<br />
material].—Bayer, 1966:98.—Le Loeuff and Intes, 1968:<br />
40, p. 2 of table 1, figs. 50, 61; 1969:64, 65.—Monod,<br />
1970:66.—Uschakov, 1970:439, 455 [listed].—Ejike, 1973:<br />
253.—Williams, 1974:735, figs. 5, 18c, 20b, 22c, 24 [review<br />
of genus].<br />
Callinectes.—Bayer, 1966:98.<br />
Lupa Smythiana.—Monod, 1970:66.<br />
SYNONYM.—Callinectes tumidus var. gladiator Benedict,<br />
1893.<br />
MATERIAL EXAMINED.—Pillsbury Material: Ivory<br />
Coast: Sta 43, surface, 29 (W).<br />
Nigeria: Sta 2, Lagos harbor, surface, 41 specimens (L,W).<br />
Sta 226: Lagos harbor, surface, 16* (W). Sta 228, Lagos<br />
harbor, surface, 16", 2$ (W). Sta 229, Lagos harbor, surface,<br />
46\ 6$ (W). Sta 250, 24 m, brackish water, mud, 9
96<br />
seum Congo Expedition, 1909-1915, Aug 1915, H. Lang,<br />
1
NUMBER 306<br />
the carapace very short, but in juveniles of the<br />
present species that spine is very elongate and<br />
straight, being much longer than the anterolateral<br />
teeth. The rest of the description also fits the<br />
present species, except for the measurements, for<br />
even C. amnicola specimens with cl 9 mm are much<br />
broader than 15 mm; one wonders whether<br />
15 mm for the width is not a typographical error<br />
for 25 or 19 mm. Because both C. amnicola and C.<br />
gladiator are estuarine species, and both are common<br />
in Senegal, it seems most likely that Neptunus<br />
pallidus belongs to one of these two species; the<br />
shape of the lateral spines and the granulation of<br />
the carapace makes clear that it can only be<br />
identified with C. gladiator Benedict, 1893. As<br />
Benedict's name gladiator was proposed 10 years<br />
after De Rochebrune's publication of pallidus the<br />
latter name has priority and has to be used for<br />
the species.<br />
As the type specimen of N. pallidus is no longer<br />
extant, we propose to fix the identity of Neptunus<br />
pallidus De Rochebrune, 1883, by selecting as its<br />
neotype the holotype of Callinectes tumidus gladiator<br />
Benedict, 1893 (USNM 14879), a male measuring<br />
29 X 63 mm (one lateral spine broken). In this<br />
way N. pallidus and C. gladiator become objective<br />
synonyms. The name pallidus is quite appropriate<br />
for the species, as preserved material usually is of<br />
a very pale color.<br />
The specimens reported as Neptunus diacanthus<br />
by De Man (1883) and Buttikofer (1890) are in<br />
the collection of the Rijksmuseum van Natuurlijke<br />
Historie, Leiden. Both author's accounts<br />
were based on material of C. amnicola as well as C.<br />
pallidus. De Man's material included two specimens<br />
of C. pallidus from the Congo (Crust. D. 374,<br />
1876), and Biittikofer's collection included one<br />
specimen of C. pallidus from Liberia (Crust. D.<br />
1871).<br />
As pointed out by Capart (1951:130, 132) the<br />
figures that Irvine (1932, 1947) published with<br />
his "Callinectes gladiator" and "Callinectes latimanus"<br />
have been interchanged: the figures indicated by<br />
Irvine in 1932 (fig. 13) as "Callanectes Sp.(?)" and<br />
in 1947 (fig. 202) as Callinectes latimanus actually<br />
represent C. pallidus. Irvine's text of "Callanectes<br />
Sp.(?)" and C. latimanus refers to Portunus validus<br />
(p. 103).<br />
BIOLOGY—Williams (1974:736) has summarized<br />
most of the available ecological information<br />
on this species, an inshore form usually found on<br />
mud, sand, or sand and shell in depths of less<br />
than about 30 m. It is less often encountered in<br />
estuaries than C. amnicola, although the majority<br />
of the Pillsbury specimens were taken in brackish<br />
water in Lagos harbor during outgoing tides.<br />
Monod (1956:208) indicated this species as being<br />
found in the sea and in brackish water.<br />
Sourie (1954b) found the species on bottoms of<br />
fine shelly sand with mud and Molgula hannensis<br />
Peres in 2-7 m in the Baie de Dakar; Uschakov<br />
(1970) found it in turbid water in depths of less<br />
than 20 m on unstable mud off Guinea. Gauld<br />
(1960) noted that off Ghana it was very common<br />
from low water mark to 18 m, usually on sand.<br />
Bruce-Chwatt and Fitz-John (1951) noted that in<br />
Nigeria it lives on muddy bottoms of creeks.<br />
Le Loeuff and Intes (1968:40) made the following<br />
observations on this species from their studies<br />
off the Ivory Coast:<br />
Extremement abondant et frequent au-dessus de 30<br />
metres, ce Callinectes est tres littoral car il ne descend pas<br />
au-dela de 35 m; son aire de repartition est tres stable: C.<br />
gladiator ne se deplace pas quelle que soit la saison et est done<br />
extremement tolerant aux variations de temperature, salinite,<br />
teneur en oxygene des eaux. CAPART (1951) a signale C.<br />
gladiator comme vivant sur des fonds vaseux. En fait l'espece<br />
est indifferente a la nature du substrat. Tous les sediments<br />
lui conviennent, sables ou vases. Sur les petits fonds C.<br />
gladiator domine nettement en nombre d'individus les autres<br />
Portunidae coders: Portunus inaequalis, Cronius ruber, Neptunus<br />
validus et contribue sans doute a repousser le maximum<br />
d'abondance des deux premiers au-dela de 25 metres et a<br />
limiter la densite de N. validus.<br />
Williams (1974) noted that earlier authors had<br />
commented on the aggressiveness of this species,<br />
a characteristic of most Portunidae.<br />
Ovigerous females have been recorded in January,<br />
February, March, April, May, June, October,<br />
and December (Capart, 1951; Forest and<br />
Guinot, 1966; Monod, 1956; Williams, 1974;<br />
Pillsbury).<br />
DISTRIBUTION.—<strong>West</strong> Africa, from numerous<br />
97
98<br />
localities between Baie de Saint-Jean (19°27'N,<br />
16°22 / W), Mauritania, to Bafa do Lobito (12°-<br />
20'S, 13°34'E), Angola, generally in depths of<br />
35 m or less. Williams (1974) reported material<br />
from numerous localities and summarized earlier<br />
records. Records not in Williams include the<br />
following.<br />
<strong>West</strong> Africa: No specific locality (Monod, 1970).<br />
Senegal: Lac de Nguer (as N'Guer), marigots de MBao<br />
[= Grand Mbao], Thionk [Pointe], Leybar (all De Rochebrune,<br />
1883). Anse de Hann, Baie de Dakar, 2-7 m (Sourie,<br />
1954b), Saloum (Sourie, 1957).<br />
Guinea: No specific locality, less than 20 m (Uschakov,<br />
1970).<br />
Sierra Leone: No specific locality, 14-19 m (Longhurst,<br />
1958).<br />
Liberia: No specific locality (Frade, 1950).<br />
Ghana: No specific locality, low water mark to 18 m<br />
(Gauld, 1960).<br />
Ivory Coast: No specific locality (Le Loeuff and Intes,<br />
1969). Off Sassandra, off Fresco, off Grand-Lahou, off<br />
Jacqueville, off Grand-Bassam, 15-35 m (Le Loeuff and<br />
Intes, 1968).<br />
Nigeria: Lagos (Bruce-Chwatt and Fitz-John, 1951). Lagos<br />
harbor, 06°28'N, 03°23'E, surface (Bayer, 1966). 7 miles<br />
[11 km] off Lagos harbor (Ejike, 1973).<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
MATERIAL EXAMINED.—Pillsbury Material: Ivory<br />
Coast: Sta 46, 38-42 m, mud with dense Jullienella, 1 juv<br />
(W). Sta 48, 22 m, 4o\ 39 (L, W).<br />
Annobon: Sta 275, 9-69 m, rubble of coralline algae, 2<br />
juv (L).<br />
Other Material: Ghana: Accra, 1868-1869, M. Sintenis, 1
NUMBER 306 99<br />
FIGURE 21.—Gonopods of two <strong>West</strong> <strong>African</strong> portunids. Cronius<br />
ruber (Lamarck), male, cb 72 mm, Pillsbury Sta 48: a,<br />
sternal view; b, apex, enlarged. Portunus validus Herklots,<br />
male, cb 151 mm, Pillsbury Sta 252: c, sternal view; d, apex,<br />
enlarged.<br />
imens in the color pattern and ornamentation of<br />
the chelae. In adults from <strong>West</strong> Africa the dark<br />
color of the movable finger extends proximally<br />
on the cutting edge, but is not expanded dorsally<br />
on the outer margin at the base of the finger. In<br />
specimens from Brazil, the dark color extends<br />
proximally as in the <strong>West</strong> <strong>African</strong> population,<br />
but at the base of the finger it expands dorsally<br />
so the finger appears to have a large, dark spot<br />
basally on the outer margin. This enlarged proximal<br />
spot is present in juveniles from both areas.<br />
The enlarged proximal spot is present in some<br />
eastern Pacific specimens, absent in others.<br />
The ornamentation of the propodus of the<br />
claw, which appears to be similar in American<br />
specimens from both coasts, differs significantly<br />
in the one large specimen from the Gulf of Guinea<br />
available to us (male, cb 72 mm, Pillsbury Sta 48).<br />
In American specimens the prominent longitudinal<br />
ridges on the propodus of the claw, especially<br />
the lower one on the outer face and the two<br />
ventral ones, are composed of short, transverse<br />
lines of tubercles or ridges which are inflated<br />
along the main axis of the longitudinal ridges so<br />
that the latter appear to be broad ridges with<br />
tuberculate edges. In our large specimen from<br />
<strong>West</strong> Africa the transverse lines of tubercules<br />
forming the main ridges are not inflated in the<br />
larger chela but are distinct throughout the<br />
length of the longitudinal ridge, as illustrated by<br />
Monod (1956, fig. 220), whereas in the smaller<br />
chela the ventral longitudinal ridges are ornamented<br />
with tubercles scattered in no apparent<br />
pattern, and there is no trace of the transverse<br />
lines of granules present on the major chela.<br />
In American specimens of C. ruber, the tubercles<br />
forming the lower, outer ridge on the chela also<br />
are inflated, so that the ridge appears smooth<br />
along its midline, with numerous marginal tubercles<br />
dorsally, fewer ventrally. In our <strong>West</strong> <strong>African</strong><br />
specimen, that ridge is made up of 2 distinct rows<br />
of tubercles with a space between them.<br />
Although the male pleopods of specimens from<br />
both sides of the Atlantic appear to be similar in<br />
shape and ornamentation, that of our largest<br />
specimen from <strong>West</strong> Africa is more curved laterally<br />
and has the apex directed anterolaterally<br />
rather than anteriorly. In available specimens<br />
from Brazil the pleopod is less strongly curved<br />
and the apex is directed anteriorly.<br />
BIOLOGY.—This is a shallow water species occurring<br />
on a variety of bottom types off <strong>West</strong><br />
Africa; it has been recorded from the intertidal<br />
zone to a depth of 69 (9-69) m, but the majority<br />
of depth records are from depths of 30 m or less.<br />
Sourie (1954a) characterized it as a sand dwelling<br />
species associated with the understone fauna of<br />
rocky shores; he also found it in the Dakar region<br />
(1954b) on coarse shelly sand, bottom with Area<br />
and Pyura, and noted that juveniles were found<br />
on fine shelly sand, with mud, bottom with Molgula<br />
hannensis Peres in 2-7 m. Off Guinea, Uschakov<br />
(1970) found it on the inferior mesolittoral<br />
part of rocky shores, associated with red algae<br />
and Padina. Buchanan (1958) characterized it as<br />
a member of the active epifauna, inshore fine<br />
sandy community, in 3-8 fm (5-15 m) off Ghana.<br />
Longhurst (1958) found it on shelly sand in 14-<br />
25 m off Sierra Leone. Le Loeuff and Intes (1968)<br />
reported that off the Ivory Coast it reached its<br />
maximum abundance at a depth of 35 m, but<br />
that on one transect in May, 465 individuals were<br />
taken at a depth of 50 m. They characterized the<br />
species as eurythermic and euryhaline and noted
100<br />
that it occurred on all types of bottom. Forest<br />
and Guinot (1966) reported several lots from the<br />
offshore islands of the Gulf of Guinea from the<br />
following habitats and depths: calcareous algae<br />
in 4-5 to 10-12 m (7 stations), calcareous algae<br />
and shells in 5-6 m, sand, algae, and calcareous<br />
algae in 8-30 m, mud, calcareous algae and shells<br />
in 31 m, and mud and sand in 35 m. The Pillsbury<br />
specimens were taken on mud bottom with Jullienella<br />
in 38-42 m and in the coralline algae habitat<br />
off Annobon in 9-69 m.<br />
Off <strong>West</strong> Africa ovigerous females have been<br />
recorded in January, March, and September (Capart,<br />
1951; Monod, 1956; Guinot and Ribeiro,<br />
1962; Ribeiro, 1964). Off <strong>West</strong> Africa, juveniles<br />
are more often found than adults; of more than<br />
80 lots recorded by Monod (1956) more than 50<br />
were made up of juveniles.<br />
DISTRIBUTION.—Atlantic-East Pacific. It is<br />
widely distributed in the eastern Pacific (Garth<br />
and Stephenson, 1966), as well as in the western<br />
Atlantic (Rathbun, 1930). Off <strong>West</strong> Africa it has<br />
been recorded from localities between Mauritania<br />
and Angola; shallow water to a depth of 69 m,<br />
usually between 10 and 30 m. Monod (1956)<br />
summarized earlier <strong>West</strong> <strong>African</strong> records and<br />
reported material from Mauritania (questionable<br />
data), Senegal, Guinea, Sierra Leone, Ivory<br />
Coast, Ghana, Gabon, and Angola; since 1956 it<br />
has been recorded from the following.<br />
<strong>West</strong> Africa: No specific locality (Monod, 1967).<br />
Cape Verde Islands: Porto da Furna, Brava, 6-20 m and<br />
Baia de Porto Grande, Sao Vicente, 4-6 m (Guinot and<br />
Ribeiro, 1962; Ribeiro, 1964).<br />
Guinea: Conakry (Uschakov, 1970).<br />
Sierra Leone: No specific locality, 14-25 m (Longhurst,<br />
1958).<br />
Ivory Coast: No specific locality (Le Loeuff and Intes,<br />
1969). O5°O5'N, 04°59.5'W, 22 m (Voss, 1966). Off Sassandra,<br />
off Fresco, off Grand-Lahou, and off Grand-Bassam,<br />
15-60 m (Le Loeuffand Intes, 1968).<br />
Ghana: Off Accra, 3-8 fm (5-15 m) (Buchanan, 1958).<br />
Off Accra, Winneba, and Tenkpobo (as Tenpobo), low water<br />
mark to 12 m (Gauld, 1960).<br />
Principe: O1°37'2O W N, 07°21'45"E, 35 m; 01°38'25"N,<br />
07°22'05"E, 31 m; between Ponta da Mina and Ilheu Santana,<br />
8-10 and 10-12 m; and in front of Praia Pequena, 5-<br />
6 m (Forest and Guinot, 1966).<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Sao Tome: 00°20'N, 06°46'E, 10 m; O0°25'15"N, 06°.<br />
43'05"E, 8-30 m; in front of Ponta Oquedelrei, 6 m; off<br />
Ponta Diogo Nunes, 4-5 m; off Sao Tome, 5 m (Forest and<br />
Guinot, 1966).<br />
Annobon: N of San Antonio, 9 m (Forest and Guinot,<br />
1966).<br />
Congo: Pointe-Indienne and Baie de Pointe-Noire (Rossignol,<br />
1957, 1962).<br />
Angola: Baia Farta, Benguela, 22-28 m, and Baia do<br />
Lobito, intertidal (Guinot and Ribeiro, 1962).<br />
Genus Portunus Weber, 1795<br />
Portunus Weber, 1795:93 [type-species: Cancer pelagicus Linnaeus,<br />
1758, by selection by Rathbun, 1926:75; see Opinion<br />
394, International Commission on Zoological Nomenclature,<br />
1956; gender: masculine; name 986 on Official<br />
List}.<br />
Portunus Fabricius, 1798:325, 363 [type-species: Cancer pelagicus<br />
Linnaeus, 1758, by selection by Latreille, 1810:94, 422;<br />
gender: masculine; name 410 on Official Index}.<br />
Lupa Leach, 1814:390 [type-species: Cancer pelagicus Linnaeus,<br />
1758, by monotypy; gender: feminine; name 411<br />
on Official Index}.<br />
Lima Leach, 1814:429 [possibly an erroneous spelling of<br />
Lupa; type-species: Cancer pelagicus Linnaeus, 1758, by<br />
present selection; gender: feminine; name 413 on Official<br />
Index}.<br />
Lupania Rafinesque, 1818:272 [substitute name for Lupa<br />
Leach, 1814; type-species: Cancer pelagicus Linnaeus, 1758;<br />
gender: feminine].<br />
Neptunus de Haan, 1833:3, 7 [type-species: Cancer pelagicus<br />
Linnaeus, 1758, by selection by Miers, 1886:172; gender:<br />
masculine; name 414 on Official Index}.<br />
Achelous de Haan, 1833:3, 8 [type-species: Portunus spinimanus<br />
Latreille, 1819, by monotypy; gender: masculine].<br />
Amphitrite de Haan, 1833:3, 8 [invalid junior homonym of<br />
Amphitrite O. F. Miiller, 1771 (Polychaeta); type-species:<br />
Portunus gladiator Fabricius, 1798, by selection by Miers,<br />
1886:172; gender: feminine].<br />
Pontus de Haan, 1833:3, 9 [type-species: Portunus (Pontus)<br />
convexus de Haan, 1833, by monotypy; gender: masculine].<br />
Lupea H. Milne Edwards, 1834:445 [erroneous spelling of<br />
Lupa Leach, 1814].<br />
Monomia Gistel, 1848:viii [substitute name for Amphitrite de<br />
Haan, 1833; type-species: Portunus gladiator Fabricius,<br />
1798; gender: feminine].<br />
Posidon Herklots, 1851:3 [invalid junior homonym of Posidon<br />
Illiger, 1801 (Crustacea); type-species: Portunus (Posidon)<br />
validus Herklots, 1851, by monotypy; gender: masculine].<br />
Xiphontctes A. Milne Edwards, 1873b: 157 [type-species: Amphitrite<br />
vigilans Dana, 1852, by selection by Rathbun, 1930:<br />
33; gender: masculine].
NUMBER 306 101<br />
Hellenus A. Milne Edwards, 1874, in 1873-1881:210, 221<br />
[type-species: Achelous spinicarpus Stimpson, 1871, by selection<br />
by Rathbun, 1930:33; gender: masculine].<br />
Lupocycloporus Alcock, 1899a:31, 32, 44 [type-species: Achelous<br />
whitei A. Milne Edwards, 1861, by monotypy; gender:<br />
masculine].<br />
Cycloachelous Ward, 1942:79 [type-species: Lupea granulata H.<br />
Milne Edwards, 1834, by monotypy; gender: masculine].<br />
Portunus hastatus (Linnaeus, 1767)<br />
Lupa hastata.—Barrois, 1888:14.<br />
Neptunus (Amphitrite) hastatus.—Lenz and Strunck, 1914:278.<br />
Neptunus hastatus.—Capart, 1951:125, fig. 44 [part, not specimen<br />
from Guinea].—Chapman and Santler, 1955:374.—<br />
Monod, 1956:203, figs. 232-235 [no material].<br />
Portunus hastatus.—Figueira, 1960:8.—Zariquiey Alvarez,<br />
1968:384, figs. 125d,e, 126c, 128a,b [Spain; references].—<br />
Turkay, 1976b:61 [listed], 64, pi. 1: figs. 1, 2.<br />
SYNONYMS.—Cancer ponticus Herbst, 1790; Portunus<br />
Dufourii Latreille, 1819; Eriphia prismaticus<br />
Risso, 1827; Neptunus hastatus rubromaculatus Steinitz,<br />
1932.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Madeira: Canical, 30 fm (55 m), seine, 18<br />
Apr 1916, A. C. de Noronha, 16* (W). Machico Bay, seine,<br />
16 Apr 1916, A. C. de Noronha, lcJ (W). Same data, 30 Jun<br />
1916, 2$ (W). Same data, 5 Nov 1921, 126\ 11? (6 ov) (W).<br />
Angola: 8 mi [13 km] W of Rio Cuanza, 09°20'S, 13°04'E,<br />
20-22 m, muddy sand, 31 Jan 1949, Expedition Oceanographique<br />
Beige, Sta A. S. 116, lcJ, 1$ ov (Brussels).<br />
DESCRIPTION.—Capart, 1951:126.<br />
Figure: Capart, 1951, fig. 44.<br />
Male Pleopod: Monod, 1956, figs. 232-235 (Lebanon)<br />
.<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 18 to 45 mm; the carapace widths<br />
of the ovigerous females from Madeira range<br />
from 18 to 26.5 mm, that from Angola is 42 mm.<br />
REMARKS.—In large males, the abdomen is narrower<br />
than in P. inaequalis; the last somite is<br />
triangular in P. hastatus, ovate in P. inaequalis. In<br />
P. hastatus the width of the terminal somite of the<br />
male abdomen is half or more than half the width<br />
of the preceding somite; in P. inaequalis it is less<br />
than half the width of the preceding somite.<br />
Finally, the ventral incision of the orbit is a closed<br />
V in P. hastatus, an open V or U in P. inaequalis, as<br />
noted by Monod (1956:203).<br />
The two specimens from Angola are those reported<br />
by Capart (1951), who correctly identified<br />
them with Portunus hastatus. We have compared<br />
these specimens with material of this species from<br />
several Mediterranean localities, as well as with<br />
the material from Madeira reported above, and<br />
we could find no constant differences in specimens<br />
from different areas. The specimens from<br />
Angola at first sight appear to be different from<br />
those from the Mediterranean, for, as figured by<br />
Capart (1951, fig. 44), the dorsal sculpturing of<br />
the carapace is much less pronounced on them.<br />
In addition, the lateral spine of the carapace is<br />
shorter than in most specimens of P. hastatus from<br />
the Mediterranean we have examined, and the<br />
chelae appear stouter with a slightly shorter movable<br />
finger.<br />
There are relatively few records for this species<br />
outside of the Mediterranean and we have tried<br />
to compile all of them here. One record, that of<br />
Miers (1886:175), as Neptunus (Amphitrite) hastatus,<br />
from Tenerife or Isla de la Gomera, Canary Islands,<br />
appears to be based on specimens of Portunus<br />
sayi. Miers noted that his specimens were<br />
"very prettily mottled with purple on a yellowish<br />
ground" (this is the color of P. sayi as described<br />
by Rathbun, 1930:42) and that the same jar<br />
contained specimens of Planes minutus. Miers' was<br />
the only record for this species from the Canary<br />
Islands that we could find.<br />
BIOLOGY.—Portunus hastatus is a sublittoral species,<br />
occurring on sandy bottoms subtidally to a<br />
depth of at least 55 m. The two specimens from<br />
Angola were taken on muddy sand in 20-22 m.<br />
Off <strong>West</strong> Africa ovigerous females have been<br />
recorded in January (Angola) and November<br />
(Madeira).<br />
DISTRIBUTION.—Eastern Atlantic, where it occurs<br />
primarily in the Mediterranean. Outside of<br />
that sea it has been recorded from the following:<br />
Azores: Ponta Delgada, Uha de Sao Miguel (Barrois,<br />
1888; Lenz and Strunck, 1914). Ilha do Faial (as Fayal)<br />
(Chapman and Santler, 1955). Ilha do Pico (Figueira, 1960).
102<br />
Madeira: Funchal, from fish market, Prainha[?], and<br />
Ponta de Sao Lourenco, 15-20 m (all Tiirkay, 1976b).<br />
Angola:8mi[13km]WofRioCuanza,09°20'S, 13°O4'E,<br />
20-22 m (Capart, 1951).<br />
* Portunus inaequalis (Miers, 1881)<br />
Neptunus hastatus.—Capart, 1951:125 [part, not fig. 44].—<br />
Rossignol, 1957:123 [key]. [Not Portunus hastatus (Linnaeus,<br />
1767).]<br />
Neptunus inaequalis.—Monod, 1956:198, figs. 225-231.—Buchanan,<br />
1958:24.—Longhurst, 1958:87.—Gauld, 1960:<br />
69.—Rossignol, 1962:116.—Le Loeuff and Intes, 1968:40,<br />
44, table 1, figs. 49, 61; 1969:63, 65.—Tiirkay, 1976b:61<br />
[listed] 64, pi. 1: figs. 3,4.<br />
Portunus inaequalis.—Guinot and Ribeiro, 1962:47.—Ribeiro,<br />
1964:6.—Forest and Guinot, 1966:63.<br />
MATERIAL EXAMINED.—Pillsbury Material: Ivory<br />
Coast: Sta 47, 37 m, bottom with Jullienella, 126*, 7$, 3 juv<br />
(W).<br />
Ghana: Sta 24, 35-37 m, dark red bryozoans, 1 juv (L).<br />
Sta 26, 27 m, shell bottom (scallops), 36*, 19 ov (L). Sta 27,<br />
33 m, 1$ (L).<br />
Nigeria: Sta 246, 37 m, lcJ (L). Sta 248, 33 m, Id, 1$ ov<br />
(L). Sta 250, 24 m, brackish water, mud, 16*, 19 (L). Sta 252,<br />
30 m, mud, Id, 19 (L).<br />
Other Material: Ghana: Chorkor, near Accra, Dec 1950, R.<br />
Bassindale, Id (L).<br />
DESCRIPTION.—Carapace more than twice as<br />
broad as long, regions distinct, surface roughened.<br />
Gastric region with long patch of raised granules,<br />
with curved branches extending onto protogastric<br />
region. Metagastric region with 2 raised submedian<br />
lobes ornamented with enlarged tubercles.<br />
Cardiac region with 2 raised submedian lobes,<br />
often united posteriorly in form of V- Epibranchial<br />
ridges raised, granular, sinuous. Mesobranchial<br />
region with raised prominence mesially,<br />
flanked laterally by 1 or more oblique granular<br />
ridges. Hepatic region with curved, granular<br />
ridges. Frontal teeth triangular, submedians<br />
smallest, sharpest, second pair larger and more<br />
triangular than blunt inner orbital spines. Interantennular<br />
spine small, visible in dorsal view.<br />
Anterolateral margin with 9 sharp, separate<br />
spines, outer orbital (first anterolateral) blunter<br />
than next spine; lateral spine long, at least as long<br />
as space occupied by preceding 4 or 5 spines,<br />
slender, curved upward and forward. Small patch<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
of iridescence present between bases of spines.<br />
Posterolateral margins unarmed. Chelae long,<br />
slender, merus longer than carapace, subequal.<br />
Dorsal surface of merus irregular, anterior margin<br />
with 4 or 5 spines, occasionally 1 smaller spine<br />
distally, posterior and ventral margin each with<br />
1 distal spine. Carpus with inner and outer spine.<br />
Propodus with 1 spine proximally and 2 spines<br />
distally on upper margin. Fingers slender, shorter<br />
than palm. Pereiopods 2-5 with iridescent lines.<br />
Merus of fifth leg unarmed. Sternum and abdomen<br />
smooth.<br />
Figures: Monod, 1956, figs. 225-231.<br />
Male Pleopod: Monod, 1956, figs. 227-231 (Senegal,<br />
Sierra Leone).<br />
Color: The following observations on color were<br />
provided by Monod in his list of material of this<br />
species:<br />
Tres marbres; en alcool une tache rouge plus ou moins<br />
marquee sur le dactyle p5 [1956:200, Senegal] . . . ; rouge<br />
brique; tache carminee sur la partie posterieure du dactyle<br />
p5; ligne carmin sur la crete inf. du propode pi, se prolongeant<br />
sur le doigt fixe; face interne du dactyle egalement<br />
carmin [1956:202, Guinea] . . . ; traces de tache rouge au<br />
dactyle p5 [1956:202, Ghana].<br />
The dark red color on the dactylus of the last leg<br />
also is characteristic of P. hastatus (Linnaeus).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 16 to 53 mm; the measurable<br />
ovigerous female has a carapace width of 25 mm.<br />
BIOLOGY.—Portunus inaequalis is a shallow water<br />
species, living near shore to a depth of 60-73 m,<br />
generally in depths of less than 40 m. It can<br />
tolerate waters of reduced salinity. The Pillsbury<br />
took it in brackish water on mud in 24 m off<br />
Nigeria, and Longhurst (1958), who reported it<br />
from Sierra Leone, characterized it as an estuarine-shelf<br />
species; he found it on stones, sand, and<br />
shelly sand in 10-25 m. Sourie (1954b) reported<br />
it from fine shelly sand with mud, bottom with<br />
Molgula hannensis Peres, in 2-7 m in the Baie de<br />
Dakar. Buchanan (1958) found it in the sandy<br />
silt community in 8-20 fm (15-37 m) off Ghana.<br />
Guinot and Ribeiro (1962) reported it from algae<br />
and rocks and on sand in 10 m. It was reported<br />
from a variety of bottom types by Forest and
NUMBER 306 103<br />
Guinot (1966): shell in 20-25 and 27 m; mud,<br />
shells and Cidaris in 60-73 m; sand or muddy<br />
sand and Foraminifera in 5 and 21-27 m; calcareous<br />
algae in 5, 6, and 12 m; rocks and coral in<br />
3-10 m; sand and calcareous algae in 4-5 m; and<br />
from a sand beach, 0-4 m.<br />
Le Loeuff and Intes (1968:44) found that off<br />
the Ivory Coast the species lived on sand or<br />
muddy sand and was rarely observed on mud.<br />
There it generally lives at depths between 30 and<br />
35 m, not going below 40 m, although during<br />
periods of upwelling or cooling part of the population<br />
was found at 15-20 m; in warm periods it<br />
generally was found below 25 m.<br />
Only two of the Pillsbury samples were taken at<br />
stations on mud; the species was found on shell<br />
bottom with scallops, on bottom with dark red<br />
bryozoans, and bottom with Jullienella.<br />
Ovigerous females have been collected in<br />
March, April, May, June, July, September, and<br />
November (Monod, 1956; Guinot and Ribeiro,<br />
1962; Ribeiro, 1964; Forest and Guinot, 1966;<br />
Pillsbury).<br />
DISTRIBUTION.—Off <strong>West</strong> Africa, from Madeira<br />
and from Senegal to Angola, in shallow<br />
water, from shore to a depth of 60-73 m, generally<br />
in less than 40 m. Monod (1956) recorded material<br />
from the Cape Verde Islands, Senegal,<br />
Guinea, Sierra Leone, Ghana, Gabon, and Principe.<br />
Since 1956 it has been recorded from the<br />
following localities.<br />
Madeira: No specific locality (Tiirkay, 1976b).<br />
Cape Verde Islands: Baia de Porto Grande and Baia de<br />
Calheta, 8-16 m, Sao Vicente; Baia da Faja di Agua (? =<br />
Porto da Faja) and Porto da Furna, 4 m, Brava; Tarrafal<br />
and Porto da Praia, Sao Tiago; and Tarrafal, Sao Nicolau<br />
(all Guinot and Ribeiro, 1962; Ribeiro, 1964).<br />
Guinea-Bissau: 10°19'N, 16°34'W, 60-73 m (Forest and<br />
Guinot, 1966).<br />
Sierra Leone: No specific locality, 10-25 m (Longhurst,<br />
1958).<br />
Ivory Coast: No specific locality (Le Loeuff and Intes,<br />
1969). 05°00'N, 05°28.5'W, 27 m; 05°03'N, 05°25"W, 20-25<br />
m; and 05°02.5'N, 05°25'W, 21-27 m (all Forest and Guinot,<br />
1966). Off Sassandra, off Fresco, off Grand-Lahou, off<br />
Jacqueville, and off Grand-Bassam, 8-40 m (Le Loeuff and<br />
Intes, 1968).<br />
Ghana: Off Accra, 8-20 fm (15-37 m) (Buchanan, 1958);<br />
10-32 m (Gauld, 1960).<br />
Principe: Between Uheu Santana and Ponta Capitao, 12<br />
m (Forest and Guinot, 1966).<br />
Sao Tome: Baia de Ana de Chaves; Praia de Santa<br />
Catarina, W coast, 3-10 m; in front of Ponta Oquedelrei, 6<br />
m; Morro Peixe, 0-4 m; Ilheu das Cabras, 4-5 m; off Sao<br />
Tome, 5 m (all Forest and Guinot, 1966).<br />
Annobon: 01°26'20"S, 05°36'25"E, 22-25 m (Forest and<br />
Guinot, 1966).<br />
Congo: W of Pointe-Noire (Rossignol, 1962).<br />
Gabon: 00°38'25"S, 08°46'E, 5 m (Forest and Guinot,<br />
1966).<br />
Angola: Baia de Caota, Benguela, 10 m (Guinot and<br />
Ribeiro, 1962).<br />
*Portunus validus Herklots, 1851<br />
FIGURES 2lc,d, 22, 23a,c,e,g<br />
Lupa Cranchiana White, 1847a:27 [nomen nudum; under<br />
Neptunus sanguinolentus].—Monod, 1970:66, 72.<br />
Neptunus validus.—Pechiiel-Loesche, 1882:287.—Bvittikofer,<br />
1890:466, 487, fig. on p. 465.—Johnston, 1906:862.—<br />
Frade, 1950:11, 26.—Capart, 1951:123, fig. 43.—Monod,<br />
1956:1%, fig. 224.—Rossignol, 1957:80, 123 [key], pi. 2:<br />
fig. 6.—Buchanan, 1958:20.—Longhurst, 1958:87.—<br />
Gauld, 1960:69.—Rossignol, 1962:115.—Crosnier, 1964:<br />
32, 87, 90, 92, 98, 105, 106, 110, 112, 120, 121.—Crosnier<br />
and Berritt, 1966:68, 100, 101, 102, 109, 123, 127, 131,<br />
132, 136.—Le Loeuff and Intes, 1968:40, 46, table 1, figs.<br />
49,61; 1969:64, 65.<br />
Callanectes sp.?.—Irvine, 1932:14.<br />
Callinectes Gladiator.—Irvine, 1932: fig. 9 [not C. gladiator<br />
Benedict, 1893 = C. pallidus (De Rochebrune, 1883)].<br />
Callinectes gladiator.—Irvine, 1947: fig. 203 [not C. gladiator<br />
Benedict, 1893 = C. pallidus (De Rochebrune, 1883)].<br />
Callinectes latimanus.—Irvine, 1947:297 [not Callineclus latimanus<br />
Rathbun — C. ammcola (De Rochebrune, 1883)].<br />
Portunus validus.—Monod, 1967:180, pi. 15: fig. 1 [no locality];<br />
1970:66, 72.—Uschakov, 1970:439, 455 [listed].—<br />
Baron, 1975a:3-13 [physiology]; 1975b: 103, figs. 1-4.<br />
MATERIAL EXAMINED.—Pillsbury Material: Nigeria:<br />
Sta 241, 59-63 m, mud and shell, 19 (L). Sta 252, 30 m, 26,<br />
2$(W).<br />
Other Material: Liberia: Off St. Paul River, Monrovia, 3-<br />
6 fm (5-11 m), 4 Mar 1953, G. C. Miller, lcj (W). Liberia,<br />
1881, J. Biittikofer, 2$ (L).<br />
Ghana: Elmina (as St. George-del-Mina), 1840-1855,<br />
H. S. Pel, lectotype, l
104<br />
1964, B. de Wilde-Duyfjes, ld\ 29 (L). Kribi, beach seine, 9<br />
Mar 1964, B. de Wilde-Duyfjes, 1$ (L). Same, 10 Aug 1964,<br />
B. de Wilde-Duyfjes, 1$ (L).<br />
Angola: No specific locality, 1879, P. Kamerman, 2$ (L).<br />
DESCRIPTION.—A. Milne Edwards, 1861:321;<br />
Capart, 1951, fig. 124.<br />
Figures: A. Milne Edwards, 1861, pi. 29: fig. 1;<br />
Capart, 1951, fig. 43; Monod, 1956, fig. 224.<br />
Color: The carapace is rather uniformly greenish<br />
gray with some brown tinges. A large conspicuous,<br />
white triangular spot is present on each side<br />
just before the middle of the posterolateral margin.<br />
This white spot is surrounded by a ring of<br />
darker gray than is on the rest of the carapace.<br />
The posterior margin also is white. The uniform<br />
color of the carapace contrasts strongly with the<br />
brightly marbled upper surface of the chelipeds<br />
(Figure 22). The upper half of each cheliped is<br />
very dark purple with numerous rather large<br />
white spots, which at some places are confluent.<br />
The lower half of the cheliped is white. The same<br />
marbled purple color is present on the upper half<br />
of the merus and carpus of the next 3 legs, the<br />
purple color and size of the white spots being<br />
very variable. The propodus and dactylus are<br />
greenish gray as is the carapace. The fifth leg has<br />
the upper surface of all segments except the dactylus<br />
purplish to greenish with white spots. The<br />
dactylus is gray dorsally with a white streak along<br />
FIGURE 22.—Portunus validus Herklots, female,<br />
Pillsbury St&2U.<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
the basal half of the outer margin. The lower<br />
surface of the body and legs is uniformly pale.<br />
Irvine (1947:297, under Callinectes latimanus) described<br />
the color of this species as follows: The<br />
carapace "is khaki in colour, with a bluish tinge.<br />
The claws are blue. . . . The legs are also<br />
blue. ..."<br />
Capart (1951:124) gave the following color<br />
account of this species: "Carapace bleu-vert, avec<br />
deux taches blanches sur les aires branchiales."<br />
Rossignol (1957:81) gave the following color<br />
description:<br />
Le 6 et la 9 sont identiques. Face dorsale:<br />
a. Carapace: marron ou brun violace plus ou moins irrise,<br />
avec deux taches arrondies d'un blanc creme sur chaque aire<br />
branchiale en arriere des dents posterieures. Sou vent, une<br />
autre petite tache blanche de forme imprecise juste au-dessus<br />
de la precedente. Extremite de la dent posterieure blanche.<br />
b. Chelipedes: brun-violace marbre de blanc (merus,<br />
carpe, propode, dactyle). Face interne de la pince: moitie<br />
inferieure 4e la paume blanche: pouce bleu turquoise.<br />
c. Pattes: bleu turquoise avec des taches blanches sur le<br />
merus (P2, P3). P4 et P5: merus et carpe brun-violace marbre<br />
de bleu et de blanc. Dactyle de P5 beige legerement rose.<br />
Face ventrale de la carapace et face externe des pinces:<br />
blanc creme.<br />
MEASUREMENTS.—The carapace width of our<br />
specimens varies between 62 and 180 mm. In the<br />
literature males with carapace lengths of 46 to<br />
114 mm, carapace widths of 85 to 205 mm,<br />
females with carapace lengths of 41.5 to 93 mm,<br />
widths 82 to 190 mm, and ovigerous females with<br />
carapace lengths of 83 to 97 mm, widths of 150<br />
to 170 mm, have been reported.<br />
REMARKS.—Portunus validus is remarkable in<br />
having the carapace finely and evenly granular,<br />
almost evenly convex, lacking sharp ridges or<br />
grooves. It falls within the genus Portunus sensu<br />
lato, as currently defined, but a careful revision<br />
of this heterogeneous genus may necessitate the<br />
recognition of a new genus for the preoccupied<br />
genus Posidon Herklots, 1851, originally established<br />
for Portunus validus. Judging from the accounts<br />
of Stephenson and Campbell (1959) and<br />
Stephenson (1972), Portunus validus shows little<br />
similarity to any of the many Indo-<strong>West</strong> Pacific<br />
species of the genus, and it shows little affinity
NUMBER 306 105<br />
with any of the American species of the genus<br />
(Rathbun, 1925; Garth and Stephenson, 1966).<br />
Indeed, a comparison of P. validus with P. pe-<br />
lagicus, the type-species of Portunus, reveals some<br />
interesting differences in addition to the ornamentation<br />
of the carapace and the structure of<br />
the male pleopod. In P. validus the interantennular<br />
spine is very short and does not extend to the<br />
front; it is not at all visible in dorsal view (Figure<br />
23a), whereas in P. pelagicus the entire spine,<br />
including its base, is visible in dorsal view, and<br />
the apex of the spine extends well beyond the<br />
level of the frontal spines (Figure 236). In P.<br />
validus there are two strong spines on the posterior<br />
margin of the merus of the claw, one subdistal<br />
(Figure 23c), whereas in P. pelagicus (and in Callinectes<br />
as well) the subdistal spine is absent (Figure<br />
23
106<br />
terminates in a bifid tip. The male abdomen has<br />
been well figured by A. Milne Edwards (1861).<br />
This species has some limited commercial importance<br />
and great commercial potential. Baron<br />
(1975b) has summarized observations on catch,<br />
size distribution, sex ratio, and growth of claw in<br />
specimens taken commercially off Senegal and<br />
Gambia.<br />
A male collected off Elmina (St. George-del-<br />
Mina), Ghana, by H. S. Pel, between 1840 and<br />
1855, is here selected as the lectotype of the<br />
species (Crust. D.395). It is in the collection of the<br />
Rijksmuseum van Natuurlijke Historic<br />
Irvine (1932) mentioned two species of Callinectes<br />
from Ghana; one he indicated correctly as<br />
C. gladiator Benedict (= C. pallidus (De Rochebrune)),<br />
the other was referred to as "Callanectes<br />
Sp.(?)" In 1947 Irvine again described and figured<br />
the two species and identified the latter as<br />
Callinectes latimanus Rathbun. The true identity of<br />
the two forms puzzled many subsequent authors,<br />
because Irvine had interchanged the figures illustrating<br />
these two species, as was first pointed out<br />
by Capart (1951:130, 132). The figures said to<br />
represent Callinectes sp. (Irvine, 1932, fig. 13) and<br />
Callinectes latimanus Rathbun (Irvine, 1947, fig.<br />
202) actually are those of C. gladiator (= C. pallidus).<br />
Irvine's text of "Callanectes Sp.(?)" (1932:14)<br />
and his figure 9, as well as his later text of C.<br />
latimanus (1947:297) do not represent Callinectes<br />
latimanus as was thought by Capart (1951), but<br />
Portunus validus. The great size of the species, the<br />
shape of the front, the fact that the merus of the<br />
chelipeds bears two distal teeth on the posterior<br />
margin, all this proves the identity of Irvine's<br />
material with P. validus. Even the peculiar white<br />
spots on the carapace are correctly indicated in<br />
the drawing, which, however, shows the carapace<br />
grooves somewhat overaccentuated. Monod<br />
(1956:196) already recognized Irvine's figures of<br />
"Callinectes gladiator" as representing Portunus validus,<br />
but evidently did not realize that they did<br />
not belong to the text of C. gladiator, but to that<br />
of the other species of "Callinectes." Williams<br />
(1974:737) recognized that something was wrong<br />
and referred to Capart and Monod, but con-<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
cluded that "since the features [of Irvine's figures]<br />
are sketchy, it is best to accept the author's designation<br />
with allowance for error."<br />
BIOLOGY.—This species inhabits shallow water<br />
in depths mostly between 0 and 40 m. Among the<br />
almost 50 depth records for this species known to<br />
us there are only two records of the capture of<br />
specimens that originate from hauls made entirely<br />
in water deeper than 40 m, viz. 50 m (Crosnier,<br />
1964:91, 92) and 50-55 m (Crosnier and Berritt,<br />
1966:135, 136); more than two-thirds of the records<br />
are from depths between 10 and 30 m.<br />
Rossignol (1957:81) therefore is mistaken in giving<br />
the range as 0-70 m and sublittoral (from 50<br />
to 99 m).<br />
The species is frequently taken with beach<br />
seines. Buchanan (1958) found it in the inshore<br />
fine sandy community in 5 to 15 m off Accra.<br />
Crosnier (1964) characterized it as a warm water<br />
species, living in depths between 0 and 30 m off<br />
Cameroon. Le Loeuff and Intes (1968:46) made<br />
the following observations on the species off the<br />
Ivory Coast:<br />
II est assez frequent dans les traits mais jamais en grandes<br />
quantites: 10 individus constituent un maximum. Espece<br />
cotiere qui ne descend pas au-dessous de 35 m, N. validus<br />
tolere de gros ecarts des facteurs physico-chimiques des eaux,<br />
ainsi que tous les types de sediments puisqu'on le peche aussi<br />
bien sur les sables de Fresco que sur les vases de Grand-<br />
Lahou.<br />
Off Guinea, Uschakov (1970) found the species<br />
in depths of less than 20 m on unstable mud in<br />
turbid water. It has been reported from bottoms<br />
consisting of shells and sand, sand and Gorgonaria,<br />
sand, muddy sand, fine sand and mud,<br />
sandy mud, mud, mud and Foraminifera, or on<br />
shells and mud.<br />
Ovigerous females have been collected in<br />
March, September, and December (Capart, 1951;<br />
Monod, 1956; Baron, 1975a).<br />
Irvine (1947:297) commented on the "ferocious<br />
disposition" of this species and the harm it can<br />
do to fishes caught in fish pots. In Ghana the<br />
crabs are caught and sold as food. According to<br />
Irvine (1947:297) "there is reason to believe that<br />
they migrate to the sea at times and it is very
NUMBER 306 107<br />
probable that they do so during the breeding<br />
season."<br />
DISTRIBUTION.—Off tropical <strong>West</strong> Africa, from<br />
Mauritania to Angola, sublittoral to about 55 m.<br />
Monod (1956), who summarized earlier records,<br />
reported material from Mauritania, Senegal,<br />
Guinea, Sierra Leone, Liberia, Ivory Coast,<br />
Ghana, and the Congo. In addition, the species<br />
has been recorded from the following localities.<br />
<strong>West</strong> Africa: No specific locality (White, 1847a; Balss,<br />
1921; Monod, 1967, 1970).<br />
Senegal: 13°25'N, 16°55'W, 15 m; 12°25'N, 17°15'W,<br />
26 m; 12°20'N, 17°5O'W (all Baron, 1975a). Saint-Louis; off<br />
the Casamance River (Baron, 1975b).<br />
Gambia: Off the Gambia River (Baron, 1975b).<br />
Guinea: No specific locality, less than 20 m (Uschakov,<br />
1970).<br />
Sierra Leone: No specific locality, 18-39 m (Longhurst,<br />
1958).<br />
Liberia: No specific locality (Biittikofer, 1890; Johnston,<br />
1906).<br />
Ivory Coast: No specific locality (Le Loeuff and Intes,<br />
1969). Off Sassandra, off Fresco, off Grand-Lahou, off<br />
Jacqueville, off Grand-Bassam, 8-35 m (Le Loeuff and Intes,<br />
1968).<br />
Ghana: Elmina [as S. Jorge da Mina] (Frade, 1950). Off<br />
Accra, 3-8 fm (5-15 m) (Buchanan, 1958); from shallow<br />
water to 25 m (Gauld, 1960).<br />
Togo: 06°15'N, 01°53'E, 15-17 m; O6°07'N, 01°53'E, 50-<br />
55 m (Crosnier and Berritt, 1966).<br />
Dahomey: 06°10'40"N, 02°02'E, 35 m; 06°10'30"N,<br />
02°02'E, 35-40 m; 06°17'N, 02°22'E, 20 m; 06°18'30"N,<br />
02°24'E, 16 m; 06°21'N, 02°37'E, 12-14 m; 06°16'N,<br />
02°37'E, 22 m; 06°12'30"N, O2°37'E, 35-36 m (Crosnier<br />
and Berritt, 1966).<br />
Cameroon: 03°55'N, 09°00'E, 50 m; 03°47'N,<br />
09°12'30"E, 20 m; 03°39'30"N, 09°14'E, 30 m; 03°32'N,<br />
09°34'E, 13-15 m; 03°27'N, 09°38'30"E, 12 m; 03°15'N,<br />
09°51'30"E, 9 m; 03°09'N, 09 o 44'30 w E, 30 m; 02°41'30"N,<br />
09°49'30"E, 20 m; 02°28'N, 09°45'30"E, 7-10 m (Crosnier,<br />
1964).<br />
Congo: Loango (Pechuel-Loesche, 1882). Off Pointe-<br />
Noire (Rossignol, 1957, 1962).<br />
Portunus vocans (A. Milne Edwards, 1878)<br />
Neptunus means.—Monod, 1956:194, figs, 222, 223 [Cape<br />
Verde Islands].—Forest, 1959:19 [Annobon, Sao Tome].<br />
Portunus vocans [Neptunus vocans].—Guinot-Dumortier and Dumortier,<br />
1960:120, 144 [table 2] [islands of Gulf of Guinea;<br />
discussion of stridulation].<br />
Portunus [Neptunus] vocans.—Guinot-Dumortier and Dumortier,<br />
1960:142 [listed].<br />
Portunus vocans.—Forest and Guinot, 1966:62, fig. 4 [Annobon,<br />
Sao Tome].—Tiirkay, 1976b:61 [listed], 66, pi. 2:<br />
figs. 1, 2 [Madeira].<br />
DISTRIBUTION.—An insular species in the Atlantic.<br />
In the eastern Atlantic it is known from<br />
Madeira, the Cape Verde Islands, and Annobon<br />
and Sao Tome Islands in the Gulf of Guinea. It<br />
also has been recorded from Ascension Island in<br />
the central Atlantic and from several western<br />
Atlantic localities; in moderate depths, 7-10 m to<br />
about 309 m (Rathbun, 1925).<br />
Genus Thalamita Latreille, 1829<br />
Thalamita Latreille, 1829:33 [type-species: Cancer admete<br />
Herbst, 1803, by monotypy; gender: feminine; name 195<br />
on Official List].<br />
Thalamita poissonii (Audouin, 1826)<br />
Portunus Poissonii Audouin, 1826:84 [Egypt].<br />
Thalamita integra var. africana Miers, 1881a:218.<br />
Thalamita africana.—Sourie, 1954b: 151.—Monod, 1956:186,<br />
figs. 213-217.—Rossignol, 1957:124 [key].—Guinot and<br />
Ribeiro, 1962:46.—Crosnier, 1962:116 [discussion].—Ribeiro,<br />
1964:5.—Forest and Guinot, 1966:61.—Uschakov,<br />
1970:455 [listed].—Stephenson, 1972:19 [key], 44 [listed].<br />
Thalamita integra africana.—Stephenson and Hudson, 1957:<br />
319 [key].<br />
Thalamita poissonii.—Holthuis and Gottlieb, 1958:89, 118, pi.<br />
2: fig. 10a,b [Eastern Mediterranean references].—Stephenson,<br />
1976:23.<br />
Thalmita africana.—Hartmann-Schroder and Hartmann,<br />
1974:19 [erroneous spelling].<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Gulf of Guinea, no specific locality, 1956,<br />
Calypso, 1
108<br />
life of this species. Monod (1956) gave the following<br />
notes in his list of material: "rougeatre" (p.<br />
187); "doigts des chelipedes (en alcool) rouges<br />
avec les apex noirs, la couleur se poursuivant, en<br />
lisere, le long des bords internes, dentigeres" (p.<br />
188).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 28 to 41 mm; the carapace widths<br />
of ovigerous females are 28 to 31 mm.<br />
REMARKS.—Crosnier (1962:117) noted the similarity<br />
between the male pleopods of T. poissonii<br />
and T. africana. Stephenson (1976:23) synonymized<br />
T. africana with T. poissonii and noted that<br />
"these [specimens from Angola] are identical in<br />
all respects with T. poissonii." We have compared<br />
our material from Angola with specimens from<br />
other areas and can find no differences.<br />
This is one of a few Indo-<strong>West</strong> Pacific species<br />
of brachyurans to occur off <strong>West</strong> Africa; it has<br />
also colonized the eastern Mediterranean via the<br />
Suez Canal (Holthuis and Gottlieb, 1958).<br />
BIOLOGY.—Thalamita poissonii is a shallow water<br />
species, occurring off <strong>West</strong> Africa from the littoral<br />
zone to a depth of about 30 m. Rathbun (1921:<br />
404) noted that it was "the most common species<br />
in the quiet stretches of the bay near town of St.<br />
Paul de Loanda, and easily caught at low tide on<br />
the many submerged sand flats. Their behavior<br />
is much the same as that of Callinectes . . . but<br />
their habitat there is typically marine." Sourie<br />
(1954b) found this species on fine shelly sand,<br />
bottom with Molgula hannensis Peres, in the Baie<br />
de Dakar. The specimens collected by the Calypso<br />
were taken on the following types of bottom<br />
(Forest and Guinot, 1966): mud, 18 m; calcareous<br />
algae, 11 m; sand, 0-4 m; sand, algae, and calcareous<br />
algae, 8-30 m; mud and calcareous algae,<br />
4 and 5 m.<br />
Off <strong>West</strong> Africa, ovigerous females have been<br />
collected in January, February, March, April,<br />
June, August, September, and October, suggesting<br />
that there the species breeds all year (Rathbun,<br />
1921; Monod, 1956; Forest and Guinot,<br />
1966).<br />
DISTRIBUTION.—Eastern Atlantic and Indo-<br />
<strong>West</strong> Pacific. In eastern Atlantic, from eastern<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Mediterranean and off <strong>West</strong> Africa at scattered<br />
localities between the Canary Islands and Angola,<br />
in depths between the littoral zone and about<br />
30 m; Indo-<strong>West</strong> Pacific from localities between<br />
Japan and Madagascar and the Red Sea (Crosnier,<br />
1962). Monod (1956) summarized the <strong>West</strong><br />
<strong>African</strong> literature and reported material from<br />
localities off Mauritania, Senegal, Guinea, and<br />
Angola. Records since then include the following.<br />
Cape Verde Islands: Bala de Porto Grande, Sao Vicente,<br />
4-6 and 8 m (Guinot and Ribeiro, 1962; Ribeiro, 1964).<br />
Senegal: Baie de Dakar (Sourie, 1954b).<br />
Guinea: No specific locality (Uschakov, 1970).<br />
Principe: In front of [Cais de] Santana, llm (Forest and<br />
Guinot, 1966).<br />
Sao Tome: 00°25'15"N, 06°43'05"E, 8-30 m; Baia de<br />
Ana de Chaves, 5 m; in front of Ponta Diogo Nunes, 4 m;<br />
Morro Peixe, 0-4 m (all Forest and Guinot, 1966).<br />
Gabon: 00°40'S, 08°46'25"E, 18 m (Forest and Guinot,<br />
1966).<br />
Angola: Baia de Luanda (Guinot and Ribeiro, 1962).<br />
Luanda (as St. Paul de Loanda) (Stephenson, 1976). Between<br />
Cacuaco and Lobito-Benguela (Hartmann-Schroder<br />
and Hartmann, 1974).<br />
Family GERYONIDAE Colosi, 1923<br />
GERYONIDAE Colosi, 1923:249.<br />
EASTERN ATLANTIC GENERA.—Due to the uncertainty<br />
of the taxonomic status of this family<br />
and of the genera sometimes assigned to it, we<br />
are unable to give here a reliable list of the eastern<br />
Atlantic geryonid genera. Guinot (1971:1077) included<br />
in the Geryonidae the following three<br />
genera that are represented in the eastern Atlantic:<br />
Geryon, Paragalene, and Progeryon; of these only<br />
Geryon is known from tropical <strong>West</strong> Africa.<br />
Paragalene Kossmann (1878:253). Type-species:<br />
Paragalene neapolitana Kossmann, 1878, a junior<br />
subjective synonym of Eriphia longicrura Nardo,<br />
1869, by monotypy; gender: feminine; name 341<br />
on Official List.<br />
Progeryon Bouvier (1922:71). Type-species: Progeryon<br />
paucidens Bouvier, 1922, by monotypy; gender:<br />
masculine.<br />
We are not convinced that these two genera<br />
are correctly assigned to the Geryonidae, but as
NUMBER 306 109<br />
we have not had the opportunity to study this<br />
question in detail, we follow Guinot, at least for<br />
the time being. The genus Platychelonion (p. 155),<br />
placed by Guinot (1971:1078) with some doubt<br />
in the Geryonidae, in our opinion finds a better<br />
place in the Xanthidae.<br />
EASTERN ATLANTIC SPECIES.—The above-mentioned<br />
three genera are represented in the eastern<br />
Atlantic area by 6 species; only two of these have<br />
been found in tropical <strong>West</strong> <strong>African</strong> waters. The<br />
other four are as follows:<br />
Geryon tridens Kr0yer, 1837. North Atlantic from<br />
N Norway and Iceland to the British Isles and<br />
the North Sea; records from the Bay of Biscay,<br />
the Mediterranean and Morocco need to be verified<br />
and at least partially pertain to G. longipes;<br />
32-690 m, usually deeper than 100 m (Christiansen,<br />
1969).<br />
Geryon longipes A. Milne Edwards, 1881. Bay of<br />
Biscay, Morocco, Mediterranean; 600-1370 m<br />
(Zariquiey Alvarez, 1968).<br />
Paragalene longicrura (Nardo, 1869). Mediterranean<br />
(Algeria, Malta, Naples, Adriatic and Aegean<br />
Seas) and Madeira; 20-ca. 120 m (Tiirkay,<br />
1976b).<br />
Progeryon paucidens Bouvier, 1922. Off Morocco,<br />
2165 m (Bouvier, 1922).<br />
Monod (1956:337) recognized a single <strong>West</strong><br />
<strong>African</strong> species: Geryon quinquedens Smith, synonymizing<br />
G. affinis with it. In the present publication<br />
two tropical <strong>West</strong> <strong>African</strong> species are distinguished:<br />
G. affinis and G. maritae, new species; G.<br />
quinquedens is thought to be restricted to East<br />
American waters.<br />
Only one species of Geryon, G. maritae, was taken<br />
by the Pillsbury in <strong>West</strong> <strong>African</strong> waters.<br />
Genus Geryon Kr0yer, 1837<br />
Geryon Kr0yer, 1837:10, 20, 21 Itype-species: Geryon tridens<br />
Kr0yer, 1837, by original designation and monotypy;<br />
gender: masculine; name 309 on Official List].<br />
Chalaepus Gerstaecker, 1856:118 [type-species: Cancer trispinosus<br />
Herbst* 1803, by monotypy; gender: masculine].<br />
REMARKS.—The place of the genus Geryon in<br />
the system of the Brachyura has been subject to<br />
many different interpretations by zoologists.<br />
Kr0yer (1837:19, 20), when describing his new<br />
genus remarked that it should be placed in the<br />
"famille des Cyclometopes" of H. Milne Edwards<br />
(1834:363) although in some respects it showed<br />
some resemblance to the Catometopes, and because<br />
of the shape of the dactyli of the last pair<br />
of pereiopods ("if0lge de bageste Tarsers Beskaffenhed")<br />
thought it should be placed in the former<br />
of the two tribes (Canceriens and Portuniens)<br />
in which H. Milne Edwards had subdivided his<br />
Cyclometopes. Miers (1886:223) placed Geryon in<br />
the subfamily Carcinoplacinae of the family Ocypodidae.<br />
Ortmann (1894:685; 1899:1176) left it<br />
in that subfamily, which he made the nominal<br />
subfamily of a new family Carcinoplacidae; he is<br />
followed in this by Stebbing (1905:35). A Milne<br />
Edwards and Bouvier (1894:41) first placed the<br />
genus in the family Canceriens sensu lato (=<br />
Cancridae + Xanthidae), but later (1899:34) assigned<br />
it to the family Galenidae. Alcock (1899c:<br />
84) accepted this disposition, but made the Galeninae<br />
a subfamily of the Xanthidae. Doflein<br />
(1904:105), without comment, ranged Geryon in<br />
the Potamidae. Colosi (1923:249) partly agreed<br />
with Doflein and erected a new (monotypic) family<br />
Geryonidae, which he placed near the Potamidae.<br />
Several authors, like Balss (1927:1020,<br />
1933a:298), Bouvier (1940:261), Stephensen<br />
(1945:222), Monod (1956:337), Guinot and Ribeiro<br />
(1962:62), Zariquiey Alvarez (1968:388),<br />
and Crosnier (1970:1216) placed the genus in the<br />
family Xanthidae; others, e.g., Rathbun (1937:<br />
265), Barnard (1950:290) and Capart (1951:173)<br />
kept it in the Goneplacidae. In recent years the<br />
family Geryonidae Colosi, 1923 (often attributed<br />
to Beurlen, 1930) is recognized by many authors.<br />
It then is usually placed between the Xanthidae<br />
and Goneplacidae (Balss, 1957:1654; Christiansen,<br />
1969:83; Glaessner, 1969:R524) or closer<br />
to the Goneplacidae than to the Xanthidae<br />
(Kaestner, 1970:349). Serological investigations<br />
by Leone (1949:284, 1951:44-48) suggested that<br />
the Geryonidae are more closely related to the<br />
Xanthidae and Portunidae on the one hand, than<br />
to the Cancridae, Ocypodidae, Grapsidae, and<br />
Majidae on the other. All in all, Stebbing's (1893:
110<br />
93) remark, made 80 years ago, still holds good:<br />
"The genus Geryon, Kr0yer, 1837, may claim a<br />
passing notice as one of those instances in which<br />
systematic arrangement finds itself at fault. It is<br />
sometimes placed among the Cyclometopa and<br />
sometimes among the Catometopa.. . . That, on<br />
the theory of the evolution of different groups<br />
from a common stem, such inosculant forms are<br />
almost sure to occur, has long been recognized.<br />
Darwin himself humorously admits that while as<br />
a theorist he delighted in coming across them, as<br />
a naturalist engaged in classification he found<br />
them an unmitigated nuisance."<br />
In studying the present material, we were<br />
struck by the great resemblance in general and in<br />
detail of Geryon with the Portunidae. In practically<br />
every point our specimens resemble Portunidae,<br />
except in the shape of the last pereiopods which<br />
lack the paddle-shape of the distal segments.<br />
Were the distal segment of these pereiopods<br />
broadened, no one would have hesitated to assign<br />
the genus to the Portunidae. The resemblance<br />
with species of Benthochascon is especially striking.<br />
The resemblance of the Geryonidae to the Portunidae<br />
is much closer, we think, than to either<br />
Xanthidae or Goneplacidae. In our opinion the<br />
family Geryonidae is close to the Portunidae,<br />
perhaps between Portunidae and Xanthidae,<br />
which incidentally would also agree with the<br />
serological findings by Leone (1949, 1951) and a<br />
study of the larvae oi Geryon tridens by Ingle (1979:<br />
230), who noted that "the larvae of G. tridens<br />
possess many portunid features...." The taxonomy<br />
of the Brachyrhynchous crabs, especially<br />
at the family level, is still highly unsatisfactory<br />
and a thorough revision is badly needed.<br />
The strong resemblance of the chelae of Geryon<br />
to those of Portunidae is shown in Figure 24. As<br />
in Portunidae the basal tooth of the dactylus of<br />
the large chelipeds in Geryon is blunt, enlarged<br />
and often directed somewhat backward. The<br />
other teeth are often flanked by small teeth; in<br />
Geryon the larger teeth usually have a single<br />
smaller tooth separating them, while in portunids<br />
the larger teeth often are flanked each with two<br />
teeth.<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 24.—Chelae: a, Geryon maritae, new species, paratype,<br />
female, cb 59.5 mm, Pillsbury Sta 51; A, Benthochascon schmitti<br />
Rathbun, female, cb 50.3 mm, Florida; c, Scylla serrata (Forskal),<br />
male, cb 55.1 mm, China.<br />
Guinot (1969c:692) mentioned the fact that<br />
between the first two abdominal segments of<br />
Geryon and the coxa of the fifth pereiopod a very<br />
small section of the 8th thoracic sternite is visible,<br />
and considered this as a goneplacid character.<br />
However, in portunid genera like Ovalipes and<br />
Benthochascon a similar situation exists, and as<br />
pointed out by Guinot (1971:1066) also in<br />
Xanthidae, like Panopeus and Rhithropanopeus, part<br />
of the 8th thoracic sternite is visible, in the last<br />
mentioned genus even "une assez importante partie<br />
laterale." In Geryon the third abdominal somite<br />
of the male covers the basal part of the coxa of<br />
the fifth pereiopod and the 8th thoracic sternite<br />
is only visible lateral to the first and second<br />
abdominal somites. All this demonstrates that the<br />
taxonomy of these crabs is still far from clear.<br />
Geryon affinis A. Milne Edwards and Bouvier,<br />
1894<br />
Geryon affinis A. Milne Edwards and Bouvier, 1894:41, figs.<br />
A, C, pi. 1; 1899:35.—Hansen, 1908:18, pi. 1: fig. 1.—
NUMBER 306 111<br />
Bouvier, 1922:70.—Rae and Lamont, 1963:24.—Kjennerud,<br />
1967:193, fig. 1.—Sankarankutty, 1968:50.—Christiansen,<br />
1969:87, fig. 35, map 29.—Mason and Davidson,<br />
1969:208.—Tiirkay, 1976b:61 [listed], 70.<br />
Geryon quinquedens.—Bouvier, 1922:70, pi. 6: fig. 7.—Monod,<br />
1956:337 [part, not fig. 441]. [Not Geryon quinquedens Smith,<br />
1879.]<br />
Geryon tridens.—Saemundsson, 1937:21 [not Geryon tridens<br />
Kr0yer, 1837].<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Azores: E of Corvo, 39°41'35"N, 31°04'-<br />
07"W of Paris (= 28°44'07"W of Greenwich), 844 m, sand<br />
and gravel, 7 Aug 1888, Princesse Alice Sta 222, syntypes, 1(5,<br />
1$ (MP).<br />
SE of Madeira: 32°42'N, 16°43'W, 670 m, tent trap, 13<br />
Mar 1976, Onversaagd Sta 63, 1(5, 19 ov (L).<br />
Cape Verde Islands: 16°44'N, 24°48'05"W, 692 m, hard<br />
bottom, 21-22 Jul 1901, Princesse Alice Sta 1138, 16* (MP).<br />
DESCRIPTION.—A. Milne Edwards and Bouvier,<br />
1894:41-45; Christiansen, 1969:87.<br />
Figures: A. Milne Edwards and Bouvier, 1894,<br />
figs. A, C, pi. 1 (color); Christiansen, 1969, fig.<br />
35.<br />
Male Pleopod: We have found no illustrations of<br />
the male pleopod of this species. Sankarankutty<br />
(1968:51) remarked that it was similar to that of<br />
G. maritae as figured by Doflein (1904).<br />
Color: A colored figure of one of the type-specimens<br />
was published by A. Milne Edwards and<br />
Bouvier (1894, pi. 1). It shows a pale brown crab<br />
with the cervical groove and posterolateral margins,<br />
as well as the spine on the carpus, with a<br />
pink hue; the central part of the carapace is<br />
somewhat greenish. Kjennerud (1967:194) remarked<br />
of her specimen: "The colour of the<br />
specimen before it was preserved in alcohol was<br />
dull yellow with patches of red and red brown,<br />
the tips and the margins of the dactyls in the<br />
walking legs were dark brown. In general it may<br />
be said that the colour was very like the colour of<br />
one of the type-specimens illustrated by Milne-<br />
Edwards and Bouvier." Christiansen (1969:85)<br />
on the other hand described the color as "red to<br />
brick-red." A. Milne Edwards and Bouvier (1899:<br />
35) mentioned a probably abnormal specimen<br />
"a pattes blanches."<br />
MEASUREMENTS.—The male and female syntypes<br />
examined have the carapace lengths 109<br />
and 112 mm, the carapace widths 134 and<br />
135 mm, respectively. In the male and female<br />
from Madeira the carapace widths are 140 and<br />
145 mm, respectively. In the male from the Cape<br />
Verde Islands the carapace length is 133 mm, the<br />
carapace width 158 mm. A. Milne Edwards and<br />
Bouvier (1894) gave the following measurements<br />
of some of the types: males, carapace lengths 15,<br />
97, and 133 mm, carapace widths 17, 112, and<br />
153 mm, respectively; females, carapace length<br />
128 mm, carapace width 14-(?) mm. (Due to a<br />
printer's error the last digit of this number was<br />
not printed so that the width may be anywhere<br />
between 140 and 149 mm). One of Saemundsson's<br />
(1937) specimens was 160 mm long by<br />
180 mm wide; Kjennerud's (1967) male had a<br />
length of 150 mm and a width of 180 mm. The<br />
5 males reported upon by Mason and Davidson<br />
(1969) had carapace widths ranging from 192 to<br />
210 mm. In contrast to these large specimens are<br />
the specimens reported upon by Hansen (1908),<br />
the male of which was 40 mm long, the ovigerous<br />
female 42 mm. The only other ovigerous female<br />
of which the measurements are known is the one<br />
(cb 145 mm) from Madeira (above). A. Milne<br />
Edwards and Bouvier, (1894:43) mentioned the<br />
eggs as very small. Hansen (1908:19) gave the egg<br />
size in his specimen as 0.5-0.6 mm.<br />
REMARKS.—The differences between the present<br />
species and G. maritae are enumerated under<br />
the latter species. Most records of Geryon qffinis or<br />
Geryon quinquedens from <strong>West</strong> Africa pertain to G.<br />
maritae. It was a surprise to find that the male<br />
from the Cape Verde Islands listed above, and<br />
already reported upon by Bouvier (1922), belongs<br />
to G. affinis rather than to G. maritae; this shows<br />
that both species do inhabit the tropical <strong>West</strong><br />
<strong>African</strong> region. The Princesse Alice specimens were<br />
examined by Holthuis in Paris on 4 October<br />
1971.<br />
The specimen from the Azores assigned by<br />
Bouvier (1922) to Geryon quinquedens is, as shown<br />
by Bouvier's illustration, a good Geryon qffinis.<br />
BIOLOGY.—In the eastern Atlantic the species<br />
is known from depths between 130 and 2047 m;<br />
almost 50% of the records is from 1165 to 2047
112<br />
m, the other half is from 130 to 844 m. The<br />
bottom on which the species was found was described<br />
as sand and gravel (A. Milne Edwards<br />
and Bouvier, 1894); muddy sand, black sand,<br />
sand and rock (A. Milne Edwards and Bouvier,<br />
1899); hard bottom (Bouvier, 1922).<br />
Ovigerous females have been collected in August<br />
(A. Milne Edwards and Bouvier, 1894); our<br />
ovigerous female from Madeira was taken in<br />
March.<br />
DISTRIBUTION.—In the eastern Atlantic the species<br />
has been reported from the following localities:<br />
Near Iceland: SW edge of Hvalsbaksbanki (approximately<br />
64°N, 13°W), 140 m (Saemundsson, 1937). SW of<br />
Geirfuglasker, Vestmanneyjar (approximately 63°N, 21°W),<br />
130 m (Saemundsson, 1937). SW of Vestmanneyjar (approximately<br />
63°N, 21°W), 240 fm (440 m) (Rae and Lamont,<br />
1963). S of Iceland, 61°30'N, 22°3O'W, 975 fm (1785 m),<br />
and 61°33'N, 19 O (AV, 1089 fm (2000 m) (Hansen, 1908).<br />
Off SW Norway: Nyegga Bank (approximately 63°38'N,<br />
05°52'E), 410 m (Kjennerud, 1967; Sankarankutty, 1968).<br />
Faroe Islands: ? Off Enniberg (approximately 62°24'N,<br />
06°33'W) (Mason and Davidson, 1969).<br />
Atlantic Ocean, NW of Scotland: Lousy Bank (approximately<br />
60°N, 13°30'W), 440 m (Mason and Davidson,<br />
1969). Rosemary Bank (approximately 59°15'N, 10°W),<br />
480 m (Mason and Davidson, 1969). George Bligh Bank<br />
(approximately 59°N, 13°30'W), 240 fm (440 m) (Rae and<br />
Lamont, 1963; Mason and Davidson, 1969). 30 miles [48<br />
km] SW of Rockall, 57°18'N, 14°25'W (Mason and Davidson,<br />
1969).<br />
Azores: 39°39'10''N, 31°03'40"W, 1300 m; 39°21'20"N,<br />
31°06'08"W, 1360 m; 38°27'N, 26°31'W, 1165 m; 38°25'-<br />
50"N, 28°34'30 w W, 785 m; 38 o 01'N, 29°22'30"W, 1260 m;<br />
37°43'N, 25°06'W, 1385 m (all A. Milne Edwards and<br />
Bouvier, 1899). Near Ilha de Sao Miguel, 37°37'N, 25°2O'-<br />
45"W, 1187 m (Bouvier, 1922). E of Ilha do Corvo, 39°41'-<br />
35"N, 28°44'07"W, 844 m; SE of Ilha do Corvo, 1386 m;<br />
between Ilha do Pico and Ilha de Sao Jorge, 38°38'N,<br />
28°08'15"W, 620 m (A. Milne Edwards and Bouvier, 1894).<br />
Madeira: No specific locality, ca. 1000 m (Tiirkay,<br />
1976b).<br />
Cape Verde Islands: W of the Cape Verde Islands, 16°-<br />
44'N, 24°48'05"W, 692 m; near Ilha de Maio, 15°17'40"N,<br />
23°02'45"W, 1300 m (Bouvier, 1922).<br />
Christiansen (1969, map 29) provided distribution<br />
charts of the species. Outside the eastern<br />
Atlantic region Geryon affinis has been reported<br />
from the east coast of America, from South Africa,<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
the Indian Ocean, and Australia. A comparison<br />
of material from those areas with specimens from<br />
the eastern Atlantic is highly desirable in order<br />
to ascertain whether or not a single species is<br />
involved in all these records. The taxonomic confusion<br />
within the genus Geryon is notorious. Many<br />
authors (including Rathbun, 1937:271; Barnard,<br />
1950:291; Monod, 1956:337) considered Geryon<br />
affinis and G. quinquedens synonymous, notwithstanding<br />
the fact that A. Milne Edwards and<br />
Bouvier (1894:41-45, figs. A-D) and Chace<br />
(1940:38-40) had convincingly shown the differences<br />
between the two species. A revision of the<br />
genus is highly desirable.<br />
* Geryon maritae, new species<br />
FIGURES 24a, 25, 26<br />
Geryon chuni Doflein, 1903:21 [nomen nudum].<br />
Geryon affinis.—Doflein, 1903:23; 1904:106, 110, 111, 175,<br />
177, 180, 186, 190, 204, 206, 208, 210, 211,213, 214, 219,<br />
220, 251, 253, 255, 257, 258, text figs. 9, 14, 32, 62,<br />
unnumbered pi.: figs. 1, 2, pis. 3, 33, 34, 38: figs. 1-5, 9,<br />
pi. 41: figs. 3-7, pi. 43: figs. 2, 8, pi. 49/50: fig. 1, pi. 52:<br />
fig. 7, pi. 55: fig. 6, pi. 58: figs. 2, 4 [part]. [Not Geryon<br />
affinis A. Milne Edwards and Bouvier, 1894.]<br />
Geryon quinquedens.—Capart, 1951:173, fig. 66.—Monod,<br />
1956:337, fig. 441 [part].—Guinot and Ribeiro, 1962:<br />
62.—Forest, 1963:628.—Monod, 1967:180, pi. 17: fig. 2<br />
[no material].—Le Loeuff and Intes, 1969:66.—Crosnier,<br />
1970:1216.—Le Loeuff, Intes, and Le Guen, 1974:73, figs.<br />
1-4.—Intes and Le Loeuff, 1976:101. [Not Geryon quinquedens<br />
Smith, 1879.]<br />
Geryon.—Voss, 1966:19.—Maurin, 1968b, figs. 1,4.<br />
Geryon quinquidens.—Maurin, 1968a:50; 1968b:484, 487, 491,<br />
492, fig. 6 [erroneous spelling.]<br />
"Third form of Geryon".—Christiansen, 1969:87.<br />
Geryon sp.—Dias and Seita Machado, 1973:1.—Bas, Arias,<br />
and Guerra, 1976, table 3.<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 74, 641-733 m, Id (holotype, L), 1$ (W).<br />
Ivory Coast: Sta 41, 641-842 m, 1(5, 1$, 10 juv (L). Sta<br />
44, 403-586 m, hard, dark gray mud, 4
NUMBER 306<br />
FIGURE 25.—Geryon maritae, new species (from Monod, 1956,<br />
fig. 441).<br />
times wider than long, greatest width at posterior<br />
pair of anterolateral teeth. Median pair of frontal<br />
teeth about as long as but narrower than lateral<br />
pair, overreaching lateral teeth by half their<br />
length. Median sinus U-shaped. Orbital margin<br />
L-shaped, short leg of L forming lateral margin of<br />
front, longer leg curving distally to join exorbital<br />
tooth. Orbital margin, apart from 2 fissures,<br />
smooth, lacking tubercles. Exorbital tooth and<br />
second and last (= fourth) anterolateral teeth of<br />
carapace distinct but low and triangular. First<br />
anterolateral tooth vaguely visible as low angle,<br />
third tooth absent or visible as low, indistinct<br />
elevation. Distances between anterolateral teeth<br />
subequal, only slightly greater than distance between<br />
exorbital tooth and first anterolateral<br />
tooth. Grooves of carapace very faint, much less<br />
distinct than in G. quinquedens. Blunt ridge extending<br />
anteriorly from near base of fifth leg, running<br />
more or less parallel to lateral margin of carapace.<br />
Anterior half of dorsal surface of carapace relatively<br />
smooth, with some groups of low, eroded<br />
tubercles on elevated parts. Posterior part of carapace<br />
with many distinct, coarse tubercles, becoming<br />
finer laterally.<br />
Suborbital margin terminating in strong inner<br />
tooth, directed anteriorly, extending to or just<br />
beyond second segment of antennular peduncle.<br />
Lower orbital margin distinctly tuberculate, ending<br />
before reaching exorbital tooth.<br />
113<br />
Abdomen triangular, closely resembling that of<br />
G. quinquedens, differing in lacking transverse carina<br />
on fourth somite. Telson almost an equilateral<br />
triangle.<br />
Epistome shorter than in G. quinquedens, with<br />
deep pit in middle of basal part.<br />
Mouth parts of this species have been well<br />
figured by Doflein (1904, pi. 38: figs. 1-5, 9).<br />
Mandible with heavy molar part and rather slender<br />
3-segmented palp, with last segment somewhat<br />
sickle-shaped. Maxillule (Figure 26) with<br />
very slender lower lacinia, upper widening distally,<br />
twice as wide at distal margin as at base;<br />
palp 2-segmented, basal segment about as wide<br />
as long or slightly wider, distal segment about<br />
half as wide as basal segment, slender in comparison.<br />
In Doflein's (1904, pi. 38: fig. 5) illustration<br />
an evidently mutilated maxillule is shown, which<br />
lacks part of the upper lacinia; therefore a new<br />
figure of the maxillule is provided here (Figure<br />
26). Maxilla with 2 inner laciniae deeply incised,<br />
forming together 4 unequal slender lobes; basal<br />
part of palp almost circular, distal part very<br />
narrow, ending in slender sharp point, somewhat<br />
curved; scaphognathite broad, truncated posteriorly.<br />
First maxilliped with 2 laciniae, upper<br />
about twice as high as lower; endopod 2-segmented,<br />
distal segment inversely triangular with<br />
FIGURE 26.—Maxillule of Geryon maritae, new species, paratype,<br />
female, cb 59.5 mm, Pillsbury Sta 51.
114<br />
distal margin wide; faint lobe visible on inner<br />
margin; exopod with well-developed flagellum;<br />
epipod large, ending in elongate narrow distal<br />
part. Second maxilliped pediform; peduncle of<br />
exopod reaching beyond endopod; well-developed<br />
flagellum present. Merus of third maxilliped<br />
with outer angle rounded, not produced.<br />
Larger of 2 chelipeds with dactylus 5/4 to 4/3<br />
as long as palm (measured along upper margin).<br />
As in Carcinus proximal teeth of fingers wide,<br />
molar-like, distal fingers flattened, tooth-like. Distal<br />
teeth alternatingly large and small. Upper<br />
surface of dactylus minutely granular, with some<br />
coarse tubercles basally; normal ridges and rows<br />
of pits present on fingers. Palm lacking anterodorsal<br />
spine, outer surface with transverse rows of<br />
tubercles and with broad longitudinal groove on<br />
upper part and 1 or 2 low, inconspicuous ridges<br />
over middle. Smaller chela resembling larger, but<br />
with fingers proportionately slightly longer and<br />
lacking molar-like teeth. Dorsal surface of carpus<br />
(of both chelipeds) with very strong, rather sharp<br />
tubercles, inner margin with large, sharp tooth,<br />
anterior and outer margins unarmed. Merus of<br />
usual shape, triangular in transverse section, with<br />
strong subdistal dorsal spine, distal spine absent.<br />
Merus less slender than in G. quinquedens.<br />
Walking legs slender, merus of last leg 4 times<br />
as long as high. Dorsal part of anterior margins<br />
of walking legs angular, unarmed. Subdistal tubercle<br />
present just behind transverse dorsal<br />
groove on distal part of merus, with smaller tubercles<br />
present on dorsal surface, restricted to<br />
distal part, more numerous and more distinct on<br />
anterior legs than on posterior ones. Distinct tubercles<br />
present on upper margin of carpus and<br />
propodus. Propodus of fifth leg about 4 times as<br />
long as high, 5/4 as long as dactylus (length<br />
measured on dorsal margin). Dactylus dorsoven-<br />
trally flattened, resembling that of G. quinquedens.<br />
Posterior margin of dactylus with distal groove,<br />
most distinct on second pereiopod (first walking<br />
leg). Lower posterior margin of dactylus of second<br />
pereiopod with row of closely placed short hairs<br />
over proximal two-thirds. Third pereiopod with<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
few scattered tufts of hairs on dactylus, tufts<br />
completely absent on fourth and fifth pereiopods.<br />
Female, carapace length 89 mm, width 95 mm:<br />
First and third anterolateral teeth of carapace<br />
more distinct than in male, but far less conspicuous<br />
than second and fourth teeth. First tooth a<br />
low triangle, third a slight elevation. Otherwise<br />
carapace similar to that of male but slightly<br />
smoother.<br />
Young male (Pillsbury Sta 41), carapace length<br />
27 mm, width 34 mm: Carapace relatively wider<br />
than in adult male, still widest at level of last pair<br />
of anterolateral teeth. Median pair of frontal<br />
teeth short, bluntly rounded, placed close together,<br />
extending about as far as slightly broader<br />
lateral frontal teeth. Exorbital tooth and second<br />
and fourth anterolateral teeth slender, spiniform,<br />
far more distinct than in adult. First and third<br />
anterolateral teeth scarcely visible, presence indicated<br />
by slight blunt hump; anterior end of<br />
hump of third tooth often abrupt, more clearly<br />
marked than that of first. Grooves of carapace<br />
more distinct than in adult, branchiocardiac<br />
grooves particularly well developed. Entire surface<br />
of carapace finely tuberculated, but elevated<br />
portions of anterior half and most of posterior<br />
part with coarse tubercles. Abdomen similar to<br />
that of adult, but with faint low transverse carina<br />
on fourth and less distinct carina on fifth somite.<br />
Orbit and epistome as in adult. Merus of third<br />
maxilliped with anterolateral angle slightly more<br />
auricularly produced.<br />
Chelipeds slenderer than in adult, with fingers<br />
somewhat longer. First tooth of dactylus large,<br />
rounded, directed proximally, exactly as in many<br />
Portunidae; other teeth also quite portunid-like.<br />
Tuberculation and pitting of chela as in adult<br />
male. No trace of spine present at broadly<br />
rounded distal end of upper margin of palm.<br />
Carpus as in large male, but spine at inner margin<br />
longer, more slender; carpus lacking any trace of<br />
spine on anterior margin. Merus only with subdistal<br />
dorsal spine, distal dorsal spine absent.<br />
Pereiopods as slender as in adult. Sharp tubercles<br />
present on upper margin of merus, carpus,<br />
and propodus, merus lacking distal dorsal spine.
NUMBER 306 115<br />
Dactylus scoop-shaped, as in adults. On second<br />
pereiopod row of hairs visible along either margin<br />
of flattened area, rows interrupted on third and<br />
fourth pereiopods, scarcely discernible on fifth<br />
pereiopod.<br />
Very small specimens, carapace length 11-<br />
17 mm: carapace slightly wider, 4/3 as wide as<br />
long. Legs slightly slenderer, merus of fifth leg<br />
about 4.3 times as long as wide. First and third<br />
anterolateral teeth of carapace visible as small<br />
lobes, first more distinct than third.<br />
Male Pleopod: Doflein, 1904, pi. 41: fig. 6 (locality<br />
not stated).<br />
Color: Color photographs were taken of the<br />
large male holotype of Pillsbury Sta 74, immediately<br />
after it came aboard. The ground color of<br />
the body is pale cream. The carapace is grayish<br />
brown in the anterior part, with a pale median<br />
longitudinal zone. Black spots of irregular size,<br />
which are irregularly arranged, are found in the<br />
middle and in the anterolateral parts of the carapace;<br />
they probably are caused by external agencies<br />
and do not form part of the actual color<br />
pattern. The chelipeds are cream with a black<br />
color along the cutting edges of the fingers. The<br />
propodi and dactyli of the walking legs are reddish<br />
brown, contrasting with the cream color of<br />
the merus; the carpus is somewhat intermediate<br />
in color. In alcohol the male specimens and the<br />
juveniles are pale cream colored, the females in<br />
several instances are more of a reddish brown.<br />
Doflein (1904, pi. 3) published a color plate of a<br />
male, showing it entirely reddish brown. Capart<br />
(1951:174) gave the following color description of<br />
his material: "Couleur generate bistre; zone rouge<br />
brique a rose sur l'arriere de la carapace et sur les<br />
pattes; dents laterales plus foncees. Les grands<br />
specimens generalement a coloration plus intense."<br />
MEASUREMENTS.—The males examined have<br />
the carapaces up to 140 mm long and 160 mm<br />
wide. The largest female has a carapace length of<br />
89 mm, a width of 95 mm. The smallest specimen<br />
has a carapace length of 11 mm, a width of<br />
15 mm. The records in the literature mostly deal<br />
with animals smaller than our largest. Doflein<br />
(1904) indicated that his largest specimen was<br />
about 90 mm long, about 110 mm wide; as<br />
Doflein's material included ovigerous females<br />
they must have been smaller than this. Le Loeuff,<br />
Intes and Le Guen (1974) gave the carapace<br />
width of their largest female as 110 mm, and that<br />
of their largest male as 164 mm. Dias and Seita<br />
Machado (1973) reported upon a male with carapace<br />
width of 165 mm. The species can attain a<br />
weight of up to 1650 g (Le Loeuff, Intes and Le<br />
Guen, 1974).<br />
REMARKS.—Geryon maritae closely resembles Ger-<br />
yon qffinis A. Milne Edwards and Bouvier, but<br />
differs in the following points.<br />
1. The front between the orbits is narrower,<br />
because the inner margins of the orbits are less<br />
strongly divergent. In G. maritae this margin is<br />
short, straight, and runs only slightly obliquely<br />
backwards. It then abruptly turns into the posterior<br />
margin of the orbit, which from there goes<br />
in an almost straight line to the exorbital tooth.<br />
In G. qffinis the inner orbital margin is much<br />
longer, diverges widely, and has a little hump in<br />
the middle. It gradually merges with the posterior<br />
margin of the orbit, which at first is directed<br />
obliquely laterally and then turns more anteriorly<br />
to the apex of the exorbital tooth. Capart's (1951,<br />
fig. 66) drawing very clearly shows the situation<br />
as it is in G. maritae.<br />
2. The first anterolateral tooth usually is more<br />
distinct in G. qffinis than in G. maritae.<br />
3. The inner infraorbital tooth in G. maritae is<br />
shorter and reaches hardly past the second segment<br />
of the antennula; in G. qffinis it is much<br />
larger and stronger and reaches far beyond that<br />
segment.<br />
4. In G. affinis the lower orbital angle is finely<br />
and closely tuberculate, in G. maritae these tubercles<br />
as a rule are placed wider apart.<br />
5. The main character distinguishing the two<br />
species, however, is shown by the dactyli of the<br />
walking legs, which in G. qffinis are laterally<br />
compressed, while in G. maritae they are dorsoventrally<br />
flattened.<br />
In the fifth character the new species resembles<br />
G. quinquedens, a species with which G. qffinis has
116<br />
often been confused; however, G. quinquedens differs<br />
from G. maritae in the following points:<br />
1. The submedian frontal teeth reach with<br />
their entire length beyond the lateral frontal teeth<br />
and are placed on a prolongation of the front.<br />
2. The upper margin of the palm of the chelipeds<br />
ends in a distinct tooth.<br />
3. The outer anterior margin of the carpus of<br />
the cheliped has a distinct sharp tooth.<br />
4. The dorsal margin of the merus of the cheliped<br />
has a distinct distal and subdistal tooth.<br />
5. The pereiopods are longer: the merus of the<br />
fifth leg is more than five times as long as high.<br />
6. The meri of the walking legs each end in an<br />
anterodorsal tooth.<br />
7. The male abdomen is broader and the lateral<br />
margin shows incisions at the lines of separation<br />
of the segments.<br />
The species was first described and figured by<br />
Doflein (1904), who had about 40 specimens from<br />
the Valdivia Bank in the South Atlantic Ocean,<br />
which he incorrectly assigned to Geryon qffinis. A<br />
specimen from E Africa also identified by him<br />
as this species evidently is specifically distinct.<br />
Doflein's extensive description and illustrations of<br />
his Atlantic specimens leave little doubt that they<br />
belong to the present new species. Doflein described<br />
many of the important characters of the<br />
species, but attached little importance to the<br />
systematic value of features as the shape of the<br />
dactylus of the walking legs; he was inclined to<br />
unite all the species of Geryon into a single variable<br />
one. His figures (1904, pi. 41: figs. 5, 7) of the<br />
dactylus of the pereiopod distinctly show that his<br />
specimens belong in the present species and not<br />
to G. affinis.<br />
A year before the publication of his large paper<br />
on the Valdivia crabs, Doflein (1903) published a<br />
short note on the eyes of deep sea crabs, using the<br />
Valdivia material as a basis. In this preliminary<br />
paper he remarked (Doflein, 1903:21) "Bei Geryon<br />
chuni konnte ich in den Eihiillen wohlentwickelte<br />
Zoeen konstatieren." Two pages beyond this he<br />
(1903:23) made the observation: "Zudem fand<br />
ich spater bei Geryon qffinis aus mehr als 1000 m<br />
Tiefe, dass die noch in den Eischalen eingeschlos-<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
senen Larven deutlich pigmentierte Augen besassen."<br />
As Doflein's (1903:4) observations are evidently<br />
based on Valdivia material, and as the only<br />
ovigerous specimens of Geryon collected by the<br />
Valdivia were those of the present species obtained<br />
from the Valdivia Bank, the specimens referred<br />
to by Doflein (1903) under both the names Geryon<br />
chuni and G. affinis evidently belong to G. maritae.<br />
This is confirmed by Doflein's (1904:258) remark<br />
in his Valdivia report: "Dagegen konnte ich z.B.<br />
bei Geryon affinis M.-E.u.Bouv., dessen Eier klein<br />
sind, die im Ausschliipfen begriffenen Zoeen auffinden."<br />
This sentence being practically identical<br />
with the one from his 1903 (p. 21) paper cited<br />
above. It seems likely that Doflein at first intended<br />
to describe the South Atlantic form as a<br />
new species Geryon chuni, but that he later changed<br />
his mind and considered it conspecific with G.<br />
affinis. Everywhere he changed the epithet chuni<br />
to affinis, but evidently forgot to do so in the entry<br />
on p. 21 of his 1903 paper; otherwise it is difficult<br />
to understand why he indicated one species in<br />
that paper with two different names. The name<br />
G. chuni was published without any description<br />
and must be considered a nomen nudum. Its<br />
identity can only be ascertained by inference.<br />
Bouvier (1922) reported a small specimen of<br />
Geryon from the Azores as G. quinquedens; the figure<br />
given by Bouvier, however, shows that it is G.<br />
affinis (p. 110). A large male and female from the<br />
Cape Verde Islands were referred by the same<br />
author (Bouvier, 1922:70) to G. affinis. Of these<br />
two specimens the male is in the collection of the<br />
Paris Museum and could there be examined by<br />
us. Contrary to our expectations, it proves to be<br />
a true Geryon affinis.<br />
Capart (1951), under the name Geryon quinquedens,<br />
gave a good description and figure of the<br />
present species. Monod (1956:337, 338) incorrectly<br />
united Geryon affinis and G. quinquedens to a<br />
single species. His specimens clearly belong to G.<br />
maritae as shown by his figure. Crosnier (1970)<br />
dealt with our above-mentioned specimens from<br />
Angola, which are now in the collection at Leiden.<br />
It also seems most likely that the material<br />
listed by Guinot and Ribeiro (1962), and Forest
NUMBER 306 117<br />
(1963) belong to the present species. All these<br />
authors thought the synonymy of Geryon quinquedens<br />
and G. affinis very likely, but they had reservations.<br />
This is the more peculiar as both A.<br />
Milne Edwards and Bouvier (1894:41-45, figs.<br />
A-D) and Chace (1940:38-40) had enumerated<br />
very clearly the differences between these two<br />
species. The cause of the confusion evidently is<br />
that the present new species is somewhat intermediate<br />
between G. affinis and G. quinquedens: it<br />
shows the dactyli of the walking legs as in G.<br />
quinquedens, but in most of the other characters it<br />
more closely resembles G. affinis.<br />
The anatomy of this species, especially that of<br />
the eye, the statocyst, etc., has been extensively<br />
dealt with by Doflein (1904).<br />
ECONOMIC IMPORTANCE.—An active trawling<br />
fishery for this species has been developed in<br />
Angola since about 1970 (Dias and Seita Machado,<br />
1973; Le Loeuff, Intes and Le Guen, 1974),<br />
the annual catch amounting to 2000 tons. Spanish<br />
trawlers, which operate off the <strong>West</strong> coast of<br />
Africa to fish for shrimps, often catch Geryon, but<br />
only sell the chelipeds; in recent years the Japanese<br />
have also developed an interest in the species<br />
and have done some exploratory fishing for it off<br />
South-<strong>West</strong> Africa (Le Loeuff, Intes and Le Guen,<br />
1974). Le Loeuff, Intes, and Le Guen (1974) and<br />
Intes and Le Loeuff (1976) described exploratory<br />
fishing for this species off the Ivory Coast, in 400<br />
to 650 m and 300 to 700 m depth. As the rough<br />
bottom configuration there made trawling impractical,<br />
lobster pots were used with reasonable<br />
success (from 1.6 to 53 kg crabs were caught per<br />
pot; the best catches being in depths between 400<br />
and 500 m, where 30 to 50 kg crabs were obtained<br />
per pot).<br />
TYPE-LOCALITY.—Off Liberia, 04°20'N, 09°.<br />
26'W to 04°30'N, 09°22'W, 641-733 m.<br />
DISPOSITION OF TYPES.—The holotype (Crust.<br />
D. 21052), a male, carapace length 134 mm, is in<br />
the Rijksmuseum van Natuurlijke Historie, Leiden.<br />
The remaining specimens are paratypes;<br />
they have been divided between the Rijksmuseum<br />
van Natuurlijke Historie, Leiden, and the<br />
National Museum of Natural History, <strong>Smithsonian</strong><br />
<strong>Institution</strong>, Washington, D.C.<br />
ETYMOLOGY.—Dr. Marit E. Christiansen, who<br />
saw our Pillsbury material, was the first to publish<br />
that the present species is distinct from G. affinis<br />
and pointed this out in her excellent book on the<br />
Scandinavian Brachyura (1969:87). For this reason<br />
we take great pleasure in naming this interesting<br />
species for her.<br />
BIOLOGY.—This species has been reported from<br />
depths between 100 and 936 m. Only three records<br />
are definitely from less than 250 m (150-<br />
245 m, Monod, 1956; no deeper than 200 m on<br />
talus, Le Loeuff and Intes, 1969; 200 m, p. 112<br />
herein), and only four are definitely from more<br />
than 600 m (936 m, Doflein, 1904; 800 m, Dias<br />
and Seita Machado, 1973; 650 m, Le Loeuff,<br />
Intes and Le Guen, 1974; and 641-842 m, p. 112<br />
herein). The majority, 24 of 31, of the records are<br />
partly or entirely from between 250 and 600 m.<br />
The bottom on which the species has been taken<br />
has been given as mud (Guinot and Ribeiro,<br />
1962; Forest, 1963); mud, sandy mud, slightly<br />
muddy sand, greenish muddy sand (Maurin,<br />
1968b); green mud and sand (three records),<br />
brown mud and sand, green mud (Capart, 1951);<br />
hard, dark gray mud (Voss, 1966; p. 112 herein);<br />
corals (Guinot and Ribeiro, 1962).<br />
Dias and Seita Machado (1973) found that at<br />
300 m depth the catch consisted of 96.8% females,<br />
but that this percentage dropped considerably in<br />
deeper waters being 38% in 500 m, 2.2% in<br />
600 m, and 1.6% m in 800 m. Le Loeuff, Intes,<br />
and Le Guen (1974) found in their catches (400-<br />
650 m) that the females were very scarce. The<br />
males were more numerous and probably represented<br />
two age classes. In 1976 Intes and Le<br />
Loeuff observed that the percentage of males<br />
increased with increasing depth.<br />
DISTRIBUTION.—Geryon maritae is a <strong>West</strong> <strong>African</strong><br />
species, known from localities between Spanish<br />
Sahara and Valdivia Bank, South-<strong>West</strong> Africa. A<br />
fishing survey reported by Dias and Seita Machado<br />
(1973) encountered the species in depths<br />
between 300 and 800 m from 29 transects made<br />
between just N of Point-Noire, Congo, and off
118<br />
the Rio Cunene, southern Angola; their individual<br />
records are not repeated below. Other records<br />
in the literature include the following.<br />
<strong>West</strong> Africa: No specific locality (Monod, 1967).<br />
Spanish Sahara: Off Villa Cisneros, 300-500 m (Maurin,<br />
1968a, 1968b). 23°35.5'N, 16°58'W to 23°32'N, 16°57'W,<br />
362-463 m (Bas, Arias, and Guerra, 1976).<br />
Mauritania: Between Cap Timiris and Tamzak (as<br />
Tamxat), 350-600 m (Maurin, 1968b).<br />
Senegal: Off Saint-Louis, 300 m (Maurin, 1968b); S of<br />
Saint-Louis, 100-300 m (Monod, 1956). Fosse de Kayar,<br />
300-350 to 600 m (Maurin, 1968b). Off Senegal, 14° 46'-<br />
51"N, 150-245 m (Monod, 1956). 11 mi [18 km] NW of<br />
Pointe des Almadies, ca. 300 m (Monod, 1956).<br />
Liberia: No specific locality (Christiansen, 1969).<br />
Ivory Coast: No specific locality (Christiansen, 1969; Le<br />
Loeuff and Intes, 1969). Numerous localities at transects<br />
across the Ivory Coast, 03° to 07.5°W, 300-700 m (Intes and<br />
Le Loeuff, 1976). Off Grand-Bassam, 03°49'W, 400-650 m<br />
(Le Loeuff, Intes and Le Guen, 1974). 05°05'N, 04°00'W to<br />
05°04'N, 04°02'W, 403-586 m (Voss, 1966). 04°32.5'N,<br />
06°31'W, 300-455 m and 04°54'N, 03°23'W, 380-400 m<br />
(Forest, 1963).<br />
Ghana: 04°39'N, 02°46'W, 300-400 m (Forest, 1963).<br />
Angola: SW of Moita Seca, 06°08'S, 11°24'E, 250-<br />
380 m; W of Ambrizete, 07°16'S, 12°02'E, 380-420 m; W of<br />
Ponta do Morro, 10°45.5'S, 13 b 07'E, 400-500 m; NW of<br />
Egito, 11°53'S, 12°23'E, 400-500 m, and 11°53'S, 13°20'E,<br />
480-510 m (all Capart, 1951). Baia dos Tigres, 320-400 m,<br />
453-478 m (Guinot and Ribeiro, 1962).<br />
South-<strong>West</strong> Africa: 17°23'S, ll°20'E, 359 m (Crosnier,<br />
1970). Valdivia Bank, 25°27'S, 06°08.2'E, 936 m (Doflein,<br />
1904).<br />
Family PLATYXANTHIDAE Guinot, 1977<br />
PLATYXANTHIDAE Guinot, 1977:1052.<br />
This family, comprising three genera, is not<br />
represented in the eastern Atlantic.<br />
Family XANTHIDAE MacLeay, 1838<br />
PILUMNIDAE Samouella, 1819:86 [by direction, under the<br />
Plenary Powers of the International Commission on Zoological<br />
Nomenclature, not to be used in preference to<br />
Xanthidae MacLeay, 1838, but available for use by those<br />
who consider that Pilumnus Leach and Xantho Leach belong<br />
to different family-group taxa; Opinion 423 in Opinions<br />
and Declarations of the International Commission on Zoological<br />
Nomenclature, volume 14, 1956; name 74 on Official List].<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
XANTHIDAE MacLeay, 1838:59 [name 73 on Official List;<br />
there dated from Dana, 1851, in error].<br />
ERIPHIDAE MacLeay, 1838:59, 60.<br />
TRICHIDEA de Haan, 1839, pi. H.<br />
CHLORODINAE Dana, 1851b: 125.<br />
POLYDECTINAE Dana, 1851b: 127.<br />
OziNAEDana, 1851b: 127.<br />
ACTUMNINAE Dana, 1851b: 128.<br />
Carpilides A. Milne Edwards, 1862a:40.<br />
Trapezides A. Milne Edwards, 1862a:40.<br />
Liagorides A. Milne Edwards, 1862a:41.<br />
Trapeziden Nauck, 1880:64.<br />
TRAPEZIINAE Miers, 1886:163.<br />
MENIPPIDAE Ortmann, 1893b:428, 431.<br />
MYOMENIPPINAE Ortmann, 1893b:429, 432.<br />
CARPILINAE Ortmann, 1893b:429, 463.<br />
ETISINAE Ortmann, 1893b:429, 470.<br />
PANOPAEINAE Ortmann, 1893b:429, 473.<br />
DOMOECIINAE Ortmann, 1893b:429, 478.<br />
ZOZYMOIDA Alcock, 1898:77, 94.<br />
EUXANTHOIDA Alcock, 1898:77, 109.<br />
HALIMEDOIDA Alcock, 1898:77, 134.<br />
ACTAEINAE Alcock, 1898:78, 137.<br />
XANTHODIOIDA Alcock, 1898:78, 156.<br />
CYMOIDA Alcock, 1898:78, 172.<br />
PSEUDOZIOIDA Alcock, 1898:176, 180.<br />
RUPPELLIOIDA Alcock, 1898:176, 186.<br />
HETEROPANOPIOIDA Alcock, 1898:177, 207.<br />
MELIOIDA Alcock, 1898:177, 230.<br />
LYBIOIDA Serene, 1965:9, 12, 26, 37.<br />
ZALASHNAE Serene, 1968:62.<br />
LIOMEROIDA T. Sakai, 1976:xvii, 385, 390.<br />
EASTERN ATLANTIC GENERA.—Thirty-two, of<br />
which all but five are represented by species<br />
occurring off tropical <strong>West</strong> Africa. The extralimital<br />
genera are the following:<br />
Atergatis de Haan (1833:4, 17). Type-species:<br />
Cancer integerrimus Lamarck, 1818, by subsequent<br />
designation by the International Commission on<br />
Zoological Nomenclature in Opinion 73 in <strong>Smithsonian</strong><br />
Miscellaneous Collections, 73(1) :26, 1922; gender:<br />
masculine; name 124 on Official List.<br />
Neopanope A. Milne Edwards (1880, in 1873-<br />
1881:329). Type-species: Neopanope pourtalesii A.<br />
Milne Edwards, 1880, a subjective junior synonym<br />
of Panopeus packardii Kingsley, 1879, by<br />
subsequent designation by Fowler (1912:400);<br />
gender: feminine.<br />
Pilumnoides H. Milne Edwards and Lucas<br />
(1843:21). Type-species: Hepatus perlatus Poeppig,
NUMBER 306 119<br />
1836, by monotypy; gender: masculine; name 347<br />
on Official List.<br />
Rhithropanopeus Rathbun (1898:273). Type-species:<br />
Pilumnus harrisii Gould, 1841, by original<br />
designation and monotypy;. gender: masculine;<br />
name 365 on Official List.<br />
Sphaerozius Stimpson (1858a:32). Type-species:<br />
Sphaerozius nitidus Stimpson, 1858, by subsequent<br />
designation by the International Commission on<br />
Zoological Nomenclature in Opinion 85 in <strong>Smithsonian</strong><br />
Miscellaneous Collections, 73(3): 17, 1925; gender:<br />
masculine; name 372 on Official List.<br />
EASTERN ATLANTIC SPECIES—Fifty-seven, of<br />
which 44 occur off tropical <strong>West</strong> Africa. Monod<br />
(1956) recorded the following species:<br />
Name in Monod Current Name<br />
Menippe nodifrons<br />
Globopilumnus africanus<br />
Globopilumnus stridulans<br />
Epixanthus helleri<br />
Pseudozius bouvieri<br />
Pilumnus stebbingi<br />
Pilumnus perrieri<br />
Pilumnus inermis<br />
Pilumnus spinifer<br />
Parapilumnus pisifer<br />
Parapilumnus boletifer<br />
Heteropanope (Heteropanope)<br />
tuberculidens<br />
Heteropanope (Pilumnopeus)<br />
caparti<br />
Heteropanope (Pilumnopeus)<br />
africana<br />
Domecia hispida<br />
Xantho (Xantho) incisa<br />
Xantho (Xantho) pilipes<br />
Xantho (Xantho) sexdentata<br />
Xantho (Xantho) denticulata<br />
Xantho (Leptodius) inaequalis<br />
punctata<br />
Xantho (Leptodius) inaequalis<br />
convexa<br />
Xantho (Leptodius) floridana<br />
Actaea (Actaea) rufopunctata<br />
Actaea (Actaea) margaritaria<br />
Actaea (Glyptoxanthus)<br />
angolensis<br />
Menippe nodifrons<br />
Globopilumnus africanus<br />
Globopilumnus stridulans*<br />
Epixanthus hellerii*<br />
Euryozius bouvieri<br />
Euryozius pagalu,<br />
new species*<br />
Pilumnus stebbingi*<br />
Pilumnus perrieri*<br />
Pilumnus inermis<br />
Pilumnus spinifer<br />
Leopoldius pisifer*<br />
Nanopilumnus boletifer*<br />
Heteropanope tuberculidens<br />
Pilumnopeus caparti<br />
Pilumnopeus africanus*<br />
Domecia acanthophora<br />
africana*<br />
Xantho incisus<br />
Xantho pilipes<br />
Xantho sexdentatus<br />
Xanthodius denticulatus*<br />
Xanthodius inaequalis<br />
inaequalis*<br />
Xanthodius inaequalis faba<br />
Cataleptodius floridanus*<br />
Paractaea rufopunctata<br />
africana*<br />
Paractaea monodi<br />
Paractaea margaritaria*<br />
Glyptoxanthus angolensis*<br />
Actaea (Glyptoxanthus)<br />
cavemosa<br />
Actaea (Glyptoxanthus) corrosa<br />
Platypodia picta<br />
Cycloxanthops occidentalis<br />
Paraxanthias eriphioides<br />
Medaeus africanus<br />
Medaeus couchi<br />
Medaeus (?) rouxi<br />
Micropanope rufopunctata<br />
Micropanope parvula<br />
Micropanope melanodactyla<br />
Panopeus africanus<br />
Panopeus parvulus<br />
Panopeus sp.<br />
Glyptoxanthus cavemosus<br />
Glyptoxanthus corrosus<br />
Platypodiella picta<br />
Cycloxanthops occidentalis<br />
Paraxanthias eriphioides<br />
Pseudomedaeus africanus*<br />
Monodaeus couchii<br />
Monodaeus rouxi*<br />
Microcassiope minor*<br />
Coralliope parvula<br />
Nanocassiope melanodactyla*<br />
Panopeus africanus*<br />
Eurypanopeus blanchardi*<br />
Panopeus africanus*<br />
The extralimital species include the following:<br />
Atergatis roseus (Riippell, 1830). Eastern Mediterranean,<br />
Israel; an immigrant from the Red<br />
Sea; littoral (Lewinsohn and Holthuis, 1964).<br />
Heteropanope laevis (Dana, 1852). Eastern Mediterranean,<br />
Egypt, an immigrant from the Red<br />
Sea; sublittoral (Holthuis and Gottlieb, 1958;<br />
Ramadan and Dowidar, 1976).<br />
Monodaeus guinotae Forest, 1976. Mediterranean,<br />
from the Balearic Isles, the Gulf of Taranto, and<br />
off Israel, shore (?) to 800 m (Forest, 1976).<br />
Neopanope sayi (Smith, 1869). An East American<br />
species introduced into England; littoral (Naylor,<br />
1960).<br />
Pilumnoides perlatus (Poeppig, 1836). An eastern<br />
Pacific species (Rathbun, 1930:537) introduced<br />
to England, South-<strong>West</strong> Africa (L), and South<br />
Africa (Barnard, 1950:257).<br />
Pilumnopeus vauquelini (Audouin, 1826). Eastern<br />
Mediterranean, from Egypt and Israel, an immigrant<br />
from the Red Sea; sublittoral (Holthuis<br />
and Gottlieb, 1958; Ramadan and Dowidar,<br />
1976).<br />
Pilumnus aestuarii Nardo, 1869. Mediterranean,<br />
from Cadaques, Spain and Venice, Italy; shallow<br />
water (Zariquiey Alvarez, 1968).<br />
Pilumnus hirsutus Stimpson, 1858. Eastern Mediterranean,<br />
Egypt, an immigrant from the Red<br />
Sea; sublittoral (Holthuis and Gottlieb, 1958;<br />
Ramadan and Dowidar, 1976).<br />
Pilumnus villosissimus (Rafinesque, 1814). Med-
120<br />
iterranean; shallow water (Zariquiey Alvarez,<br />
1968).<br />
Rhithropanopeus harrisii (Gould, 1841). Scattered<br />
localities between the Baltic, the southern North<br />
Sea, NW and SW France, Black Sea, Sea of Azov,<br />
and Caspian Sea; introduced from North America;<br />
estuarine (Christiansen, 1969).<br />
Sphaerozius nitidus Stimpson, 1858. Eastern<br />
Mediterranean, Egypt, an immigrant from the<br />
Red Sea; shallow water (Ramadan and Dowidar,<br />
1976).<br />
Xantho granulicarpus Forest, 1953. Mediterranean;<br />
sublittoral (Zariquiey Alvarez, 1968).<br />
Xantho poressa (Olivi, 1792). Canary Islands,<br />
Portugal, and Mediterranean; shallow water<br />
(Zariquiey Alvarez, 1968).<br />
We believe that two species included in the<br />
<strong>West</strong> <strong>African</strong> fauna by Monod (1956) are based<br />
either on misidentifications or erroneously labeled<br />
specimens:<br />
Chlorodiella longimana (H. Milne Edwards,<br />
1834). A western Atlantic species reported from<br />
Sao Tome by Osorio (1887:221) (see Monod,<br />
1956:303), and not subsequently recollected<br />
there.<br />
Pilumnoides hassleri A. Milne Edwards, 1880. A<br />
western Atlantic species reported by Monod<br />
(1956:262) on the basis of a specimen supposedly<br />
from Gabon collected by Heurtel. The origin of<br />
most or all of the Heurtel collection from Gabon<br />
is doubtful.<br />
REMARKS.—The status of subfamilies and genera<br />
within the family Xanthidae is currently<br />
being studied by Mme. D. Guinot of the Museum<br />
national d'Histoire naturelle, Paris. Preliminary<br />
observations of Mme. Guinot have appeared in a<br />
series of shorter papers published between 1967<br />
to 1971 and the first of more comprehensive<br />
papers, detailing observations on three subfamilies<br />
of xanthids, Actaeinae, Polydectinae, and<br />
Trichiinae, was published in 1976. Rather than<br />
try to accomodate the <strong>West</strong> <strong>African</strong> genera within<br />
the partial framework established by Mme.<br />
Guinot to date or one of the unsatisfactory older<br />
classifications, we have arranged accounts of all<br />
of the <strong>West</strong> <strong>African</strong> genera alphabetically.<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Genus Cataleptodius Guinot, 1968<br />
Cataleptodius Guinot, 1968a:704 [type-species: Chlorodius floridanus<br />
Gibbes, 1850, by original designation; gender: masculine].—Guinot,<br />
1971:1068 [list of species].<br />
* Cataleptodius floridanus (Gibbes, 1850)<br />
Xantho (Leptodius) flondana.—Monod, 1956:291, figs. 353-<br />
355.<br />
Xantho (Leptodius) floridanus.—Forest and Guinot, 1966:75.—<br />
Guinot, 1968a:706 [discussion].<br />
Cataleptodius a.ft. floridanus.—Guinot, 1968a:706, 708 [discussion],<br />
fig. 23; 1971:1068 [listed].<br />
MATERIAL EXAMINED.—Pillsbury Material: Annobon:<br />
Sta 275, 9-69 m, rubble of coralline algae, lcJ, 2$ (1 ov)<br />
(L,W).<br />
DESCRIPTION.—We have found no full description<br />
of this species based on <strong>West</strong> <strong>African</strong> specimens,<br />
but we anticipate that Mme. Guinot, in<br />
future studies, will provide a description. A description<br />
based upon western Atlantic specimens<br />
may be found in Rathbun (1930:297).<br />
Figure: Monod, 1956, figs. 353-356.<br />
Male Pleopod: Monod, 1956, figs. 354-355 (Annobon),<br />
fig. 356 (Florida); Guinot, 1968a, fig. 23<br />
(Annobon).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 6 to 7 mm; that of the ovigerous<br />
female is 6.5 mm. Monod (1956) mentioned specimens<br />
from <strong>West</strong> Africa 8 to 20 mm wide, and<br />
Rathbun (1930) reported a carapace width of<br />
33 mm in an American specimen.<br />
REMARKS.—In her generic revision, Guinot<br />
(1968a: 706) designated the <strong>West</strong> <strong>African</strong> population<br />
of this species as Cataleptodius aff. floridanus<br />
and noted:<br />
Sous ce nom, nous separons les representants ouest-africains<br />
rapportes a Xantho (Leptodius) floridanus par Monod<br />
(1956, p. 291, fig. 353-356) et par nous-meme (Forest et<br />
Guinot, 1966, p. 75), car nous relevons de petites differences,<br />
tout au plus d'ordre sous-specifique, entre les echantillons<br />
americains et ouest-africains: aff. floridanus serait de plus petit<br />
taille et en particulier presenterait des fosses orbitaires moins<br />
arrondies et moins profondes, plus inclinees lateralement et<br />
munies a Tangle externe d'une dent moins saillante; de plus,<br />
les pattes ambulatoires semblent un peu plus greles. II y aura
NUMBER 306 121<br />
lieu de revenir sur cette question a Paide d'un materiel plus<br />
important que celui dont nous disposons aujourd'hui.<br />
Unfortunately, the Pillsbury collections are so<br />
small that we cannot add anything to the solution<br />
of this problem at this time. The question can<br />
only be settled when much more material becomes<br />
available.<br />
BIOLOGY.—The <strong>West</strong> <strong>African</strong> population of<br />
this species lives sublittorally, in depths between<br />
4 and 12 or more meters; the Pillsbury specimens<br />
were taken in the beds of coralline algae off<br />
Annobon in 9 to 69 m. All of the specimens taken<br />
by the Calypso also were taken from bottoms with<br />
coralline algae (Forest and Guinot, 1966).<br />
Off <strong>West</strong> Africa ovigerous females have been<br />
collected in May and June (Forest and Guinot,<br />
1966; Pillsbury).<br />
DISTRIBUTION.—Atlantic Ocean: western Atlantic<br />
from Bermuda and Florida to Brazil; eastern<br />
Atlantic from off Gabon and the offshore<br />
islands of the Gulf of Guinea: Annobon, Principe<br />
and Sao Tome; sublittoral in the eastern Atlantic,<br />
in depths between 4 m and more than 12 m (9-<br />
69 m). Monod (1956) was the first to report the<br />
species from the eastern Atlantic, based on material<br />
from Gabon and Annobon. The only other<br />
records that we have found in the literature are:<br />
Principe: Between Ponta da Mina and Ilheu Santana,<br />
10-12 m (Forest and Guinot, 1966).<br />
Sao Tome: 00°20'N, 06°46'E, 10 m; in front of Ponta<br />
Diogo Nunes, 4 m; off Ponta Diogo Nunes, 4-5 m; in front<br />
of Ponta Oquedelrei, 6 m; in front of Praia Lagarto, 5-6 m;<br />
Iogoiogo (all Forest and Guinot, 1966).<br />
Genus Coralliope Guinot, 1967<br />
Coralliope Guinot, 1967c:353 [type-species: Actumnus parvulus<br />
A. Milne Edwards, 1869, by original designation; gender:<br />
feminine].—Guinot, 1971:1076 [list of species].<br />
Coralliope parvula (A. Milne Edwards, 1869)<br />
Micropanope parvula.—Monod, 1956:317, figs. 393-400.—<br />
Guinot and Ribeiro, 1962:59.—Ribeiro, 1964:11.—Forest<br />
and Guinot, 1966:82.—Guinot, 1967c:348 [discussion].<br />
Actumnus parvulus.—Guinot, 1967c:355 [listed].—Garth,<br />
1968:314 [discussion].<br />
Coralliope parvula.— Guinot, 1967c, figs. 3, 11; 1971:1076<br />
[listed].<br />
SYNONYM.—Xanthodes talismani A. Milne Edwards<br />
and Bouvier, 1898.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Cape Verde Islands: Sao Vicente, 20 m, 26<br />
Jul 1883, Talisman, syntypes of Xanthodes talismani A. Milne<br />
Edwards and Bouvier, 1898, 3c5, 7? (2 ov) (L, Crust. D.1567;<br />
W, USNM 22957).<br />
DESCRIPTION.—A. Milne Edwards and Bouvier,<br />
1900:91 (as Xanthodes talismani).<br />
Figures: Monod, 1956, figs. 393-400.<br />
Male Pleopod: Monod, 1956, figs. 397-400 (Cape<br />
Verde Islands, Senegal); Guinot, 1967c, fig. 1 la,b<br />
(Senegal).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 2.4 to 6.0 mm; the carapace widths<br />
of the ovigerous females are 6.0 mm.<br />
BIOLOGY.—Coralliope parvula is a sublittoral species,<br />
inhabiting relatively shallow water, 4-5 m,<br />
to a depth of 355 m. More than 90% of the<br />
records in the recent literature are from depths of<br />
less than 50 m. The deepest record is that from<br />
Cabo Bojador from 355 m on shelly sand and<br />
coral (Monod, 1956). The Calypso collected the<br />
species on calcareous algae in 4-5 m, calcareous<br />
algae, sand and coral in 7-10 m, and rocks and<br />
coral in 25-40 m.<br />
Ovigerous females have been collected in May,<br />
July, September, November, and December<br />
(Monod, 1956; Guinot and Ribeiro, 1962; Ribeiro,<br />
1964; Forest and Guinot, 1966).<br />
DISTRIBUTION.—<strong>West</strong> Africa, from the Cape<br />
Verde Islands and Cabo Bojador, Spanish Sahara<br />
southward to Angola, including the offshore islands<br />
Principe, Sao Tome, and Annobon in the<br />
Gulf of Guinea. On the mainland it is known<br />
only from off Cabo Bojador, several localities off<br />
Senegal, and one locality off Angola. Monod<br />
(1956) summarized the literature and reported<br />
material from the Cape Verde Islands, Spanish<br />
Sahara, and Senegal. Subsequent records include<br />
the following:<br />
Cape Verde Islands: Sao Vicente (Guinot, 1967c). Baia<br />
da Matiota, Sao Vicente; Baia do Porto Grande, Sao Vicente,<br />
(3)3.5-11 m, 4-6 m, 8-10 m; Baia do Monte Trigo,
122 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Santo Antao; Porto dos Carvoeiros (as Baia do Porto Novo),<br />
Santo Antao, 12 m; Baia do Tarrafal, Sao Nicolau, 16 m;<br />
channel between Santa Luzia and Branco, 18 and 30 m;<br />
Porto Ingles, Maio, 14 m; S of Ilheu Luis Carneiro, 22 m;<br />
Baixo Joao Leitao, 25 m; Porto da Furna, Brava, 6-20 m<br />
(all Guinot and Ribeiro, 1962; Ribeiro, 1964).<br />
Senegal: Dakar (Guinot, 1967c).<br />
Principe: Tinhosa Grande (as Hermano Grande) island,<br />
12 mi [19 km] SSW of Principe, 01°20'45"N, O7°17'37"E,<br />
25-40 m (Forest and Guinot, 1966).<br />
Sao Tome: Off Ponta Diogo Nunes, 4-5 m (Forest and<br />
Guinot, 1966).<br />
Annobon: 01°24'04"S, 05°36'45"E, 7-10 m (Forest and<br />
Guinot, 1966).<br />
Angola: Praia Azul, Ponta da Caruita (Guinot and Ribeiro,<br />
1962).<br />
Genus Cycloxanthops Rathbun, 1897<br />
Cycloxanthus A. Milne Edwards, 1863:278 [invalid junior<br />
homonym of Cycloxanthus H. Milne Edwards, 1850 (Crustacea);<br />
type-species: Xantho sexdecimdentatus H. Milne Edwards<br />
and Lucas, 1843, by original designation and<br />
monotypy; gender: masculine].<br />
Cycloxanthops Rathbun, 1897b: 164 [substitute name for Cycloxanthus<br />
A. Milne Edwards, 1863; type-species: Xantho<br />
sexdecimdentatus H. Milne Edwards and Lucas, 1843; gender:<br />
feminine].—Guinot, 1968a:699; 1971:1067 [list of<br />
species, C. occidentalis not included].<br />
Cycloxanthops occidentalis (A. Milne Edwards,<br />
1867)<br />
Cycloxanthops occidentalis.—Monod, 1956:301, figs. 368-370<br />
[Senegal; references].—Gauld, 1960:70 [Ghana].—Guinot<br />
and Ribeiro, 1962:57 [Angola].—Guinot, 1968a:700 [discussion].—Crosnier,<br />
1969:530 [Congo].—Guinot, 1971:<br />
1068 [listed].<br />
DISTRIBUTION.—<strong>West</strong> Africa, from the Cape<br />
Verde Islands and Senegal to Angola; intertidal<br />
to 23 m.<br />
Genus Domccia Eydoux and Souleyet, 1842<br />
Domecia Eydoux and Souleyet, 1842:234 [type-species: Domecia<br />
hispida Eydoux and Souleyet, 1842, by monotypy;<br />
gender: feminine; name 144 on Official List].<br />
Neleus Desbonne and Schramm, 1867:35 [type-species: Neleus<br />
acanthophorus Desbonne and Schramm, 1867, by monotypy;<br />
gender: masculine].<br />
* Domecia acanthophora africana Guinot, 1964<br />
Domecia hispida.—Monod, 1956:273.—Forest, 1959:22.—<br />
Guinot and Ribeiro, 1962:52. [Not Domecia hispida Eydoux<br />
and Souleyet, 1842.]<br />
Domecia sp.—Ribeiro, 1964:8.<br />
Domecia acanthophora.—Forest and Guinot, 1966:73 [not Domecia<br />
acanthophora Desbonne and Schramm, 1867].<br />
Domecia acanthophora forma africana Guinot, 1964:272, figs. 6,<br />
10-12, 16, 17.—Forest and Guinot, 1966:73 [discussion].<br />
MATERIAL EXAMINED.—Pillsbury Material: Annobon:<br />
Sta 273, shore, Id (L). Sta 282, 18-37 m, nodular coralline<br />
algae, 1$ (W).<br />
Other Material: Annobon: S coast, 01°28.5'S, 05°37.5'E,<br />
35-55 m, 16 Jun 1967, F. Poinsard, 19 (W).<br />
DESCRIPTION.—Guinot, 1964:273-278 (comparison<br />
with the Indo-<strong>West</strong> Pacific D. hispida<br />
Eydoux and Souleyet, 1842 and D. glabra Alcock,<br />
1899).<br />
Figures: Guinot, 1964, figs. 6, 10-12, 16.<br />
Male Pleopod: Guinot, 1964, figs. 11, 12 (Cape<br />
Verde Islands; Annobon).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 4 mm.<br />
REMARKS—In 1964 Guinot clarified the status<br />
of the species of Domecia and recognized three<br />
species, two of which, D. hispida Eydoux and<br />
Souleyet, 1842 and D. glabra Alcock, 1899, are<br />
restricted to Indo-<strong>West</strong> Pacific localities. She<br />
pointed out that the oldest available name for the<br />
Atlantic species, previously thought to be D. hispida,<br />
actually was D. acanthophora (Desbonne and<br />
Schramm, 1867), a species originally described<br />
from Guadeloupe in the Antilles. She further<br />
recognized Domecia acanthophora forma africana, an<br />
infrasubspecific name without standing in nomenclature,<br />
for the smaller <strong>West</strong> <strong>African</strong> population<br />
of D. acanthophora, and noted (Guinot, 1964:<br />
281) that "les petites differences (pilosite, spinulation<br />
[and size]) separant nos exemplaires ouestafricains<br />
des specimens est-americains sont sans<br />
doute liees a des conditions ecologiques differentes."<br />
Our material is too limited to add any new<br />
information but we believe that east and west<br />
Atlantic subspecies should be recognized in this<br />
case.
NUMBER 306 123<br />
BIOLOGY.—This species usually is associated<br />
with coral; it appears to be quite abundant in<br />
rocks and coral as well as in the beds of coralline<br />
algae off Sao Tome and Annobon islands in the<br />
Gulf of Guinea (Forest and Guinot, 1966), based<br />
on collections made by the Calypso. Off <strong>West</strong><br />
Africa it occurs from the intertidal zone to a<br />
depth of 35-55 m; of 14 stations at which this<br />
species was taken by the Calypso, 10 were in depths<br />
of 10 m or less.<br />
Off <strong>West</strong> Africa, ovigerous females have been<br />
collected in June and August (Guinot and Ribeiro,<br />
1962; Ribeiro, 1964; Guinot, 1964; Forest<br />
and Guinot, 1966).<br />
DISTRIBUTION.—Atlantic; an insular form off<br />
<strong>West</strong> Africa, occurring in the Cape Verde Islands<br />
and the offshore islands of the Gulf of Guinea,<br />
Annobon, Principe, and Sao Tome, in depths<br />
from shore to about 35 m (15-40 m and 35-<br />
55 m). Records in the literature include the following:<br />
Cape Verde Islands: No specific locality (Monod, 1956;<br />
Guinot, 1964). Channel between Santa Luzia and Ilheu<br />
Branco, 18 m (Guinot and Ribeiro, 1962; Ribeiro, 1964).<br />
Baia de Santa Clara, Sao Tiago, 15-35 m; Porto da Praia,<br />
Sao Tiago, 10 m; Ponta da Areia, Fogo, 2.5 m; Ponta<br />
Garbeiro [?], Brava; Ilheu de Sal Rei, Boavista, 7 m (all<br />
Guinot, 1964).<br />
Gulf of Guinea: No specific locality (Forest, 1959).<br />
Principe: Ilheu Caroco, 2-8 m; 01°38'25"N, 07°21'35"E,<br />
35 m (Forest and Guinot, 1966).<br />
Sao Tome: Between Ponta Oquedelrei and Ponta de Sao<br />
Sebastiao, 5 m (Forest and Guinot, 1966). Ponta Diogo Vaz,<br />
W coast, 0-6 and 30 m; Praia de Santa Catarina, W coast,<br />
3-10 m; Baia de Sao Miguel, 6-10 m; Ilheu Macaco [as ilot<br />
dos Cocos], 3-8 m; Morro Peixe, 2-6 m; Ilheu das Cabras,<br />
0-2 m; and Ponta Furada, 3-8 m (all Guinot, 1964; Forest<br />
and Guinot, 1966).<br />
Annobon: Off Annobon, 12 m (Monod, 1956; Guinot,<br />
1964). Santa Cruz bay, 8-12 m (Guinot, 1964). 01°24'04"S,<br />
05°36'45"E, 7-10 m; Isla Tortuga, NW side, 15-40 m;<br />
01°25'10"S, 05°36'10"E, 20-25 m (Guinot, 1964; Forest and<br />
Guinot, 1966).<br />
Genus Epixanthus Heller, 1861<br />
Epixanthus Heller, 1861:323 [type-species: Epixanthus kolschii<br />
Heller, 1861, a subjective junior synonym of Ozius frontalis<br />
H. Milne Edwards, 1834, by monotypy; gender: masculine;<br />
name 299 on Official List].<br />
* Epixanthus hellerii A. Milne Edwards, 1867<br />
Epixanthus hellen.—Monod, 1956:236, figs. 270-272.—Longhurst,<br />
1958:88.—Gauld, 1960:70.—Guinot and Ribeiro,<br />
1962:51.—Ribeiro, 1964:8.—Forest and Guinot, 1966:<br />
68.—Garth, 1968:314 [discussion].—Uschakov, 1970:445,<br />
447, 455 [listed].<br />
SYNONYM.—Ozius corrugatus Osorio, 1887.<br />
MATERIAL EXAMINED.—Pillsbury Material: Fernando<br />
Poo: Sta 258, shore, 36*, 4$, 4 juv (L,W).<br />
Other Material: Sierra Leone: Aberdeen, J. Cadenat, Mar<br />
1948, 19 (W).<br />
DESCRIPTION.—A. Milne Edwards and Bouvier,<br />
1900:83.<br />
Figures: Monod, 1956, figs. 270-272.<br />
Male Pleopod: Monod, 1956, figs. 271, 272<br />
(Sierra Leone).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 5 to 25 mm.<br />
BIOLOGY.—Epixanthus hellerii is a primarily intertidal<br />
species which may also occur subtidally<br />
in shallow water; Forest and Guinot (1966) reported<br />
one specimen from a depth of 3-10 m off<br />
Principe. Sourie (1954a) noted that this species<br />
was a characteristic component of the understone<br />
fauna of rocky shores of Senegal, and Uschakov<br />
(1970) found it in the inferior mesolittoral zone<br />
of rocky shores off Guinea, a zone with red algae<br />
and Padina.<br />
Ovigerous females have been recorded in<br />
March and November (Monod, 1956).<br />
DISTRIBUTION.—<strong>West</strong> Africa, from the Cape<br />
Verde. Islands and Senegal to Angola, including<br />
Fernando Poo, Principe, Sao Tome, and Annobon.<br />
Monod (1956), who reported material from<br />
the Cape Verde Islands, Senegal, Guinea, Sierra<br />
Leone, and Ghana, summarized earlier records.<br />
Records in the literature since 1956 include the<br />
following:<br />
Cape Verde Islands: Baia das Gatas, Sao Vicente (Guinot<br />
and Ribeiro, 1962; Ribeiro, 1964).<br />
Guinea: lie Tamara, lies de Los; Conakry (Uschakov,<br />
1970).<br />
Sierra Leone: No specific locality (Longhurst, 1958).<br />
Ghana: Tenkpobo (as Tenpobo), Dixcove (Gauld, 1960).<br />
Principe: Ponta da Mina, beach; Ilheus dos Mosteiros, 3-<br />
10 m (Forest and Guinot, 1966).
124<br />
Sao Tome: Off Sao Tome; Iogoiogo; W of Ponta Diogo<br />
Nunes; Ilheu das Cabras, 0-2 m; and Santana (Forest and<br />
Guinot, 1966).<br />
Annobon: No specific locality (Forest and Guinot, 1966).<br />
Genus Eriphia Latreille, 1817<br />
Eriphia Latreille, 1817a:404 [type-species: Cancer spinifrons<br />
Herbst, 1785, a subjective junior synonym of Cancer verrucosus<br />
Forskal, 1775, by selection by H. Milne Edwards,<br />
1842, in 1836-1844, pi. 14: fig. 1; gender: feminine; name<br />
1621 on Official List}.<br />
Eriphia verrucosa (Forskal, 1775)<br />
Cancer verrucosa Forskal, 1775:93.<br />
Eriphia spinifrons.—Stimpson, 1907:72 [Madeira].—Bouvier,<br />
1911:226 [Mauritania].—Chapman and Santler, 1955:374<br />
[Azores].<br />
Eriphia verrucosa.—Figueira, 1960:9 [Azores].—Forest and<br />
Games, 1960:352 [Morocco].—Zariquiey Alvarez, 1968:<br />
393, figs, li, 135b [Spain; references].—Tiirkay, 1976b:61<br />
[listed], 67 [Madeira, Ilhas Desertas].<br />
SYNONYMS.—Cancer spinifrons Herbst, 1782; Eriphia<br />
spinifrons angusta Czerniavsky, 1884; Eriphia<br />
spinifrons mediterranea Czerniavsky, 1884; Eriphia<br />
spinifrons orientalis Czerniavsky, 1884.<br />
DISTRIBUTION.—Eastern Atlantic from the Bay<br />
of Biscay to Mauritania, including Madeira, the<br />
Azores, and the Mediterranean; intertidal.<br />
Genus Euryozius Miers, 1886<br />
Euryozius Miers, 1886:142 [type-species: Xantho bouvieri A.<br />
Milne Edwards, 1869, by monotypy; gender: masculine].—Guinot,<br />
1968c:320 [listed], 325 [discussion][genus<br />
restricted to type-species]; 1971:1077 [included species].<br />
Gardineria Rathbun, 1911:236 [type-species: Gardinena canora<br />
Rathbun, 1911, by monotypy; gender: feminine].—<br />
Guinot, 1967a:264; 1968b: 156; 1968c:325; 1971:1077.<br />
DISCUSSION.—Guinot (1967a: 264, footnote)<br />
noted that "Gardineria canora appartient au meme<br />
genre que l'espece ouest-africaine Pseudozius bouvieri<br />
(A. MILNE EDWARDS). Ces deux especes appartiennent,<br />
selon nous, au genre Euryozius etabli<br />
par MIERS en 1886 et dont le plus proche parent<br />
serait Carpilius." In that same paper Gardineria was<br />
assigned to the Xanthinae whereas Pseudozius<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
sensu stricto, was assigned to the Menippinae. In<br />
another paper, part of an overall review of some<br />
of the more difficult xanthid genera, Guinot<br />
(1968b: 156) resurrected Euryozius from the synonymy<br />
of Pseudozius and placed both Euryozius<br />
and Gardineria in the Menippinae. In a subsequent<br />
paper in that same series, Guinot (1968c:320, 325)<br />
recognized the subfamily Carpilinae Ortmann,<br />
1893, for the Recent genera Carpilius, Euryozius,<br />
and Gardineria, and for two other genera based on<br />
fossils.<br />
In her account of Gardineria and Euryozius,<br />
Guinot (1968c:325-330) pointed out the numerous<br />
similarities and some differences between the<br />
species of these two monotypic genera, concluding<br />
(p. 328): "la plupart des ces differences ne semblent<br />
que d'ordre specifique." Earlier (p. 326) she<br />
noted,<br />
En fait, canora est tellement proche de bouvieri que notre<br />
premier mouvement fut d'unir les deux genres Gardineria et<br />
Euryozius. Pratiquement, tous les caracteres concordent. Nous<br />
avons maintenu separe Gardineria surtout pour la raison que<br />
le male de l'espece de RATHBUN n'est pas connu.<br />
Finally, Guinot (1971:1077), in her summary<br />
of her xanthid revision, noted that Gardineria is<br />
"tres proche, sinon synonyme, d'Euryozius."<br />
We have examined the unique female holotype<br />
of Gardineria canora (USNM 41535) from Providence<br />
Island, western Indian Ocean, and conclude<br />
that it should be known as Euryozius canorus<br />
(Rathbun, 1911). As Guinot pointed out, E. canorus<br />
shares all essential features with E. bouvieri,<br />
including the stridulating ridge under the carapace,<br />
as well as the short branchial ridges. Miers<br />
(1886:142) and Rathbun (1911:236) both commented<br />
on the well-developed but incomplete<br />
branchial ridges, a unique feature of this genus.<br />
The Indian Ocean species differs from its Atlantic<br />
counterparts in having two faint notches in the<br />
orbit; at most one such notch is visible in our<br />
material of the Atlantic species.<br />
Three species of Euryozius can be recognized in<br />
the Atlantic, E. sanguineus (Linnaeus, 1767) from<br />
Ascension and Saint Helena islands in the central<br />
Atlantic; E. bouvieri (A. Milne Edwards, 1869)<br />
from Madeira, the Azores, and the Cape Verde
NUMBER 306<br />
Islands; and a new species, E. pagalu, from the have carapace lengths of 7.4 and 8.0 mm. The<br />
Gulf of Guinea. Euryozius canorus, an insular spe- male from Madeira has a carapace length of<br />
cies like those in the Atlantic, is the only repre- 28.8 mm, carapace width of 46.5 mm. Barrois'<br />
sentative of the genus from the Indo-<strong>West</strong> Pacific (1888) specimen was 25.8 mm long, 38.5 mm<br />
region.<br />
wide. Guinot (1968c) figured a female 30 mm<br />
long, 47 mm wide. This species, like E. sanguineus,<br />
Euryozius bouvieri (A. Milne Edwards, 1869) is a large species.<br />
FIGURE 27a<br />
REMARKS.—See comments under account of E.<br />
pagalu, new species.<br />
Xantho Bouvieri A. Milne Edwards, 1869:377.—Stebbing, BIOLOGY.—Almost nothing is known of the<br />
1893:63 [discussion].<br />
ecology of this species. Barrois's (1888) specimen<br />
Pseudozius bouvieri.—Miers, 1886:142 [listed].—Monod, 1956:<br />
239, figs. 275-278 [part].—Figueira, 1960:9.—Guinot-Du-<br />
from the Azores was taken on a jetty among<br />
mortier and Dumortier, 1960:121, 144 [table 2], fig. 4 rocks. The specimen reported by Figueira (1960)<br />
[stridulation].—Guinot, 1967a:264 [footnote], 272 [discus- from the Azores was found dead on a beach. One<br />
sion]; 1968b: 156 [discussion]; 1968c:325 [discussion]. of the specimens collected by the Talisman in the<br />
Ozius Edwardsi Barrois, 1888:10, pi. 1 [fig. 1 in color].— Cape Verde Islands was taken at a depth of<br />
Stebbing, 1893:63 [discussion].<br />
Pseudorius Bouvieri.—A. Milne Edwards and Bouvier, 1898:<br />
20 m, and one of the specimens identified with<br />
185 [discussion; error for Pseudozius].<br />
this species by Tiirkay (1976b:67) was taken on<br />
Pseudozius Bouvieri.—A. Milne Edwards and Bouvier, 1900: an "aus 5-6 m Tiefe gegreiferten Steinblock."<br />
82, pi. 15: figs. 11-14.—Bohn, 1901:297, 298, 302, 308, DISTRIBUTION.—Euryozius bouvieri is an insular<br />
309, 315, 316, 318 [morphology].—Monod, 1933b:529<br />
[part of synonymy only; listed].<br />
species occurring in the central eastern Atlantic,<br />
Euryozius bouvieri.—Guinot, 1968b: 156 [discussion], fig. 8 [nofrom<br />
the littoral zone to a depth of about 20 m.<br />
locality]; 1968c:325, 326, 327, 329, 330 [all discussion], fig. Records in the literature include the following:<br />
8 [no locality], pi. 2: fig. 2; 1971:1077 [listed].—Tiirkay,<br />
1976b:61 [listed], 67.<br />
Azores: No specific locality (A. Milne Edwards and Bouvier,<br />
1900). Ponta Delgada, Ilha de Sao Miguel, jetty<br />
MATERIAL EXAMINED.—Pillsbury Material: None. (Barrois, 1888). Volcano Capelhinos [= Rochas dos Capel-<br />
Other Material: Madeira: No specific locality, 1(5 (BM). hinos], on beach (Figueira, 1960).<br />
Cape Verde Islands: No specific locality, syntypes, 2 dry Madeira: No specific locality (Monod, 1956). Funchal,<br />
specimens (MP).<br />
fish market (from stomachs of littoral fishes and Phycis phycis<br />
DESCRIPTION.—A. Milne Edwards, 1869:377;<br />
Barrois, 1888:10 [as Ozius Edwardsi].<br />
(Linnaeus)); between Ponta da Oliveira and Ponta da Garajau,<br />
5-6 m (Tiirkay, 1976b).<br />
Cape Verde Islands: No specific locality (A. Milne Ed-<br />
Figures: Barrois, 1888, pi. 1; Monod, 1956, figs. wards, 1869; A. Milne Edwards and Bouvier, 1900; Bohn,<br />
275-278; Guinot, 1968c, pi. 2: fig. 2.<br />
1901; Monod, 1956; Guinot, 1968c). Baia das Gatas (as Baie<br />
Male Pleopod: Monod, 1956, figs. 276-278<br />
(Azores).<br />
Color: Barrois (1888:12, pi. 1: fig. 1) described<br />
the color in the type of Ozius edwardsi as follows:<br />
de la Gatta), Sao Vicente [indicated as type-locality]<br />
(Monod, 1956). Between Ilheu Branco and Ilheu Raso (as<br />
ilot Razzo), Sao Vicente (A. Milne Edwards and Bouvier,<br />
1900; Monod, 1956).<br />
Un beau rouge orange, legerement teinte de jaune en<br />
arriere; les pattes ravisseuses sont de meme teinte que la<br />
carapace, sauf les doigts qui sont d'une noir intense, tandis<br />
According to Barrois (1888:12), A. Milne Edwards<br />
informed him that Barrois's specimen was<br />
conspecific with a species collected by the Talis-<br />
que les pattes ambulatoires sont d'un nuance plus claire et<br />
plus rosee.<br />
man in the Canary Islands in 1883. Monod (1956:<br />
240, footnote) correctly pointed out that the Ca-<br />
The pereiopods are shown to be of uniform nary Islands were not among the localities listed<br />
color in his figure.<br />
for the species by A. Milne Edwards and Bouvier<br />
MEASUREMENTS.—The two syntypes, juveniles, (1900) in their report on the Talisman collections.<br />
125
126<br />
h<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 27.—Atlantic species of Euryozius. Euryozius bouvieri (A. Milne Edwards), dry syntype, cl<br />
7.4 mm: a, carapace. Euryozius pagalu, new species, paratype, male, cl 9.0 mm, Pillsbury Sta 275:<br />
b, carapace; c, second pereiopod; d, fifth pereiopod; e, abdomen. Euryozius sanguineus (Linnaeus),<br />
male, cl 11.7 mm, Ascension Island:/, carapace; g, second pereiopod; h, fifth pereiopod; i,<br />
abdomen.<br />
* Euryozius pagalu, new species<br />
FIGURES 27b-e, 28<br />
Pseudozius Bouvieri.—Balss, 1921:63 [not Xantho bouvieri A.<br />
Milne Edwards, 1869].<br />
Pseudozius caystrus.—Capart, 1951:19 [error for bouvieri] [not<br />
Panopeus caystrus Adams and White, 1848].<br />
Pseudozius bouvieri.—Capart, 1951:164, fig. 62.—Monod,<br />
1956:239 [part, not figs. 275-278].—Rossignol, 1962:<br />
116.—Forest and Guinot, 1966:68.<br />
young crab.—Cousteau, 1958:393 [color photograph].<br />
Euryozius bouvieri.—Guinot, 1968c, figs. 15, 16, pi. 1: fig. 3.<br />
MATERIAL EXAMINED.—Pillsbury Material: Annobon:<br />
Sta 275, 9-69 m, rubble of coralline algae, 66, 4$, 16 juv (L,<br />
W) (largest male is holotype). Sta 282, 18-37 m, nodular<br />
coralline algae, 36, 10$ (1 ov) (L, W). Sta 283, 51-55 m,<br />
nodular coralline algae, 36, 19, 9 juv (L).<br />
Other Material: Annobon: 01°28'S, 05°38'30"E, 80 m, 11<br />
Dec 1965, A. Crosnier, Ombango, \6 (W). 01°28'40"S, 05°<br />
35'50"E, 11 Dec 1965, A. Crosnier, 39 (W).<br />
DESCRIPTION.—Size small, maximum carapace<br />
width about 18 mm. Carapace (Figure 21b, 28a)<br />
oval, length 0.6 to 0.7 times width. Front broad,<br />
fronto-orbital margin 0.54 to 0.7 times carapace<br />
width. Front composed of 2 sinuate lobes, almost<br />
straight in dorsal view, sinuate in oblique anterior<br />
view in adults (convex mesially, concave laterally),<br />
less sinuate in smaller specimens; lobes<br />
separated by a distinct median notch. Front<br />
flanked dorsally by shallow groove and low, irregular<br />
ridge above groove. Orbits smooth, with<br />
indications of 1 incision in dorsal margin of<br />
smaller specimens, upraised margins continuing<br />
laterally and ventrally into suborbital margin,
NUMBER 306 127<br />
FIGURE 28.—Euryoziuspagalu, new species, paratypes. Female,<br />
cb 13.5 mm, Pillsbury Sta 282: a, dorsal view. Male, cb 14.4<br />
mm, Pillsbury Sta 275: b, gonopod; c, apex of gonopod.<br />
lateral part of ridge appearing as tubercle in<br />
dorsal view. Anterolateral margin cristate, unarmed<br />
anterior to lateralmost point of carapace,<br />
there produced into 2 low anterolateral teeth,<br />
anteriormost, anterior to midlength of carapace,<br />
occasionally not developed, posteriormost distinct<br />
behind midlength, each with short ridge on carapace,<br />
posteriormost the longer. Low irregular<br />
swelling extending laterally from orbit above lateral<br />
margin. Carapace regularly convex from<br />
front to back, regions indistinct, dorsal surface<br />
punctate, anterior surface, in arc behind front,<br />
rougher than posterior surface. Stridulating ridge<br />
under anterolateral border of carapace arising<br />
under first anterolateral tooth, extending about<br />
to midlength of lower margin of orbit, composed<br />
of about 60 striae.<br />
Chelipeds stout, unequal. Carpus with dorsal<br />
and outer surface irregular, deeply punctate; inner<br />
margin produced into rectangular lobe (the<br />
stridulating lobe) terminating anteriorly in blunt<br />
inner tooth. Surface of palm deeply punctate and<br />
longitudinally sulcate dorsally, outer surface<br />
smooth. Fingers dark, evenly toothed, with bilobed<br />
enlarged basal tooth.<br />
Walking legs slender, third (Figure 27c) much<br />
the longest, overreaching second by about 1/2 to<br />
1/3 of dactylus. First walking leg with at most a<br />
low tubercle distally on inner margin of carpus.<br />
Dactylus of each walking leg longer than respective<br />
propodus, grooved laterally and dorsally,<br />
ornamented with a few slender, simple setae.<br />
Propodi of third and fourth walking legs grooved<br />
laterally, length of propodus of third about 3<br />
times depth, length of that of fourth 2 times depth<br />
(measured dorsally).<br />
Abdomen (Figure 27*) of 7 distinct somites in<br />
both sexes, terminal somite of male abdomen<br />
longer than penultimate. Male pleopods as figured<br />
by Guinot (1968c, figs. 15, 16).<br />
Color: Like the other Atlantic species of Euryozius,<br />
E. pagalu is reddish. Capart (1951:165) noted<br />
that in his material the general color was "rose<br />
orange, les doigts bruns." Cousteau (1958) illustrated<br />
a juvenile from Sao Tome; it is red with<br />
the pereiopods banded red and white, the latter<br />
at the articulations. The color pattern of our<br />
material has completely faded.<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths ranging from 8.0 to 16.2 mm; the<br />
carapace width of the ovigerous female is 11.2<br />
mm. The largest specimens examined by Capart<br />
(1951) were a male, 10.5 X 17 mm, and a female,<br />
11.5 X 18 mm; the latter is the largest known<br />
representative of the species. Guinot (1968c) figured<br />
a male, 9.7 X 14.5 mm, from the Calypso<br />
collection.<br />
REMARKS.—Specimens from different geographic<br />
areas identified with Euryozius bouvieri in<br />
the past have exhibited an order of magnitude<br />
difference in size. For example, a syntype of Xantho<br />
bouvieri from the Cape Verde Islands measured<br />
8 X 12 mm (A. Milne Edwards, 1869), whereas<br />
another specimen from the same area measured<br />
30 X 47 mm (Guinot, 1968c). The type of Ozius<br />
edwardsi, from the Azores, measured 25.8 X 38.5
128<br />
mm (Barrois, 1888), and syntypes of Pseudozius<br />
mellissi, from Ascension and Saint Helena Islands<br />
in the central Atlantic, measured 25 X 38 mm<br />
and 35.5 X 59 mm (Miers, 1881b). Our specimens<br />
from Annobon, more than 33 in all, comprising<br />
an ovigerous female as well as adult males, do not<br />
exceed 10 X 16 mm, and are obviously adult at<br />
a size of 7 X 11 mm.<br />
This size disparity, considered together with<br />
two features reported in the literature, pereiopod<br />
length and color, suggested to us that the species<br />
now known as E. bouvieri actually might consist of<br />
a complex of species. Euryozius bouvieri is considered<br />
to be a widely distributed insular species in<br />
the central and eastern Atlantic (Monod, 1956:<br />
240), and, as noted by Monod "Cette distribution<br />
presque uniquement insulaire est singuliere."<br />
Such an insular distribution pattern for a single<br />
species is difficult to understand or explain. We<br />
decided to investigate the possibility that different<br />
species occurred in each of the three geographic<br />
areas from which E. bouvieri was reported:<br />
the mid-Atlantic, on Ascension and Saint Helena<br />
islands; the central eastern Atlantic, from Madeira,<br />
the Azores, and the Cape Verde Islands; and<br />
the Gulf of Guinea.<br />
The problem is further complicated by the fact<br />
that large specimens from both the mid-Atlantic<br />
{Pseudozius mellissi), as well as the central eastern<br />
Atlantic (Ozius edwardsi and a female from the<br />
Cape Verde Islands taken by the Talisman, see<br />
Guinot, 1968c, pi. 2: fig. 2), have a very narrow<br />
front, which in the specimens from the mid-Atlantic,<br />
at least, is distinctly four-lobed. In our<br />
specimens from Annobon, as well as in the syntypes<br />
of Xantho bouvieri, the front is comparatively<br />
quite broad and is produced into two transverse<br />
or sinuous lobes.<br />
In his original account of Xantho bouvieri from<br />
the Cape Verde Islands, A. Milne Edwards (1869:<br />
378) remarked that in his specimens the fourth<br />
pereiopods were quite long, exceeding all of the<br />
other legs by the length of the dactylus. This is<br />
not the case in our series from Annobon or in the<br />
young male (6.6 X 9.8 mm) from Ascension that<br />
we have examined; in our specimens the fourth<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
legs exceed the third by no more than half the<br />
dactylus. The small specimens from the Cape<br />
Verde Islands reported by Bouvier appear to<br />
represent a species distinct from that occurring<br />
on Annobon or that occurring in the mid-Atlantic.<br />
Those small specimens from the Cape Verde<br />
Islands represent either a distinct small species or<br />
the juveniles of the large species also known from<br />
the Cape Verde Islands. We believe that it is<br />
unlikely that two species of Euryozius occur in the<br />
Cape Verde Islands and that the types of Xantho<br />
bouvieri are based on the young of a species, which<br />
in that area attains a size of at least 30 X 47 mm.<br />
The second feature mentioned in the literature<br />
that attracted our attention is color. Euryozius<br />
bouvieri, sensu lato, is known to be a bright red<br />
crab. Melliss (1875:275) referred to an orange-red<br />
crab. A. Milne Edwards (1869), Miers (1886),<br />
Barrois (1888), and Capart (1951) all referred to<br />
the red color of this species. Barrois (1888, pi. 1:<br />
fig. 1) illustrated his Ozius edwardsi in color and<br />
showed the pereiopods to be uniform in color, not<br />
banded. Cousteau (1958: 393 photograph) illustrated<br />
a juvenile specimen from Sao Tome in<br />
color, showing walking legs distinctly banded red<br />
and white. The walking legs in all of our specimens<br />
from Ascension are distinctly banded red<br />
and white.<br />
The available material suggests to us that there<br />
are three species of Euryozius in the Atlantic: a<br />
large species with banded legs which lives on<br />
Ascension and Saint Helena; a large species with<br />
uniform coloration which is found at Madeira,<br />
the Azores, and the Cape Verde Islands; and a<br />
small species with banded legs which lives in the<br />
Gulf of Guinea.<br />
Four names are available for two of these taxa.<br />
The large central Atlantic species from Ascension<br />
and Saint Helena was named Pseudozius mellissi<br />
by Miers (1881b:432); until now it was considered<br />
a synonym of Xantho bouvieri A. Milne Edwards<br />
from the Cape Verde Islands. However, Miers'<br />
name is not the oldest available name for this<br />
species. Linnaeus (1767:2970) described an<br />
oceanic crab, Cancer sanguineus, a species with an<br />
obsoletely bidentate carapace and a four-lobed
NUMBER 306 129<br />
front that, judging from the name, is red. Herbst<br />
(1785, in 1782-1804) noted that this species occurred<br />
at Ascension. The central atlantic form is<br />
identifiable as Euryozius sanguineus, which, as noted<br />
by Linnaeus, has two low anterolateral teeth on<br />
the carapace and a distinctly four lobed front.<br />
Two names also are available for the species<br />
occurring in the central part of the eastern Atlantic,<br />
Xantho bouvieri A. Milne Edwards, 1869 and<br />
Ozius edwardsi Barrois, 1888, the former, based on<br />
juveniles, having priority.<br />
The third species, from the Gulf of Guinea, is<br />
named herein.<br />
The possibility exists that Barrois was incorrect<br />
in showing his species to have a uniform color<br />
pattern or that A. Milne Edwards was incorrect<br />
in his account of the walking legs of his species.<br />
Moreover, all of the Atlantic species of Euryozius<br />
might prove to be conspecific in which case the<br />
name E. sanguineus would have to be used. Alternatively,<br />
the same large species might occur in<br />
the central as well as in the eastern Atlantic and<br />
E. bouvieri also might occur in the Cape Verde<br />
Islands and the Gulf of Guinea. Additional fresh<br />
material will have to be studied before this problem<br />
can be settled.<br />
A detailed account of E. sanguineus is in preparation<br />
by Manning in collaboration with Fenner<br />
A. Chace, Jr., in a review of the marine decapods<br />
of Ascension Island.<br />
We have been able to compare our material of<br />
E. pagalu with three male specimens of E. sanguineus<br />
(Figure 21f-i) from Ascension Island (6.6 X<br />
9.8 mm, 17.6 X 27.8 mm, and 28.6 X 45.8 mm)<br />
and one female of E. sanguineus from Saint Helena<br />
(42.5 X 71.1 mm) in the collections of the <strong>Smithsonian</strong><br />
<strong>Institution</strong>. The specimens from Ascension,<br />
newly collected, all have the walking legs<br />
banded red and white. The fronto-orbital width<br />
of the three larger specimens of E. sanguineus<br />
ranges from 0.45 times the carapace width in the<br />
smaller male (17.6 X 27.8 mm) to 0.37 times the<br />
carapace width in the large female. In the smallest<br />
male from Ascension, of comparable size with our<br />
specimens of E. pagalu from Annobon, it is 0.67<br />
times the carapace width; it ranges from 0.54 to<br />
0.7 times the carapace width in E. pagalu.<br />
A comparison of our material with the smallest<br />
specimen of E. sanguineus from Ascension shows<br />
the following differences: the frontal lobes are less<br />
sinuous; the carapace and dorsal surfaces of the<br />
carpus and propodus of the claw are rougher; the<br />
third pereiopod overreaches the second by less of<br />
its length, 1/3 to 1/2 its length in E. pagalu, 2/3<br />
of its length in E. sanguineus; the pereiopods appear<br />
to be slenderer in E. pagalu; there is a distinct<br />
tubercle (almost a spine in larger specimens) on<br />
the inner, distal surface of the carpus of the<br />
second and third periopods in E. sanguineus; this<br />
tubercle is scarcely or not at all detectable in even<br />
the largest specimens of E. pagalu.<br />
Guinot (1968c:330) commented on differences<br />
in pilosity, possibly related to sex or size, in<br />
material available to her. Our material exhibits<br />
no obvious differences.<br />
TYPE-LOCALITY.—Off Annobon Island, Gulf of<br />
Guinea, 01°24'S, 05°37'E to 01°24'S, 05°38'E, in<br />
9 to 69 m.<br />
DISPOSITION OF TYPES.—The holotype (Crust<br />
D. 31539), the largest male from Pillsbury Sta 275,<br />
measuring 10.3 X 16.2 mm, is in the Rijksmuseum<br />
van Natuurlijke Historie, Leiden. Paratypes are<br />
in the collections of that museum and in the<br />
<strong>Smithsonian</strong> <strong>Institution</strong>.<br />
ETYMOLOGY.—The specific name is derived<br />
from the modern name for Annobon Island, Pagalu.<br />
BIOLOGY.—Euryozius pagalu is generally a sublittoral<br />
species, living from shore to a depth of<br />
80 m. Of 16 depth records available, only one is<br />
from shore (shore to 2 m); that and two others,<br />
2-6 m and 7-10 m, are in 10 m or less. Five<br />
records (9-69 m, 51-55 m, 60 m, 60 m, and<br />
80 m) are from depths generally greater than 50<br />
m. Fifty percent of the records are from depths<br />
between 15 and 40 m, suggesting a preference for<br />
that depth range.<br />
Only two records are from localities other than<br />
Sao Tome and Annobon islands: one specimen<br />
was found dead on a beach off the Congo (Rossi<br />
gnol, 1962), and Capart (1951) reported 4 males
130<br />
and 3 females from sandy mud in 20-22 m off<br />
Angola.<br />
Apparently the preferred habitat for this species<br />
is the nodular coralline algae found off Annobon<br />
and Sao Tome; all of the Pillsbury specimens<br />
and eight of the ten lots (25 specimens)<br />
collected by the Calypso were taken in coralline<br />
algae. Other kinds of bottom noted for the Calypso<br />
collections (Forest and Guinot, 1966) include:<br />
rocks and coral; one specimen from a sponge on<br />
rocky bottom with calcareous algae; sand, with<br />
calcareous algae; sand, algae, rocks, or coral.<br />
The only ovigerous female known was taken<br />
by the Pillsbury in May.<br />
DISTRIBUTION.—Euryozius pagalu is primarily an<br />
insular species occurring in the Gulf of Guinea; it<br />
lives from the shore to a depth of 80 m. Records<br />
in the literature include the following:<br />
Sao Tome: No specific locality (Cousteau, 1958). Ilheu<br />
Gago Coutinho (as Ilha das Rolas) (Balss, 1921). 00°25-<br />
15"N, 06°43'E, 8-30 m; Morro Peixe, 2-6 m (Forest and<br />
Guinot, 1966). W of Ponta Diogo Nunes, shore to 2 m (Forest<br />
and Guinot, 1966; Guinot, 1968c).<br />
Annobon: OfT Annobon, 60 m (Monod, 1956). 01°24'-<br />
04"S, 05°36'45"E, 7-10 m; 01°25'10"S, 05°36'10"E, 20-<br />
25 m; 01°25'12"S, 05°36'05"E, 20 m; 01°26'15"S,<br />
05°35'40"E, 60 m; O1°27.5'S, 05°36.5'E, 35 m; N of San<br />
Antonio, 23 m; Isla Tortuga, NW coast, 15-40 m (all Forest<br />
and Guinot, 1966).<br />
Congo: Pointe Indienne, beach (Rossignol, 1962).<br />
Angola: 8 mi [12.8 km] W of Rio Cuanza, 09°20'S,<br />
13°04'E, 20-22 m (Capart, 1951).<br />
Genus Eurypanopeus A. Milne Edwards, 1878<br />
Eurypanopeus A. Milne Edwards, 1878, in 1873-1881:318<br />
[type-species: Panopeus crenatus H. Milne Edwards and<br />
Lucas, 1843, by subsequent designation by Fowler, 1912:<br />
394; gender: masculine].<br />
* Eurypanopeus blanchardi<br />
(A. Milne Edwards, 1881)<br />
FIGURE 296<br />
Xantho parvulus.—Dana, 1852b: 170.—Miers, 1881a:214 [discussion].<br />
[Not Cancer parvulus Fabricius, 1793 = Xanthodius<br />
parvulus; not Xantho parvulus sensu H. Milne Edwards, 1834<br />
= Eurypanopeus abbreviate (Stimpson, 1860); not Actumnus<br />
parvulus A. Milne Edwards, 1869 = Coralliope parvula.]<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Xantho parvula.—Stimpson, 1858a:31; 1907:47 [= Xantho parvulus<br />
sensu Dana, 1852, not parvulus auctorum].<br />
Panopeus Blanchardi A.. Milne Edwards, 1880, in 1873-1881:<br />
308, footnote [nomen nudum]; 1881, in 1873-1881:323.<br />
Panopeus blanchardi.—Miers, 1886:129 [discussion].<br />
Eurypanopeus parvulus.—A. Milne Edwards and Bouvier, 1900:<br />
99, pi. 17: fig. 7 [not Cancer parvulus Fabricius, as in first<br />
entry above].<br />
Eurypanopeus blanchardiQ).—Rathbun, 1921:440, fig. 20, pi.<br />
36: figs. 2, 3.<br />
Panopeus parvulus.—Monod, 1956:329, figs. 416-434.—Gauld<br />
and Buchanan, 1959:127.—Gauld, 1960:70.—Guinot and<br />
Ribeiro, 1962:62—Rossignol, 1962:118.—Monod, 1963,<br />
fig. 35 [no material].—Ribeiro, 1964:13.—Forest and<br />
Guinot, 1966:85.—Uschakov, 1970:447, 455 [listed].<br />
MATERIAL EXAMINED.—Pillsbury Material: Fernando<br />
Poo: Sta 257, shore, 5(5 (W). Sta 258, shore, 5
NUMBER 306 131<br />
Elle [E. abbreviatus] ressemble beaucoup a un Panope de<br />
la cote occidentale d'Afrique que j'ai designe dans les cata-<br />
logues du Museum sous le nom de Panopeus Blanchardi, dont<br />
les pinces sont depourvues de dent basilaire et dont les bords<br />
latero-anterieurs sont un peu plus courts.<br />
Rathbun (1921:440, 441) who used the name<br />
Eurypanopeus blanchardi (?) for the species and who<br />
gave the only description of the species, commented<br />
in her account based on a specimen from<br />
Angola:<br />
I think, however, that this is with little doubt the species<br />
named blanchardi. Bouvier makes it a synonym of the Amer-<br />
ican parvulus or abbreviatus [in A. Milne Edwards and Bouvier,<br />
1900:99]. Our <strong>African</strong> specimen, however, differs from abbreviatus<br />
of equal size as follows. The carapace is slightly<br />
narrower in proportion to its length, but is wider in its<br />
posterior half, the posterolateral margins less convergent, so<br />
that the carapace appears less regularly oval than in abbreviatus.<br />
The anterolateral teeth are more pronounced and more<br />
elevated; the chelipeds more rugose, fingers more deeply<br />
grooved. A small piece of the sternum shows at each end of<br />
the second abdominal segment, between it and the coxa of<br />
the last leg; this is not the case in abbreviatus.<br />
Rathbun (1930) did not synonymize E. blanchardi<br />
with E. abbreviatus in her account of the<br />
latter species.<br />
With one exception, the characters pointed out<br />
by Rathbun to differentiate E. blanchardi from E.<br />
abbreviatus are well marked in the materal available<br />
to us. In addition to the difference in proportion<br />
of the body, in our specimens of E. blanchardi<br />
on the carapace the regions are more distinct,<br />
the surface is much more uneven, and the<br />
lines of granules are much better developed, as<br />
shown by Monod (1956, fig. 417). A character<br />
that appears to have some value is the width of<br />
the third anterolateral tooth. In Eurypanopeus abbreviatus<br />
this tooth is broader than the fourth<br />
tooth, whereas in E. blanchardi it is subequal in<br />
length to the fourth tooth. The male pleopods of<br />
the two species, as figured by Monod (1956, figs.<br />
422-434) for E. blanchardi and by Williams (1965,<br />
fig. 183K) for E. abbreviatus are essentially similar,<br />
except that in E. blanchardi the slender, subterminal,<br />
triangular lobe overreaches the irregularly<br />
shaped median lobe, whereas in E. abbreviatus it<br />
does not.<br />
The character mentioned by Rathbun, which<br />
is of relatively little value, is the exposure of the<br />
small piece of sternum between the second abdominal<br />
somite and the coxa of the last leg. In E.<br />
blanchardi the part of the sternum is clearly visible,<br />
but in material of E. abbreviatus examined for<br />
comparison the sternum is usually concealed in<br />
specimens from the Antilles (USNM 15658)<br />
whereas it usually is visible in specimens from<br />
Brazil (USNM 59844).<br />
That A. Milne Edwards (1881, in 1873-1881)<br />
described this species as lacking an enlarged basal<br />
tooth on the dactylus of the chela is puzzling, for<br />
it is clearly present in our material. Monod (1956:<br />
334), who examined what might be the types of<br />
E. blanchardi from Gabon, did not mention<br />
whether or not he observed the tooth.<br />
Although Rathbun (in Stimpson, 1907:47,<br />
footnote) identified Dana's Xantho parvulus and<br />
Stimpson's Xantho parvula, both from the Cape<br />
Verde Islands, with Xanthias melanodactylus [= Nanocassiope<br />
melanodactylus, below], we believe that<br />
Monod (1956:329) was correct in identifying<br />
those records with E. blanchardi (as Panopeus parvulus).<br />
Dana's description fits E. blanchardi quite<br />
well.<br />
The carapace of E. blanchardi (Figure 296) does<br />
appear to be less oval in shape than that of E.<br />
abbreviatus (Figure 29a) as noted by Rathbun, but<br />
the length/width ratio of representatives of the<br />
two species overlaps broadly. Dana (1852b) noted<br />
that in his specimens from the Cape Verde Islands<br />
the length/width ratio was 1 to 1.53; his reported<br />
measurements were 5 X 7% lines (10.6 X 16.2<br />
mm). Monod (1956) included measurements of<br />
numerous specimens in his account; the length/<br />
width ratio for all material reported by Monod<br />
ranged from 1:1.17 to 1:1.64. The ratio for 22<br />
larger specimens, carapace length 8.5 mm or<br />
more, ranged from 1:1.33 to 1:1.64, with a mean<br />
of 1.44. The ratio for 13 specimens from the<br />
western Atlantic from Brazil and Trinidad ranges<br />
from 1:1.47 to 1:1.56; the mean is 1:1.53. The<br />
difference in shape of the carapace of E. blanchardi<br />
and E. abbreviatus is shown in Figure 29.<br />
The specific epithet parvulus has been applied
132<br />
a<br />
FIGURE 29—Carapaces: a, Eurypanopeus abbreviatus (Stimpson),<br />
male, cb 20.0 mm, Brazil; b, Eurypanopeus blanchardi (A. Milne<br />
Edwards), male, cb 19.1 mm, Pillsbury Sta 257.<br />
to three distinct species of xanthid crabs from the<br />
Atlantic by three different authors. Fabricius<br />
(1793:451) named Cancer parvulus which Rathbun<br />
(1930:305), who had examined the type, placed<br />
in Leptodius. H. Milne Edwards (1834:395) identified<br />
a second species as Xantho parvulus (Fabricius);<br />
Rathbun (1930:404) placed this species in<br />
Eurypanopeus. Actumnus parvulus was described by<br />
A. Milne Edwards (1869), was placed in Micropanope<br />
by Monod (1933b, 1956), and finally was<br />
assigned to Coralliope by Guinot (1967c).<br />
At least three authors, Dana (1852b: 170), A.<br />
Milne Edwards (1881, in 1873-1881:322), and<br />
Benedict and Rathbun (1891:369), considered<br />
Cancer parvulus Fabricius and Xantho parvulus sensu<br />
H. Milne Edwards to be synonymous. Rathbun<br />
(1930), without discussing the problem, correctly<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
replaced parvulus sensu H. Milne Edwards with<br />
the next oldest available name, Panopeus abbreviatus<br />
Stimpson, 1860, which she, as A. Milne Edwards<br />
(1881, in 1873-1881:320) had before her, considered<br />
to be a Eurypanopeus.<br />
The problem of the two identical specific<br />
names has extended even further. Monod (1956:<br />
285), in discussing the synonymy and affinities of<br />
Xanthodius inaequalis [as Xantho (Leptodius)], compared<br />
it with an American species which Rathbun<br />
(1930:305) called Leptodius parvulus (Fabricius,<br />
1793), and concluded that the eastern and<br />
western Atlantic populations represented distinct<br />
species. Because of the confusion over the use of<br />
parvulus, Monod suggested that the American<br />
population be called Xantho (Leptodius) americana<br />
(Saussure, 1858): "Comme par ailleurs le binome<br />
"Xantho parvulus" ne peut que provoquer de nouvelles<br />
confusions, je serais d'avis d'utiliser Xantho<br />
(Leptadius) [sic] americana (SAUSSURE)." Guinot<br />
(1968a: 712; 1971:1068) followed Monod in substituting<br />
americanus for parvulus; she assigned De<br />
Saussure's species to Xanthodius. That species<br />
should be known as Xanthodius parvulus (Fabricius,<br />
1793).<br />
BIOLOGY.—Eurypanopeus blanchardi is an intertidal<br />
species, which may occur subtidally to a depth<br />
of 10-30 m; the majority of records in the literature<br />
are based on specimens taken intertidally on<br />
rocky shores. Gauld (1960) noted that it is common<br />
on rocky shores off Ghana, and Gauld and<br />
Buchanan (1959) characterize it as a member of<br />
the understone fauna. Uschakov (1970) found it<br />
in the mesolittoral zone of rocky shores of Guinea,<br />
in habitats with red algae and Padina.<br />
It has been taken together with Panopeus ajricanus,<br />
but in general that species prefers lagoons<br />
and estuaries rather than rocky shores.<br />
Ovigerous females have been collected in January,<br />
February, March, May, June, and November<br />
(Monod, 1956; Forest and Guinot, 1966; present<br />
paper). The species probably spawns all year.<br />
DISTRIBUTION.—<strong>West</strong> Africa, where it occurs<br />
from the Cape Verde Islands and Mauritania<br />
southward to Angola, including the offshore islands<br />
in the Gulf of Guinea, Principe, Sao Tome,
NUMBER 306 133<br />
and Fernando Poo (new record); usually intertidal,<br />
rarely to a depth of 10-30 m. Records in the<br />
literature include the following.<br />
<strong>West</strong> Africa: No specific locality (A. Milne Edwards,<br />
1880, 1881, in 1873-1881; Monod, 1963).<br />
Cape Verde Islands: No specific locality (Dana 1852b).<br />
Sao Tiago (as St. Jago) (Stimpson, 1858a). Porto da Praia,<br />
Sao Tiago, 12 fm (22 m) (Stimpson, 1907); beach (Guinot<br />
and Ribeiro, 1962; Ribeiro, 1964); same, (as La Praya), 10-<br />
30 m (A. Milne Edwards and Bouvier, 1900). Sao Vicente<br />
(Monod, 1956).<br />
Mauritania: No specific locality; Bilaouakh [?] (Monod,<br />
1956).<br />
Senegal: Goree (?); Ngazobil, shore (Monod, 1956).<br />
Guinea: Conakry, shore (Monod, 1956). lie Tamara and<br />
ile Roume, lies de Los, shore (Uschakov, 1970).<br />
Sierra Leone: Aberdeen, near Freetown, shore (Monod,<br />
1956).<br />
Ghana: Accra, shore (Monod, 1956). Christiansborg, Accra<br />
(Monod, 1956). Tenkpobo (as Tenpobo), shore or reef<br />
(Monod, 1956; Gauld and Buchanan, 1959). Apam, Prampram;<br />
Dixcove (Monod, 1956). Tenkpobo to Dixcove, shore<br />
(Gauld, 1960).<br />
Principe: Praia Ponta da Mina, shore (Forest and Guinot,<br />
1966).<br />
Sao Tome: Harbormaster's dock, shore (Forest and<br />
Guinot, 1966).<br />
Gabon: No specific locality (Monod, 1956).<br />
Congo: Baie de Pointe-Noire (Rossignol, 1962).<br />
Angola: Santo Antonio do Zaire (as San Antonio), less<br />
than 20 ft [6.1 m] (Rathbun, 1921). Lucira, beach (Guinot<br />
and Ribeiro, 1962).<br />
Genus Globopilumnus Balss, 1933<br />
Globopilumnus Balss, 1933b:6 [type-species: Pilumnus globosus<br />
Dana, 1852, by original designation; gender: masculine].<br />
Globopilumnus afiricanus<br />
(A. Milne Edwards, 1867)<br />
MATERIAL EXAMINED—Pillsbury Material: None.<br />
Other Material: Senegal: No other data, 2? (W).<br />
DESCRIPTION.—Capart, 1951:142.<br />
Figures: Capart, 1951, fig. 51, pi. 1: fig. 15, pi.<br />
3: figs. 7, 8; Monod, 1956, figs. 249-257.<br />
Male Pleopod: Capart, 1951, pi. 3: figs. 7, 8<br />
(Angola); Monod, 1956, figs. 254-257 (Senegal);<br />
Turkay, 1976b, fig. 4 (Madeira).<br />
Color: "Rougeatre plus ou moins fonce, a maculatures<br />
brun chocolat ou violettes; epines antero-laterales<br />
et doigts de chelipedes noirs"<br />
(Monod, 1956:235). The black color of the fixed<br />
finger of the chela extends well onto the palm<br />
(Monod, 1956, fig. 251).<br />
MEASUREMENTS—Our specimens have carapace<br />
widths of 20 and 22 mm.<br />
BIOLOGY.—Globopilumnus africanus is a shallow<br />
water species, generally found intertidally or subtidally;<br />
of 22 records for which depth is given in<br />
the recent literature, seven are from shore, 11 are<br />
from depths between shore and 12 m, and four<br />
are from depths greater than 12 m (15, 20-22, 30,<br />
and 35 m). Intertidally it appears to be part of<br />
the rocky shore fauna, where it lives among rocks<br />
or in algae; some specimens have been taken with<br />
bryozoans. Subtidally it has been taken on sandy<br />
mud, rocks, rocks and coral, rocks and sand, or<br />
on sand with calcareous algae, rocks and coral.<br />
Ovigerous females have been collected in January<br />
and September (Capart, 1951; Guinot and<br />
Ribeiro, 1962; Ribeiro, 1964).<br />
DISTRIBUTION.—Off <strong>West</strong> Africa, from Madeira,<br />
the Cape Verde Islands, Senegal, offshore<br />
islands of the Gulf of Guinea, and Angola, intertidal<br />
to 35 m, generally in shallower water.<br />
Monod (1956:230) commented on its apparent<br />
absence on the mainland between Senegal and<br />
Pilumnus qfricanus.—Capart, 1951:142, fig. 51, pi. 1: fig. 15,<br />
pi. 3: figs. 7, 8.—Garth, 1968:314 [discussion].<br />
Angola; he summarized earlier records and re-<br />
Globopilumnus qfricanus.—Monod, 1956:227, figs. 249-257.— ported material from the Cape Verde Islands,<br />
Guinot-Dumortier, 1959:98, 99, 116, 118, fig. 14 [discus- Senegal, Sao Tome, and Angola. Subsequent recsion;<br />
review of Indo-<strong>West</strong> Pacific Globopilumnus].—Guinot- ords in the literature include the following:<br />
Dumortier and Dumortier, 1960:138, fig. 21 [stridulation].—Guinot<br />
and Ribeiro, 1962:51.—Ribeiro, 1964:7.— <strong>West</strong> Africa: No specific locality (Guinot-Dumortier and<br />
Forest and Guinot, 1966:66.—Garth, 1968:312, 313 [dis- Dumortier, 1960).<br />
cussion].— Turkay, 1976b:61 [listed], 67, fig. 4, pi. 2: figs. Madeira: Funchal, fish market (from stomach of Phycis<br />
3,4.<br />
phycis (Linnaeus)) (Turkay, 1976b).
134<br />
DESCRIPTION.—Monod, 1956:231.<br />
Figures: Monod, 1956, figs. 258-269.<br />
Male Pleopod: Monod, 1956, figs. 263-267 (Senegal,<br />
Guinea, Annobon).<br />
Color: "Blanc sale" (Monod, 1956:235). Our<br />
specimens have a reddish body, with red and<br />
white banded legs.<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 2 to 20 mm.<br />
REMARKS.—The larger specimens taken by the<br />
Pillsbury off Annobon differ from Monod's account<br />
in having the anterolateral spines of the<br />
carapace and those on the outer face of the wrist<br />
and palm of the chela dark; in none of the<br />
specimens does the dark color of the fixed finger<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Cape Verde Islands: Matiota, Sao Vicente (Guinot and of the chela extend onto the palm. The large male<br />
Ribeiro, 1962; Ribeiro, 1964).<br />
Principe: Ilheu Caroco (as ilot Carioco), 2-8 m; 01°38'-<br />
35"N, O7°21'35"E, 35 m; and S of Ilheu Santana (as tie Sta.<br />
Ana), 12 m (all Forest and Guinot, 1966).<br />
Sao Tome: 00°25'N, 06°37'E, 5-6 m; Ponta Diogo Vaz,<br />
0-6 m; Praia de Santa Catarina, 3-10 m; Baia de Sao<br />
from Pillsbury Sta 282 (carapace width 17 mm)<br />
differs from the specimen from Annobon illustrated<br />
by Monod (1956, fig. 261) in having the<br />
terminal segment of the abdomen semicircular<br />
rather than subtriangular.<br />
Miguel, 6-10 m; Ilheu Macaco (as Hot dos Cocos), 3-8 m; E<br />
of the beach, Morro Peixe, 2-6 m; Ilheu das Cabras, 0-2 m;<br />
Santana, shore; Ponta Furada, 3-8 m (all Forest and Guinot,<br />
1966).<br />
Annobon: Bay of Santa Cruz, 8-12 m (Forest and Guinot,<br />
1966).<br />
BIOLOGY.—In contrast with G. africanus, this<br />
species prefers sublittoral habitats, as noted by<br />
Forest and Guinot (1966:67, 68). Of 16 depth<br />
records in the literature, 6 are from depths ranging<br />
from 0-6 m to 10 m whereas 10 are from<br />
Angola: Baia da Caota, Benguela, shore; Baia das Vacas, depths ranging from 15-40 m to 73 m; none was<br />
shore; Lucira; Praia das Conchas, Mocamedes, shore (all<br />
Guinot and Ribeiro, 1962).<br />
from the intertidal. The Calypso took both species<br />
together at three stations, on sand with calcareous<br />
algae, rocks, and coral in 35 m and on rocks and<br />
* Globopilumnus stridulans Monod, 1956 coral in 0-6 m and 30 m. This species, like G.<br />
Globopilumnus stridulans Monod, 1956:230, figs. 258-269.— africanus, apparently prefers rough bottom with<br />
Guinot-Dumortier, 1959:98, 99, 116, 118, fig. 14 [discus- rocks and coral or coralline algae. Crosnier (1964:<br />
sion; review of Indo-<strong>West</strong> Pacific Globopilumnus].—Guinot- 31) found it off Cameroon in sponges, coral, and<br />
Dumortier and Dumortier, 1960:138, 139, figs. 20, 21<br />
[stridulation].—Rossignol, 1962:116.—Crosnier, 1964:<br />
31.—Forest and Guinot, 1966:67.—Garth, 1968:312, 313,<br />
314, 315, 316 [discussion].<br />
marl on rocky bottom with gorgonians. Part of<br />
Monod's material was taken from a wash of<br />
sponges.<br />
No ovigerous females have been recorded.<br />
MATERIAL EXAMINED.—Pillsbury Material: Annobon:<br />
Sta 275, 9-69 m, rubble of coralline algae, 4(3, 12$ (L). Sta<br />
282, 18-37 m, nodular coralline algae, 16\ 2$ (W).<br />
Other Material: Principe: 12 mi [19 km] SSW of Tinhosa<br />
Grande (as Hermano Grande) Island, 01°20'45"N, 07° 17'-<br />
37"E, 25-40 m, rocks and coral, 7 Jul 1956, Calypso Sta 110,<br />
DISTRIBUTION.—Off <strong>West</strong> Africa, from a few<br />
mainland localities between Senegal and Gabon<br />
and from the offshore islands of Principe, Sao<br />
Tome, and Annobon; intertidal to a depth of<br />
73 m. Monod (1956) described the species on<br />
13, 1$ (W).<br />
Annobon: 01°28.5'S, 05°37.5'E, 35-55 m, F. Poinsard,<br />
2(5, 89, 8juv(W).<br />
material from Senegal, Guinea (the type-locality),<br />
and Annobon; the holotype from Guinea was<br />
taken in 40 m. Subsequent records in the literature<br />
include the following:<br />
<strong>West</strong> Africa: No specific locality (Guinot-Dumortier and<br />
Dumortier, 1960).<br />
Cameroon: No specific locality (Crosnier, 1964).<br />
Principe: 01°36'50"N, O7°22'1O"E, 19 m; S part, Baia<br />
das Agulhas, 4-8 m; 01°38'35"N, 07°21'35"E, 35 m; 01°-<br />
43'10"N, 07°28'20"E, 73 m; 01°43'N, O7°28'55"E, 37 m;<br />
and Tinhosa Grande (as Hermano Grande) Island, 12 mi<br />
[19 km] SSW of Principe, O1°2O'45"N, O7°17'37"E, 25-<br />
40 m (all Forest and Guinot, 1966).<br />
Sao Tome: 00°20'N, 06°46'E, 10 m; Ponta Diogo Vaz,<br />
W coast, 0-6 m and 30 m; in front of Ponta Oquedelrei, 6<br />
m; and off Ponta Diogo Nunes, 6 m (all Forest and Guinot,<br />
1966).<br />
Annobon: 01°24'04"S, 05°36'45"E, 7-10 m; 01°25'-<br />
12"S, 05°36'05"E, 20 m; and NW coast, Isla Tortuga, 15-<br />
40 m (all Forest and Guinot, 1966).
NUMBER 306 135<br />
Gabon: W of Cap Santa Clara, Libreville, 45-47 m<br />
(Rossignol, 1962).<br />
Genus Glyptoxanthus A. Milne Edwards, 1879<br />
Glyptoxanthus A. Milne Edwards, 1879, in 1873-1881:253<br />
[type-species: Actaea erosa Stimpson, 1859, by selection by<br />
Rathbun, 1930:263; gender: masculine].—Guinot, 1971:<br />
1072 [list of species].<br />
* Glyptoxanthus angolensis<br />
(De Brito Capello, 1866)<br />
Actaea angolensis.—Capart, 1951:157, fig. 59.—Longhurst,<br />
1958:88.—Gauld, 1960:70.—Rossignol, 1962:117.<br />
Actaea (Glyptoxanthus) angolensis.—Monod, 1956:296, fig.<br />
361.—Serene, 1961:198 [listed].<br />
Glyptoxanthus angolensis.—Forest and Guinot, 1966:77, fig. 6a,<br />
b.—Guinot, 1971:1073 [listed].<br />
Glyptoxanthus anglolensis.—Guinot, 1967d:556 [listed; erroneous<br />
spelling].<br />
MATERIAL EXAMINED.—Pillsbury Material: Annobon:<br />
Sta 282, 18-37 m, nodular coralline algae, let, 19 (L, W).<br />
Other Material: Annobon: S coast, 01°28.5'S, O5°37.5'E,<br />
35-55 m, 16 Jun 1967, F. Poinsard, \6 (W).<br />
DESCRIPTION.—Capart, 1951:158.<br />
Figures: Capart, 1951, fig. 59; Monod, 1956, fig.<br />
361; Forest and Guinot, 1966, fig. 6a,b.<br />
Male Pleopod: Forest and Guinot, 1966, fig. 6a,b<br />
(Annobon).<br />
Color: "Rouge orange avec des taches plus<br />
claires" (Capart, 1951:158).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 12, 17 and 25 mm.<br />
BIOLOGY.—Glyptoxanthus angolensis is a shallow<br />
shelf species, occurring sublittorally in depths<br />
between 4-5 m and 40 m, with one record of 35-<br />
55 m. Like the <strong>West</strong> <strong>African</strong> species of Paractaea,<br />
habitats on rough bottom are preferred. The<br />
Pillsbury specimens were collected in nodular calcareous<br />
algae off Annobon, and the material<br />
collected by the Calypso (Forest and Guinot, 1966)<br />
was found on the following types of bottom:<br />
calcareous algae, 4-5 m; calcareous algae, sand<br />
and coral, 7-10 m; rocks and calcareous algae,<br />
20 m; and rocks and coral, 15-40 m. Longhurst<br />
(1958) found the species on shelly sand in 25 m<br />
off Sierra Leone.<br />
Ovigerous females have been recorded in January<br />
and June (Monod, 1956; Forest and Guinot,<br />
1966).<br />
DISTRIBUTION.—<strong>West</strong> <strong>African</strong> coast, from scattered<br />
localities between Senegal and Angola, including<br />
Annobon and Sao Tome islands in the<br />
Gulf of Guinea; sublittoral, from 4-5 m to 40 m<br />
(also 35-55m). Monod (1956) summarized earlier<br />
records and reported material from Senegal,<br />
Sierra Leone, Congo (Congo Francais), and Annobon;<br />
records in the literature since 1956 include<br />
the following:<br />
Sierra Leone: No specific locality, in 25 m (Longhurst,<br />
1958).<br />
Ghana: Off Accra, sublittorally; Tenkpobo (asTenpobo),<br />
intertidal (Gauld, 1960).<br />
Sao Tome: Off Ponta Diogo Nunes, 4-5 m (Forest and<br />
Guinot, 1966).<br />
Annobon: 02°24'04"S, 05°36'45"E, 7-10 m; 01°25'12"S,<br />
05°36'05"E, 20 m; and Isla Tortuga, NW coast, 15-40 m<br />
(Forest and Guinot, 1966).<br />
Congo: Pointe Indienne (Rossignol, 1962).<br />
Angola: No specific locality (Forest and Guinot, 1966).<br />
Glyptoxanthus cavernosus<br />
(A. Milne Edwards, 1878)<br />
Xantho vermiculatus.—Osorio, 1906:149 [Cape Verde Islands;<br />
see Forest and Guinot, 1966:79] [not Cancer vermiculatus<br />
Lamarck, 1818].<br />
Actaea (Glyptoxanthus) cavemosa.—Monod, 1956:298, fig. 362<br />
[Cape Verde Islands; references].—Serene, 1961:198<br />
[listed].<br />
Glyptoxanthus cavemosa.—Forest and Guinot, 1966:79 [Cape<br />
Verde Islands].<br />
Glyptoxanthus cavemosus.—Guinot, 1967d:551, 553 [discussion],<br />
556 [listed], fig. 31 [Cape Verde Islands]; 1971:1073<br />
[listed].<br />
DISTRIBUTION.—Known only from the Cape<br />
Verde Islands.<br />
Glyptoxanthus corrosus<br />
(A. Milne Edwards, 1869)<br />
Actaea (Glyptoxanthus) corrosa.—Monod, 1956:298 [Cape<br />
Verde Islands; references].<br />
Glyptoxanthus corrosus.—Guinot, 1967d:556 [listed], fig. 21<br />
[Cape Verde Islands]; 1971:1073 [listed].<br />
DISTRIBUTION.—Cape Verde Islands.
136 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Genus Heteropanope Stimpson, 1858<br />
Heteropanope Stimpson, 1858a:33 [type-species: Heteropanope<br />
glabra Stimpson, 1858, by subsequent selection by Balss,<br />
1933b:32; gender: feminine; name 1627 on Official List].<br />
Heteropanope acanthocarpus Crosnier, 1967<br />
Heteropanope acanthocarpus Crosnier, 1967:325, figs. 3, 4, 7-10,<br />
28 [Dahomey]; 1971:570 [Congo].<br />
DISTRIBUTION.—<strong>West</strong> Africa, from Dahomey<br />
(06°10'N, O2°27'E), 55 m, and off Pointe-Noire,<br />
Congo, 20 m.<br />
Heteropanope tuberculidens Monod, 1956<br />
Heteropanope tuberculidens Monod, 1956:265, figs. 309-313.—<br />
Gauld, 1960:70.—Crosnier, 1967:325, figs. 1, 2, 6, 27.—<br />
Uschakov, 1970:444, 455 [listed].<br />
Heteropanope tubulicidens.—Longhurst, 1958:88 [erroneous<br />
spelling].<br />
MATERIAL EXAMINED.—PMsbury Material: None.<br />
Other Material: Togo: 06° 1 l'N, 01 °28'30"E, 14-17 m, sand<br />
with gorgonians, mud, 17 Oct 1963, A. Crosnier, 16\ 2$ (W).<br />
Congo: Pointe-Noire, 5-10 m, lobster net, 27 Jan 1967, J.<br />
Marteau, Id, 2$, 1 juv (W).<br />
DESCRIPTION.—Monod, 1956:265.<br />
Figures: Monod, 1956, figs. 309-313; Crosnier,<br />
1967, figs. 1, 2, 6, 27.<br />
Male Pleopod: Monod, 1956, figs. 312, 313<br />
(Ghana); Crosnier, 1967, fig. 27 (Guinea-Bissau).<br />
Color: "Plus ou moins brun rouge, traces<br />
d'annelures aux pattes" (Monod, 1956:268).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 2.6 to 5.2 mm.<br />
REMARKS.—Our specimens agree well with accounts<br />
of this species, except that in our smaller<br />
specimens from the Congo the outer face of the<br />
chela is relatively smooth, and in one of the<br />
females from the same locality the tubercles on<br />
the palm are aligned in rows.<br />
BIOLOGY.—Heteropanope tuberculidens is a sublittoral<br />
species living in relatively shallow water.<br />
Depth records for the species range from 0-2.5 m<br />
off Guinea (Uschakov, 1970) to 72 m off Sierra<br />
Leone (Longhurst, 1958); the latter is the only<br />
record for the species in depths below 25-30 m,<br />
and the majority of depth records are from less<br />
than 20 m; records include: 0-2.5, 8-16, 10-12,<br />
14-17, 15 (2X), 25-30, and 72 m. The species<br />
apparently prefers rough bottom; it has been<br />
recorded from bottom with coral; gravel, sand<br />
and shell; muddy sand with gorgonians; sand,<br />
rock or mud, and gorgonians; and shelly mud.<br />
Ovigerous females have been collected in<br />
March, April, and June (Monod, 1956; Crosnier,<br />
1967).<br />
DISTRIBUTION.—Off <strong>West</strong> Africa, from a few<br />
localities between Guinea-Bissau and the Congo,<br />
in depths to 72 m, generally in less than 20 m.<br />
Records in the literature include the following:<br />
Guinea-Bissau: Channel between Ilha de Rubane (as<br />
Rouban) and Ilha de Bubaque, Bissagos Islands, 25-30 m,<br />
and channel near Ilha de Soga, Arquipelago dos Bijagos (as<br />
Bissagos Islands), 8-16 m (Monod, 1956; Crosnier, 1967).<br />
Guinea: Conakry, 0-2.5 m (Uschakov, 1970). Between lie<br />
Tamara and lie de Corail, 10-12 m (Monod, 1956).<br />
Sierra Leone: No specific locality, in 72 m (Longhurst,<br />
1958).<br />
Ghana: Off Accra, 15 m (Gauld, 1960). 2 mi [3 km] off<br />
Densu, near Accra, 15 m (type-locality) (Monod, 1956).<br />
Togo: 06°ll'N, 01°28'30"E, 14-17 m (Crosnier, 1967).<br />
Congo: Off Pointe-Noire (Crosnier, 1967).<br />
Genus Leopoldius Serene, 1971<br />
Leopoldius Serene, 1971:908 [an invalid junior homonym of<br />
Leopoldius Rondani, 1843 (Insecta); type-species: Parapilumnus<br />
leopoldi Gordon, 1934, by original designation; gender:<br />
masculine].<br />
REMARKS.—R. Serene has informed us (in litt.)<br />
that he will propose a replacement name for this<br />
genus.<br />
•Leopoldius pisifer (MacLeay, 1838),<br />
new combination<br />
Parapilwnnus pisifer.—Capart, 1951:146, fig. 53, pi. 3: fig.<br />
18.—Sourie, 1954b: 150.—Monod, 1956:254, figs. 298-<br />
301.—Longhurst, 1958:88.—Gauld, 1960:70.—Rossignol,<br />
1962:117.—Guinot and Ribeiro, 1962:52.—Forest and<br />
Guinot, 1966:71.—Crosnier, 1969:535.—Uschakov, 1970:<br />
439, 455 [listed].—Takeda, 1974:216 [discussion].<br />
Synonym.—Pilumnus verrucosipes Stimpson,<br />
1858.
NUMBER 306 137<br />
MATERIAL EXAMINED.—Pillsbury Material: Ivory<br />
Coast: Sta 46, 38-42 m, mud with dense Julltenella, 66, 5$<br />
(4 ov) (W). Sta 47, 37 m, bottom with Jullienella, 26, 2$<br />
(1 ov) (L).<br />
Nigeria: Sta 248, 33 m, 16, 1$ (L, W). Sta 253, 33-40 m,<br />
mud, 1(5 (L).<br />
Other Material: Senegal: Les Almadies, Dakar, 0-0.5 m,<br />
under stones, 9 Jun 1964, F. M. Bayer, R. B. Manning, and<br />
L. B. Holthuis, Ic5, 1$ (L). Goree, lobster net, 15 Sep 1950,<br />
M. Paraiso, 216, 26$ (12 ov) (W). Same locality, 6 Oct 1950,<br />
M. Keita, 26, 19 ov (W).<br />
Congo: Pointe-Noire, rocks with gorgonians, 25 Jan 1967,<br />
A. Crosnier, 46 (W).<br />
DESCRIPTION.—Capart, 1951:146.<br />
Figures: Capart, 1951, fig. 53, pi. 3: fig. 18;<br />
Monod, 1956, figs. 298-301.<br />
Male Pleopod: Capart, 1951, pi. 3: fig. 18 (Senegal);<br />
Monod, 1956, figs. 300, 301 (Senegal).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 3 to 10 mm; the carapace widths<br />
of ovigerous females are 6 to 9 mm.<br />
REMARKS.—As suggested by Takeda (1974:<br />
216), this species appears to belong to Leopoldius<br />
Serene, 1971, rather than to Parapilumnus Kossmann,<br />
1877. In L. pisifer the carapace is covered<br />
with a thick coat, arranged more or less symmetrically,<br />
the lobes of the front are sinuous rather<br />
than truncated, the three posterior anterolateral<br />
teeth of the carapace are broad, not spiniform,<br />
and their borders are granulated, the greatest<br />
width of the carapace is at the level of the posteriormost<br />
tooth, the pereiopods are short and irregularly<br />
carinate, and the lateral palatal ridges<br />
are completely absent.<br />
This is the only species of Leopoldius to occur in<br />
the Atlantic; the remainder of the species are<br />
from the Indo-<strong>West</strong> Pacific region.<br />
BIOLOGY.—Like Nanopilumnus boletifer (p. 145),<br />
Leopoldius pisifer is a shallow shelf species, living<br />
from the intertidal zone to a depth of 50 m; the<br />
majority of records in the literature are from<br />
depths of 30 m or less. It can utilize a wide range<br />
of substrates. Sourie (1954b) found it on shelly<br />
sand in the Baie de Dakar and Longhurst (1958)<br />
reported in from muddy sand in 22 m off Dakar.<br />
Uschakov (1970) reported it from hard sand bottom<br />
in depths greater than 20 m in clear water.<br />
It was reported from rocks ornamented with gorgonians<br />
in 10 m off Pointe-Noire, and was taken<br />
by the Calypso on mud and shells in 18 m, on mud<br />
in 18-30 m and 50 m, on muddy sand with<br />
Foraminifera in 21-27 m, and on mud with Area<br />
in 32 m (Forest and Guinot, 1966). The Pillsbury<br />
specimens were taken on mud or mud with Jullienella.<br />
Ovigerous females have been recorded from all<br />
months (Monod, 1956; Forest and Guinot, 1966).<br />
DISTRIBUTION.—<strong>West</strong> Africa, from Mauritania<br />
southward to Gabon, including Ilha do Principe<br />
in the Gulf of Guinea; and southern Africa, from<br />
South Africa and Mozambique (Barnard, 1954);<br />
intertidal zone to a depth of 50 m. Monod (1956)<br />
summarized earlier records and reported material<br />
from Mauritania, Senegal, Gambia, Guinea,<br />
Guinea-Bissau, Sierra Leone, Ghana, Gabon, and<br />
Principe. In addition the species has been reported<br />
from the following localities:<br />
Senegal: Baie de Dakar (Sourie, 1954b). Dakar, 22 m<br />
(Longhurst, 1958).<br />
Guinea: No specific locality, in depths greater than 20 m<br />
(Uschakov, 1970). 09°40'N, 14°05'W, 18 m, and 09°36'N,<br />
13°57'W, 18-30 m (Forest and Guinot, 1966).<br />
Ivory Coast: 05°02.5'N, 05°25'W, 21-27 m (Forest and<br />
Guinot, 1966).<br />
Ghana: Tenkpobo (as Tenpobo), Christiansborg, Apam,<br />
Winneba and Axim, rarely sublittoral to 37 m (Gauld, 1960).<br />
04°40'N, 02°08'W to 04°39'N, 02°05'W, 50 m (Forest and<br />
Guinot, 1966).<br />
Nigeria: Off the mouths of the Niger River, 04°03'N,<br />
06°12'E, 32 m (Forest and Guinot, 1966).<br />
Gabon: W of Libreville, 20-40 m (Rossignol, 1962).<br />
Congo: Baie de Pointe-Noire, 6-8 m (Rossignol, 1962).<br />
Pointe-Noire, 10 m (Crosnier, 1969).<br />
Angola: Baia da Caota, Benguela, 13 m and 30 m (Guinot<br />
and Ribeiro, 1962).<br />
Genus Menippe de Haan, 1833<br />
Menippe de Haan, 1833:21 [type-species: Cancer rurnphii Fabricius,<br />
1798, by subsequent designation by Glaessner,<br />
1929:253; gender: feminine].—Guinot, 1971:1076 [list of<br />
species].<br />
Menippe nodifrons Stimpson, 1859<br />
Menippe nodifrons.—Frade, 1950:11, 26 [Congo].—Capart,<br />
1951:138, fig. 49 [Guinea].—Monod, 1956:222, figs. 244-
138<br />
248 [Senegal, Guinea, Sierra Leone, Ghana, Rio Muni;<br />
references].—Longhurst, 1957:374 [Sierra Leone].—Rossignol,<br />
1957:82 [Congo].—Buchanan, 1958:20 [Ghana].<br />
—Longhurst, 1958:88 [Sierra Leone].—Gauld and Buchanan,<br />
1959:127 [Ghana].—Gauld, 1960:70 [Ghana].—<br />
Rossignol, 1962:116 [Congo].—Guinot and Ribeiro, 1962:<br />
50 [Angola].—Monod, 1967:180, pi. 17: fig. 3 [no material].—Le<br />
Loeuff and Intes, 1968, table 2 [Ivory<br />
Coast].—Guinot, 1968b: 156 [discussion].—Uschakov,<br />
1970:445, 446, 447, 455 [listed; Guinea].—Guinot, 1971:<br />
1076 [listed].<br />
Menippe nanus.—Capart, 1951:140 [discussion].<br />
Menippe.— Gauld and Buchanan, 1959:128 [Ghana].<br />
SYNONYMS.—Menippe rudis A. Milne Edwards,<br />
1879; Menippe nanus A. Milne Edwards and Bouvier,<br />
1898.<br />
DISTRIBUTION.—Eastern Atlantic from the<br />
Cape Verde Islands and Senegal to Angola; western<br />
Atlantic from Florida to Brazil (Rathbun,<br />
1930); littoral and sublittoral, from shore to about<br />
20 m.<br />
Genus Microcassiope Guinot, 1967<br />
Microcossiope Guinot, 1967c:358 [type-species: Xanthodes rufopunctatus<br />
A. Milne Edwards, 1869, by original designation;<br />
gender: feminine]; 1971:1076 [list of species].<br />
* Microcassiope minor (Dana, 1852)<br />
FIGURE 30<br />
Xmtho minor Dana, 1852b: 169; atlas, 1855, pi. 8: fig. 7.—<br />
Miers, 1881a:214; 1886:124 [listed].<br />
Pilumnus granulimanus Stimpson, 1871a: 143 [Cuba].<br />
Muropanopt granulimanus.—Rathbun, 1930:439, pi. 180: figs.<br />
1, 2 [western Atlantic records].<br />
Micropanope granosa.—Chapman and Santler, 1955:374.<br />
Micropanope rufopunctata.—Monod, 1956:313, figs. 386-392.—<br />
Gauld, 1960:70.—Guinot and Ribeiro, 1962:59.—Monod,<br />
1963, fig.34 [no material].—Ribeiro, 1964:10—Forest and<br />
Guinot, 1966:81.—Chace, 1966:639, fig. 8.—Guinot,<br />
1967c:348 [discussion], 358 [listed].—Le Loeuff and Intes,<br />
1968, table 1—Tiirkay, 1976b:61 [listed], 69.<br />
Microcassiope rufopunctata.—Guinot, 1967c, figs. 10, 15; 1971:<br />
1076 [listed].<br />
Xanthodes rufopunctata.—Guinot, 1967c:359 [listed].<br />
Xanthodes rufopunctatus.—Garth, 1968:314 [discussion].<br />
Microcassiope granulimanus.— Guinot, 1971:1076 [listed; considered<br />
to be distinct from M. rufopunctata].<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
SYNONYMS.—Xanthodes rufopunctatus A. Milne<br />
Edwards, 1869; Xanthodes granosus A. Milne Edwards<br />
and Bouvier, 1898.<br />
MATERIAL EXAMINED.—Pillsbury Material: Annobon:<br />
Sta 282,18-37 m, nodular coralline algae, 3d, 59, 2 juv (L,<br />
W).<br />
Other Material: Madeira: Ponta de Sao Lourenco, 32°44'N,<br />
16°44'W, littoral, rocky shore with tide pools, 29 Feb 1976,<br />
Onversaagd Sta 16, 19 (L). SE coast, near Canical, 32°44'N,<br />
16°44'W, 0-22 m, shore collecting, snorkeling, diving, 11<br />
Mar 1976, Onversaagd Sta 48, Ic5 (L).<br />
Cape Verde Islands: Porto da Praia (as La Praya), Sao<br />
Tiago, 10-30 m,Jun-Jul 1883, Talisman, syntype of Xanthodes<br />
granosus A. Milne Edwards and Bouvier, 1898, Id, 19 (L, W).<br />
Annobon: S coast, 01°28.5'S, 05°37.5'E, 35-55 m, 16 Jun<br />
1967, F. Poinsard, 19 (W).<br />
DESCRIPTION.—A. Milne Edwards and Bouvier,<br />
1900:87.<br />
Figures: Monod, 1956, figs. 386-392.<br />
Male Pleopod: Monod, 1956, figs. 387-392<br />
(Azores, Cape Verde Islands); Chace, 1966, fig. 8<br />
(St. Helena, Curacao, Cape Verde Islands);<br />
Guinot, 1967c, fig. 15 (Cape Verde Islands).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 2.5 to 9.1 m.<br />
REMARKS.—In his account of the Crustacea of<br />
the U.S. Exploring Expedition, Dana (1852b: 169)<br />
described a small xanthid crab that had been<br />
collected at Madeira or the Cape Verde Islands,<br />
Xantho minor. His account is based on an ovigerous<br />
female 2.1 lines long, 3.1 lines wide (approximately<br />
4.5 X 6.6 mm):<br />
Near X. parvulus [= Eurypanopeus blanchardi\. Carapax anteriorly<br />
areolate, areolets slightly raised, 2M, 3M, 5L, 6L<br />
circumscribed behind; antero-lateral margin thin, fourtoothed,<br />
three posterior teeth subtriangular. Anterior feet of<br />
moderate size, subequal, carpus and hand above slightly<br />
granulous, hand exteriorly faint granulato-costate, and<br />
above sulcate, moveable finger not armed with a large basal<br />
tooth. Eight posterior feet sparsely pubescent.<br />
Two small species of xanthids, both formerly<br />
placed in Micropanope, are now known to live in<br />
the Cape Verde Islands and one or more of the<br />
more northern islands or island groups, Madeira,<br />
the Azores, and the Canary Islands: Microcassiope<br />
rufopunctata (A. Milne Edwards, 1869) and Nanocassiope<br />
melanodactyla A. Milne Edwards, 1867).<br />
Earlier authors have identified Xantho minor with
NUMBER 306 139<br />
FIGURE 30.—Microcassiope minor (Dana) (from Dana, 1852b,<br />
pi. 8: fig.7).<br />
one or the other of these species without adopting<br />
the specific epithet minor. A third small species of<br />
xanthid, also at one time placed in Micropanope,<br />
CoralHope parvula (A. Milne Edwards, 1869), also<br />
occurs in the Cape Verde Islands; that species, as<br />
noted above, has but three anterolateral teeth<br />
whereas X. minor was described as having four,<br />
and has an additional inner spine on the carpus<br />
of the chela; the spine is absent in X. minor.<br />
Miers (1886:124) listed Xantho minor with other<br />
species now assigned to Xantho sensu stricto, and<br />
retained both melanodactyla and rufopunctata in Xanthodes,<br />
the genus to which they were assigned in<br />
Milne Edwards' original accounts.<br />
A. Milne Edwards and Bouvier (1898:190), in<br />
their description of Xanthodes talismani [= Coralliope<br />
parvula], pointed out similarities between that species<br />
and Xantho minor:<br />
Cette espece se rapproche beaucoup du Xantho minor<br />
Dana, de Madere et des lies du Cap Vert; mais cette derniere<br />
appartient a un genre different; sa dent anterieure est encore<br />
nette, sa carapace est depourvue de touffes de poils, ses<br />
pinces sont un peu costulees et il n'y a pas d'epines sur le<br />
bord interne du carpe.<br />
Later A. Milne Edwards and Bouvier (1900:<br />
93) repeated this observation in their account of<br />
Xanthodes talismani, in spite of the fact that on p.<br />
85 of the same work they identified Xantho minor<br />
Dana, 1852 with Xanthodes melanodactylus A. Milne<br />
Edwards, 1867 (and used the junior synonym!)<br />
and on page 87 of the same work reported Xan-<br />
thodes melanodactylus var. rufopunctata and noted: "il<br />
y a probablement lieu d'y rattacher [that mate-<br />
rial] plus specialement le Xantho minor de Dana."<br />
They also synonymized Dana's Xantho parvulus,<br />
which we believe should be identified with Eurypanopeus<br />
blanchardi, with Xanthodes melanodactylus.<br />
Monod (1956:320, in synonymy) identified<br />
Xanthodes melanodactylus var. rufopunctatus of A.<br />
Milne Edwards and Bouvier, 1900, with Micropanope<br />
melanodactyla. Three females from one of<br />
the lots (in 75 m) reported by A. Milne Edwards<br />
and Bouvier and labeled Xanthodes melanodactylus<br />
var. rufopunctata are in the collection of the <strong>Smithsonian</strong><br />
<strong>Institution</strong>; they are Nanocassiope melanodactyla.<br />
Monod (1933b:519) pointed out the multiple<br />
use of the epithet minor by A. Milne Edwards and<br />
Bouvier and further noted: "En fait la figure de<br />
Xantho minor donnee par DANA semblerait plutot<br />
representer Xanthias granosus [ = Microcassiope rufopunctata]<br />
ou X. tuberculatus [— Monodaeus couchii]<br />
que X. melanodactylus. L'examen du type est indispensable."<br />
We believe that Monod (1956:314, synonymy)<br />
was correct in identifying Xantho minor Dana with<br />
Xanthodes rufopunctatus A. Milne Edwards. Dana's<br />
account mentioned the small size, the wellmarked<br />
regions, the triangular anterolateral teeth<br />
of the carapace, the granular chela lacking a large<br />
basal tooth, and, in addition, clearly showed in<br />
his plate 8: figure 7 (see Figure 30) several transverse<br />
lines of granules on the carapace. All of<br />
these features are characteristic of the species now<br />
known as Microcassiope rufopunctata. We believe<br />
that the older name, Microcassiope minor (Dana,<br />
1852), must be used for the species.<br />
Chace (1966:639), in recording this species<br />
from Saint Helena in the south central Atlantic,<br />
showed that Pilumnus granulimanus Stimpson, 1871,<br />
from the western Atlantic, placed in Micropanope<br />
by Rathbun (1930), is conspecific with M. minor<br />
(as M. rufopunctata); the latter is now known to<br />
have a relatively wide range on both sides of the<br />
Atlantic. Guinot (1971:1076), in her summary of<br />
nomenclatural changes resulting from her studies<br />
of the xanthids, removed M. granulimanus from the<br />
synonymy of M. minor without stating her reasons.<br />
She noted: "Cette espece a ete mise en synonymie
140<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
avec Micropanope rufopunctata par CHACE (1966, p. insohn and Holthuis, 1964:59, for records), the<br />
639, fig. 8), mais pour l'instant nous la mainten- Azores, the Canary Islands, Madeira, the Cape<br />
ons separee de l'espece ouest-africaine."<br />
Verde Islands, the <strong>African</strong> mainland from Span-<br />
In discussing the distribution patterns of some ish Sahara, Ivory Coast, and Ghana, and the<br />
Atlantic-East Pacific xanthids, Garth (1968:314) offshore islands of the Gulf of Guinea, Principe,<br />
was in error in stating that Microcassiope did not Sao Tome, and Annobon; central Atlantic from<br />
occur in the western Atlantic. It is represented Saint Helena (Chace, 1966); western Atlantic<br />
there by M. minor (as Micropanope granulimanus). from the Bahamas, Cuba, Curacao, and Islas Los<br />
Garth apparently was misled by the omission of Roques (Chace, 1966); intertidal to a depth of ca<br />
this species in Guinot's (1967c:358) original ac- 220 m, commoner in shallower water. Monod<br />
count of Microcassiope.<br />
(1956) summarized earlier <strong>West</strong> <strong>African</strong> records<br />
A male syntype of Xanthodes granosus (see "Syn- and reported material from the Azores, the Caonyms")<br />
is in the collection of the National Munary Islands, the Cape Verde Islands, and Ghana.<br />
seum of Natural History, <strong>Smithsonian</strong> Institu- Other <strong>West</strong> <strong>African</strong> records in the literature intion,<br />
under catalog number USNM 22956. A clude the following:<br />
female syntype is in the collection of the Rijksmuseum<br />
van Natuurlijke Historie, Leiden, under<br />
Crust. D.1564.<br />
<strong>West</strong> Africa: No specific locality (Monod, 1963).<br />
Azores: Pasteleiro, Feteira, and Horta, Ilha do Faial,<br />
intertidal (Chapman and Santler, 1955).<br />
BIOLOGY.—Off <strong>West</strong> Africa, Microcassiope minor Madeira: Porto do Funchal, ca 220 m, on telegraph cable;<br />
lives from the intertidal zone to a depth of ca<br />
Ilheu do Gorgulho, littoral; between Ponta da Garajau and<br />
Ponta da Oliveira, 5-6 m (all Tiirkay, 1976b).<br />
220 m. Of the recent records for which depth is<br />
Cape Verde Islands: No specific locality (Guinot, 1967c).<br />
available, 19 out of 26 (or 73%) are from depths Baixo Joao Leitao, 25 m (Guinot and Ribeiro, 1962; Ribeiro,<br />
of 10 m or less, and seven (or 27%) are from 1964). Porto da Praia (as La Praya), Sao Tiago (Chace,<br />
depths ranging from between 15 and 45 m to ca 1966).<br />
220 m (15-40 m, 43-45 m, and ca 220 m). Eight<br />
of the 19 shallower collections are from shore.<br />
Forest and Guinot (1966:84) pointed out that<br />
Nanocassiope melanodactyla (as Micropanope melanodactyla)<br />
and Microcassiope minor both are common<br />
in the Gulf of Guinea, where they may occur<br />
Spanish Sahara: 21°05'N, 17°14'W, 43-45 m (Forest and<br />
Guinot, 1966).<br />
Ivory Coast: Lagoon of Abidjan, 05°16'N, 04<br />
together, but that the former lives in deeper water<br />
generally on less stable substrates whereas "la<br />
seconde, plus littorale, a souvent ete capturee<br />
dans la zone intercotidale ou jusqu'a une dizaine<br />
de metres, principalement sur des fonds durs,<br />
roches et coraux," a statement borne out by the<br />
capture of these two species off the offshore islands<br />
of the Gulf of Guinea. The deepest record is that<br />
of Tiirkay (1976b) from Madeira, in ca 220 m on<br />
a telegraph cable; he also recorded specimens<br />
from the littoral zone, under stones.<br />
Off <strong>West</strong> Africa, ovigerous females have been<br />
collected in June (Forest and Guinot, 1966).<br />
Chace (1966) recorded ovigerous females from<br />
Saint Helena in January and April.<br />
DISTRIBUTION.—Atlantic Ocean. Eastern Atlantic<br />
from the eastern Mediterranean (see Lew-<br />
o 01'20 w W,<br />
(Forest and Guinot, 1966). Off Jacqueville, 40 m (Le Loeuff<br />
and Intes, 1968).<br />
Ghana: Off Tenkpobo (as Tenpobo), littoral; off Accra,<br />
37 m (Gauld, 1960).<br />
Principle: Ilheu Caroco, 2-8 m; Ponta da Mina, beach<br />
(Forest and Guinot, 1966).<br />
Sao Tome: 00°20'N, 06°46'E, 10 m; Ponta Diogo Vaz,<br />
W coast, 0-6 m; Praia de Santa Catarina, W coast, 3-10 m;<br />
off Sao Tome, 8 m; off Ponta Diogo Nunes, shore and<br />
4-5 m; in front of Ponta Oquedelrei, 6 m; Morro Peixe, 2-<br />
6 m; Ilheu das Cabras, shore; in front of the harbormaster's<br />
office, shore; in front of Praia Lagarto, 5-6 m (all Forest and<br />
Guinot, 1966).<br />
Annobon: No specific locality (Guinot, 1967c). 01°24'<br />
04"S, 05°36'45"E, 7-10 m; O1°25'12"S, 05°36'05"E, 20 m;<br />
Isla Tortuga, NW coast, 15-40 m (all Forest and Guinot,<br />
1966).<br />
Saint Helena: Rupert's Bay, 0-75 m; James Bay (Chace,<br />
1966).<br />
Genus Monodaeus Guinot, 1967<br />
Monodaeus Guinot, 1967c:369 [type-species: Xantho couchii<br />
Couch, 1851, by original designation; gender: masculine];<br />
1971:1074 [ list of species].
NUMBER 306 141<br />
Monodaeus couchii (Couch, 1851)<br />
Xantho couchii Couch, 1851:13.<br />
Medaeus couchi.—Monod, 1956:310, figs. 383, 877, 878.—<br />
Peres, 1964:20.—Forest and Guinot, 1966:80.—Crosnier,<br />
1967:331, figs. 13, 14.—Zariquiey Alvarez, 1968:400, figs.<br />
9, 15e, 133, 134a [Spain; references].—Crosnier, 1970:1215<br />
[listed], 1216.<br />
Micropanope (?) couchi.—Forest and Gantes, 1960:352.<br />
Xantho couchi.—Peres, 1964:27, 28, 29.—Guinot, 1967c:348,<br />
349, 371 [all discussion], 373 [listed].—Maurin, 1968a: 19,<br />
116 [Spain, Mediterranean].—Forest, 1976:66 [discussion].<br />
Monodaeus couchi.—Guinot, 1967c:372 [discussion], figs. 23,<br />
32 [Spain]; 1971:1074 [listed].—Turkay, 1976a:25 [listed],<br />
38, fig. 24 [Portugal, in part].—Forest, 1976:68 [discus-<br />
SYNONYM.—Xantho tuberculatus Bell, 1852.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Undaunted Material: South-<strong>West</strong> Africa: Sta 106, 225 m, 1$<br />
(L).<br />
Other Material: Morocco: Off Cap de Mazagan, 33°40'N,<br />
08°45'W, Agassiz trawl, 28 Mar 1976, Onversaagd Sta 154, 1
142<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 68, 70 m, broken shell, 16* (W).<br />
Ivory Coast: Sta 45, 73-97 m, 16* (L). Sta 49, 73-77 m,<br />
16*, fragments (L). Sta 50, 128-192 m, 1(5, 2$ ov (W). Sta<br />
62, 46 m, brown, branched and foliate Foraminifera, 16*, 1$<br />
(L).<br />
Nigeria: Sta 237, 101 m, coral ground, rough, 16* (L). Sta<br />
239, 73 m, 19 (L).<br />
Other Material: Ivory Coast: Off Sassandra, 11 m, 3 Apr<br />
1964, Guinean Trawling Survey, Tr 22, Sta 1, 16* (L).<br />
04°33'N, 06°36'W, 100-109 m, sand, mud, shells, rocks, 21<br />
May 1956, Calypso Sta 16, 26*, 39 (1 ov) (W).<br />
Congo: Off Pointe-Noire, 04°56'S, 11°31'E, 95-97 m,<br />
trawl, 21-22 Sep 1965, Id (W).<br />
DESCRIPTION.—Capart, 1951:153.<br />
Figures: Capart, 1951, fig. 57, pi. 3: fig. 17;<br />
Monod, 1956, figs. 384, 385; Guinot and Ribeiro,<br />
1962, fig. 26; Crosnier, 1967, figs. 25, 26.<br />
Male Pleopod: Capart, 1951, pi. 3: fig. 17<br />
(Congo); Monod, 1956, fig. 385 (Congo); Guinot<br />
and Ribeiro, 1962, fig. 26 (Ivory Coast).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 5 to 15 mm; the carapace widths<br />
of ovigerous females is 8 mm. Crosnier (1967:336)<br />
pointed out that this is a relatively small species;<br />
he examined one ovigerous female 6.6 mm wide,<br />
and the ovigerous female reported by Capart<br />
(1951) was only 6 mm wide.<br />
BIOLOGY.—Monodaeus rouxi is an offshore species<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Ivory Coast: SW of Grand-Bassam, 100-400 m (Crosnier, which lives on the continental shelf and upper<br />
1967).<br />
slope. It usually occurs in depths between 46 m<br />
Congo: 51.5 mi [83 km] WNW of Banana [Zaire],<br />
05°50'S, 11°32'E, 215-220 m (Capart, 1951). Off Pointe-<br />
Noire, 80-100 m (type-locality) (Crosnier, 1967); in 255 m<br />
(Crosnier, 1969). 04°53'S, 11°38'E, 75 m; 05°00'S, 11°26'E,<br />
115 m (Crosnier, 1969).<br />
and 215-220 m; the extremes of depth recorded<br />
so far, 11 m herein and ?500 m (Guinot and<br />
Ribeiro, 1962), require verification. Nine of the<br />
14 available depth records are for depths below<br />
100 m, and seven of those are for depths between<br />
64 and 97 m. The five deeper records are 100-<br />
* Monodaeus rouxi (Capart, 1951)<br />
109 m, 101 m, 128-192 m, 215-220 m, and<br />
Micropcmope rouxi Capart, 1951:153, fig. 57, pi. 3: fig. 17.— ?500 m. The species has been collected on brown<br />
Forest, 1965a:380.—Forest and Guinot, 1966:81.— mud in 215-220 m and on muddy sand and rocks<br />
Guinot, 1967c:348, 349 [discussion], 371 [listed].—Forest, in 80 m (Capart, 1951); mud, sand and com-<br />
1976:66 [discussion].<br />
pacted sand (sable construit) in 65-75 m, sand,<br />
Medaeus (?) rouxi.—Monod, 1956:312, figs. 384, 385.—Guinot<br />
mud, and shells in 64 m, and on sand, mud,<br />
and Ribeiro, 1962:58, fig. 26.—Crosnier, 1967: 335, figs.<br />
25, 26. [Not Medaeus rouxi Balss, 1935.]<br />
shells, and rocks in 100-109 m by the Calypso<br />
Medaeus rouxi.—Forest, 1959:15 [not Medaeus rouxi Balss, (Forest and Guinot, 1966). In the latter depth a<br />
1935].<br />
large series of 56 specimens was collected. The<br />
Monodaeus rouxi.— Guinot, 1967c:371, 372, 373 [all discus- specimens taken by the Pillsbury were taken on<br />
sion], fig. 24; 1971:1074 [listed].—Forest, 1976:68 [discusion].<br />
brown, branching, and foliate Foraminifera in 46<br />
m, on broken shell in 70 m, and on coral on rough<br />
bottom in 101 m.<br />
Ovigerous females have been collected in May<br />
and December (Capart, 1951; Forest and Guinot,<br />
1966; Pillsbury).<br />
DISTRIBUTION.—<strong>West</strong> Africa, from scattered localities<br />
between Senegal and Angola, in depths<br />
between 46 m and 215-220 m (11 m to ?500 m).<br />
No material was available to Monod (1956). Records<br />
in the literature include the following:<br />
<strong>West</strong> Africa: No specific locality (Forest, 1976).<br />
Senegal: No specific locality (Forest, 1965a); in 65-75 m<br />
(Forest, 1959). 12°55.5'N, 17°33'W, 65-75 m (Forest and<br />
Guinot, 1966).<br />
Liberia: 04°34.5'N, 08°31'W, 64 m (Forest and Guinot,<br />
1966).<br />
Ivory Coast: No specific locality (Guinot and Ribeiro,<br />
1962; Forest, 1965; Crosnier, 1967). 04°33'N,06°36'W, 100-<br />
109 m (Forest and Guinot, 1966).<br />
Congo: 51.5 mi [83 km] WNW of Banana [Zaire],<br />
05°50'S, 11°32'E, 215-220 m (Capart, 1951). Off Pointe-<br />
Noire (Crosnier, 1967).<br />
Angola: 11 mi [18 km] W of Cap Ledo, 09°40'S, 13°02'E,<br />
80 m (Capart, 1951). Benguela, ?500 m (Guinot and Ribeiro,<br />
1962).<br />
Genus Nanocassiope Guinot, 1967<br />
Nanocossiope Guinot, 1967c:355 [type-species: Xanthodes melonodactylus<br />
A. Milne Edwards, 1867, by original designation;
NUMBER 306<br />
gender: feminine]; 1971:1075 [list of species].—Takeda,<br />
1976:85 [definition].<br />
* Nanocassiope melanodactyla<br />
(A. Milne Edwards, 1867)<br />
Micropanope polita Rathbun, 1893b:238; 1930:440, fig. 70, pi.<br />
180: figs. 3, 4.—Garth, 1946:459, pi. 77: fig. 4 [eastern<br />
Pacific; considered distinct by Guinot, 1971].<br />
Panopeus tanneri Faxon, 1893:154 [eastern Pacific].<br />
143<br />
Cape Verde Islands: Sao Vicente, 75 m, 29 Jul 1883,<br />
Talisman, 39 (W) [labelled Xanthodes melanodactylus var rufopunctatus].<br />
Annobon: 01°24'S, 05°37.5'E, 11 Dec 1965, A. Crosnier,<br />
26, 39 (W). O1°27'S, 05°35'50"E, 50-60 m, 11 Dec 1965, A.<br />
Crosnier, 26*, 39 (W). O1°27'S, 05°35'48 w E, 50-60 m, 11 Dec<br />
1965, Ombango, A. Crosnier, 16*, 89 (2 ov) (W).<br />
DESCRIPTION.—Capart, 1951:152.<br />
Figures: Capart, 1951, fig. 56, pi. 3: fig. 16;<br />
Xanthodes melanodactylus var. rufopunctatus.—A. Milne Edwards Monod, 1956, figs. 401-405.<br />
and Bouvier, 1900:87, pi. 16: figs. 4, 5 [part?] [not Xan- Male Pleopod: Capart, 1951, pi. 3: fig. 16 (Anthodes<br />
rufopunctatus A. Milne Edwards, 1869 = Xantho minor<br />
gola); Monod, 1956, figs. 403-405 (Senegal);<br />
Dana, 1852].<br />
Micropanope melanodactylus.—Capart, 1951:151, fig. 56, pi. Chace, 3; 1966, fig. 7 (Cape Verde Islands, Baja<br />
figs. 15, 16.—Chace, 1966:637, fig. 7 [Saint Helena]. California, Cocos Island); Guinot, 1967c, fig. 13<br />
Micropanope melanodactyla.—Monod, 1956:320, figs. 401- (Senegal).<br />
405.—Gauld, 1960:70.—Guinot and Ribeiro, 1962:60.— Color: "Coloration dans l'alcool, gris tachete de<br />
Ribeiro, 1964:12.—Forest and Guinot, 1966:83.—Guinot,<br />
rose; les doigts des chelipedes parfois bruns, par-<br />
1967c:348 [discussion], 355 [listed].—Le Loeuff and Intes,<br />
1968, table 1.<br />
fois noirs" (Capart, 1951:152).<br />
Xanthodes melanodactylus.—Guinot, 1967c:358 [listed]. MEASUREMENTS.—Our specimens have cara-<br />
Nanocassiope melanodactyla.—Guinot, 1967c, figs. 8, 13; 1971: pace widths of 2.5 to 13 mm; the carapace widths<br />
1075 [listed].—Tiirkay, 1976b:61 [listed], 68.<br />
of ovigerous females ranges from 4 to 9 mm.<br />
Nanocassiope polita.—Guinot, 1971:1075 [listed].<br />
REMARKS.—Chace (1966:637-638) reported<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 70, 33 m, branched Foraminifera, 36*, 1$ ov (L).<br />
Micropanope melanodactylus from Saint Helena in<br />
the south central Atlantic and noted: "I agree<br />
Ivory Coast: Sta 42, 62-75 m, mud with brown, branched with Monod (1956, p. 324) that M. melanodactylus<br />
Foraminifera, 16* (W). Sta 46, 38-42 m, mud with dense<br />
Jullienella, 16*, 29 (1 ov) (L). Sta 47, 37 m, bottom with<br />
Jullienella, 66, 9$ (3 ov) (L).<br />
Ghana: Sta 22, 51 m, rough bottom, 16*, 1? (L). Sta 23,<br />
42 m, foliate brown to orange bryozoans, 136*, 79 (1 ov) (L).<br />
Sta 24, 35-37 m, dark red bryozoans, 26*, 179 (6 ov), 1 juv<br />
is probably not a synonym of Xantho minor Dana,<br />
1852b. Unfortunately, the type-specimen of<br />
Dana's species is probably no longer extant, and<br />
his name is therefore likely to remain a nomen<br />
dubium indefinitely." We agree that Xantho minor<br />
(L,W).<br />
cannot be identified with Xanthodes melanodactylus.,<br />
Nigeria: Sta 250, 24 m, brackish water, mud, 19 (W).<br />
Annobon: Sta 271, shore, sand beach, 16* (W). Sta 275,<br />
9-69 m, rubble of coralline algae, 86, 89 (1 ov), 2 juv (L).<br />
Sta 282, 18-37 m, nodular coralline algae, 26, 19 (L, W).<br />
Sta 283, 51-55 m, nodular coralline algae, 426\ 369 (7 ov)<br />
we have identified it with Microcassiope rufopunctata,<br />
above.<br />
We believe that Chace (1966:637) correctly<br />
synonymized Micropanope polita Rathbun, 1893,<br />
(W). Sta 284, 73 m, black basaltic rocks, 29 ov (L). from the eastern Pacific, with M. melanodactyla.<br />
Other Material: Madeira: Porto da Abra, SE coast, Guinot (1971:1075) did not accept this action,<br />
32°45'N, 16°41'W, to 12 m, diving, 13 Mar 1976, Onversaagd apparently preferring, without stating her rea-<br />
Sta 68, 16 (L). Near Canical, SE coast, 32°44' N, 16°44' W,<br />
sons, to keep the two taxa separate. As Garth<br />
0-22 m, shore collecting, snorkeling, diving, 10 Mar 1976,<br />
Onversaagd Sta 39, 1 juv
144<br />
this species. The front is transversely grooved in<br />
all specimens examined by us.<br />
BIOLOGY.—Nanocassiope melanodactyla is a sublittoral,<br />
usually shallow shelf species that apparently<br />
prefers a bottom with coralline algae or Foraminifera<br />
on mud or muddy sand. Of 70 recent depth<br />
records for the species off <strong>West</strong> Africa, 56 or 80%<br />
are from depths of 50 m or less; two of those are<br />
from shore, the remainder in depths from 3-11 m<br />
to 40-54 m or 9-69 m. Of the 14 records from<br />
deeper water, two, 200 m and 225 m, are from<br />
depths greater than 100 m and the remainder<br />
range from 60 to 85 m. The deepest records in<br />
the literature are in A. Milne Edwards and Bouvier<br />
(1900), who provided 11 depth records. Only<br />
three of these are from depths of less than 50 m,<br />
three are from depths generally more than 50 m<br />
but less than 100 m (75, 80, and 80-115 m), and<br />
the remainder are 100-150 m, 110-180 m, 225 m,<br />
355 m, and 627 m. The reliability of these deeper<br />
records is questionable, but the species may stray<br />
into water that deep. Subsequently, Tiirkay<br />
(1976b) recorded it from 80 m on a telephone<br />
cable in the harbor of Funchal; in other samples<br />
he recorded it from blocks of stone. It generally<br />
occurs in depths between 20 and 50 m, as pointed<br />
out by Forest and Guinot (1966:84). They compared<br />
the apparent habitat preferences of N.<br />
melanodactyla and Microcassiope minor and noted:<br />
"La premiere a ete recueillie a des profondeurs<br />
comprises entre 9 et 65-75 m, mais surtout entre<br />
20 et 50 m, sur des fonds meubles, le plus souvent<br />
parmis les algues calcaires."<br />
Apparently this species spawns all year off<br />
<strong>West</strong> Africa. Ovigerous females have been collected<br />
in all months but April, August, and December<br />
(Capart, 1951; Monod, 1956; Guinot and<br />
Ribeiro, 1962; Ribeiro, 1964; Forest and Guinot,<br />
1966; Pillsbury).<br />
DISTRIBUTION.—Eastern Pacific, from Baja California,<br />
Mexico, Cocos Island, and the Galapagos<br />
Islands; central Atlantic, from Saint Helena and<br />
Ascension Island; and eastern Atlantic, from the<br />
Azores, Madeira, Ilhas Desertas, the Canary Islands,<br />
the Cape Verde Islands and Senegal south-<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
ward to Angola, including the offshore islands of<br />
the Gulf of Guinea, Principe, Sao Tome, and<br />
Annobon; sublittoral, subtidal to a depth of more<br />
than 600 m. Monod (1956) summarized the literature<br />
and reported material from the Azores,<br />
Madeira, the Cape Verde Islands, the mainland<br />
from localities between Senegal and Gabon, and<br />
from the offshore islands of Principe and Annobon;<br />
other records, including those published<br />
since 1956, include the following.<br />
Madeira: No specific locality (Dana, 1852b; Miers,<br />
1881a). Funchal harbor, ca. 80 m; near Ponta da Garajau;<br />
between Ponta da Garajau and Ponta da Oliveira; and<br />
Ponta de Sao Louren^o (all Tiirkay, 1976b).<br />
Cape Verde Islands: No specific locality (Dana, 1852b).<br />
Porto Grande, Sao Vicente (Chace, 1966). Baia de Fateixa,<br />
Sao Vicente, shore; Baia de Porto Grande, Sao Vicente,<br />
4-6 m, 8 m, 8-10 m, 3-11 m (3.5-11 m); Baia do Tarrafal,<br />
Sao Tiago, 14-23 m and 9-17 m; Porto de Sao Francisco,<br />
Sao Tiago, 9 m; Porto da Furna, Brava, 6-20 m (all Guinot<br />
and Ribeiro, 1962; Ribeiro, 1964). Baia de Porto Novo,<br />
Santo Antao, 12 m; Porto Ingles, Maio (Ribeiro, 1964).<br />
Senegal: No specific locality (Guinot, 1967c). 12°55.5'N,<br />
17°33'W, 65-75 m (Forest and Guinot, 1966).<br />
Guinea-Bissau: 10°19'N, 16°34'W, 60-73 m (Forest and<br />
Guinot, 1966).<br />
Ivory Coast: Off Sassandra, 100 m (Le Loeuff and Intes,<br />
1968).<br />
Ghana: Off Accra, 10 m (Gauld, 1960). 04°40'N,<br />
02°08'W, 48 m; 04°40'N, 02°08'W to 04°39'N, 02°05'W)<br />
50 m; and 04°36.5'N, 01°31'W, 50 m (Forest and Guinot,<br />
1966).<br />
Principe: No specific locality (Guinot, 1967c). 01°35'N,<br />
07°28'E, 45 m; 01°38'25"N, 07°22'05"E, 31 m; 01°38'35"N,<br />
07°21'35"E, 35 m; 01°43'10"N, 07°28'20"E, 73 m; 01°43'N,<br />
07°28'55"E, 37 m; off Tinhosa Grande (as Hermano<br />
Grande) Island, 12 mi [19 km] SSW of Principe, 01°20'45''N,<br />
O7°17'37"E, 25-40 m; in front of [Cais de] Santana, 11 m<br />
(all Forest and Guinot, 1966).<br />
Sao Tome: 00°20'N, 06°47'E, 40-54 m; 00°20'N,<br />
06°46'E, 10 m; 00°25'40"N, 06°40'10"E, 50 m; 00°25'15"N,<br />
06°43'05"E, 8-30 m; in front of Ponta de Sao Sebastiao,<br />
11 m (all Forest and Guinot, 1966).<br />
Annobon: 01°27.5'S, 05°36.5'E, 35 m; 01°26'15"S,<br />
05°35'40"E, 60 m; 01°25'10"S, 05°36'10"E, 20-25 m; N of<br />
San Antonio, 9 and 23 m (all Forest and Guinot, 1966).<br />
Angola: Baia Farta, Benguela, 22-28 m (Guinot and<br />
Ribeiro, 1962).<br />
Saint Helena: Off Rupert's Bay, 0-75 m, 0-2 m (Chace,<br />
1966).
NUMBER 306 145<br />
Genus Nanopilumnus Takeda, 1974<br />
Nanopilumnus Takeda, 1974:215 [type-species: Medaeus rouxi<br />
Balss, 1935; gender: masculine].<br />
* Nanopilumnus boletifer (Monod, 1956)<br />
Pampilumnus boletifer Monod, 1956:260, fig. 302; 1963, fig. 36<br />
[no material].—Forest and Guinot, 1966:72, fig. 5a,b—<br />
Takeda, 1974:216 [listed; transferred to Nanopilumnus].<br />
MATERIAL EXAMINED.—Pillsbury Material: Annobon,<br />
Sta 275, 9-69 m, rubble of coralline algae, 1$ ov (L).<br />
DESCRIPTION.—Monod, 1956:260.<br />
Figure: Monod, 1956, fig. 302.<br />
Male Pleopod: Forest and Guinot, 1966, fig. 5a,b<br />
(Sao Tome).<br />
Color: Cream (Monod, 1956:261).<br />
MEASUREMENTS.—Our ovigerous female has a<br />
carapace width of 5 mm.<br />
BIOLOGY.—Nanopilumnus boletifer is a sublittoral<br />
species, with a recorded depth range extending<br />
from shore to 9-69 m. All depth records other<br />
than that of the Pillsbury are from depths of less<br />
than 12 m. The Pillsbury specimen was collected<br />
in the rubble of coralline algae. The following<br />
bottom types were noted for the Calypso collections<br />
(Forest and Guinot, 1966): rocks and coral;<br />
rocks and sand; mud and calcareous algae; and<br />
calcareous algae.<br />
Ovigerous females have been collected in May,<br />
June, and July (Forest and Guinot, 1966;<br />
Pillsbury).<br />
DISTRIBUTION.—<strong>West</strong> Africa, from the offshore<br />
islands of the Gulf of Guinea, Annobon (the typelocality)<br />
, Principe, and Sao Tome, from shore to<br />
a depth of 9-69 m, usually in 12 m or less.<br />
Records in the literature include the following:<br />
Annobon: No specific locality, in 12 m (Monod, 1956).<br />
Principe: Ilheu Caroc.o, 2-8 m; Ilheus dos Mosteiros, 3-<br />
10 m (Forest and Guinot, 1966).<br />
Sao Tome: Off Sao Tome, 8 m; Baia de Ana de Chaves,<br />
5 m; W of Ponta Diogo Nunes, shore; off Ponta Diogo<br />
Nunes, 4-5 m; in front of Ponta Oquedelrei, 6 m; Ilheu das<br />
Cabras, 0-2 m; and in front of Praia Lagarto, 5-6 m (all<br />
Forest and Guinot, 1966).<br />
Genus Panopeus H. Milne Edwards, 1834<br />
Panopeus H. Milne Edwards, 1834:403.<br />
REMARKS.—When H. Milne Edwards (1834:<br />
403) erected the genus Panopeus, he assigned two<br />
species to it, viz., Panopeus herbstii (a new species)<br />
and Panopeus limosus (a new combination based on<br />
Cancer limosa Say, 1817, a species now known as<br />
Eurytium limosum (Say)). Two other species were<br />
doubtfully assigned by H. Milne Edwards to the<br />
new genus: Cancer trispinosus Herbst, 1803, and<br />
Cancer ochtodes Herbst, 1783. In the synonymy of<br />
his Panopeus herbstii H. Milne Edwards cited Cancer<br />
panope Herbst (1801:40, pi. 54: fig. 5) and Say's<br />
(1817:58, pi. 4: fig. 3) use of Herbst's name Cancer<br />
panope for an East American species. H. Milne<br />
Edwards indicated that his material of Panopeus<br />
herbstii came from "les cotes de l'Amerique septentrionale."<br />
H. Milne Edwards did not indicate a typespecies<br />
for his genus Panopeus, and the first valid<br />
type selection that we know of is the one by E.<br />
Desmarest (1852:17), who stated: "Panopeus:<br />
genre americain, ayant pour type le Cancer panope,<br />
Herbst, que M. Milne Edwards nomme Panope<br />
d'Herbst." The type-species of Panopeus thus is<br />
Cancer panope Herbst, 1801, as H. Milne Edwards,<br />
1834, cited that species by name in the synonymy<br />
of Panopeus herbstii. That this type selection was<br />
most unfortunate is shown below.<br />
As was first pointed out by S. I. Smith (1869a:<br />
278), Cancer panope Herbst, 1801, is an Indo-<strong>West</strong><br />
Pacific species, different from the East American<br />
form which was figured by Say (1817) and described<br />
by H. Milne Edwards (1834). Von Martens<br />
(1872:87) examined Herbst's type material<br />
and came to the conclusion that Herbst's Cancer<br />
panope belongs to the genus Menippe de Haan,<br />
1833; he also showed that the type locality of<br />
Cancer panope is Tranquebar, India, where the<br />
material was collected by Ingobert Karl Daldorff,<br />
an officer of the Danish garrison at Tranquebar<br />
(1790-1793) and a pupil of J. C. Fabricius for<br />
whom he collected (cf., Zimsen, 1964:12). Balss<br />
(1932:513) removed Herbst's species from Menippe
146<br />
and placed it in the genus Sphaerozius; he pointed<br />
out that Sphaerozius panope is a rare species; apart<br />
from the type only three specimens were known<br />
to him.<br />
Unpublished notes in the Division of Crustacea,<br />
National Museum of Natural History, <strong>Smithsonian</strong><br />
<strong>Institution</strong>, Washington, D.C., made by<br />
the late Mary J. Rathbun, show that Cancer panope<br />
Herbst, 1801, is a junior subjective synonym of<br />
Cancer scaber Fabricius (1798:336). Miss Rathbun<br />
came to this conclusion after having examined<br />
the type-specimens of both species. Like Herbst's<br />
material of C. panope, the type-specimens of Cancer<br />
scaber were collected in India by Daldorff, and it<br />
thus is possible that the types of both species<br />
originally formed part of a single lot. The correct<br />
name of the species thus should be Sphaerozius<br />
scaber (Fabricus, 1798).<br />
The acceptance of Desmarest's (1852) selection<br />
of Cancer panope Herbst as the type-species of<br />
Panopeus H. Milne Edwards, would cause an enormous<br />
confusion. Not only would the generic name<br />
Sphaerozius Stimpson, 1858 (type-species, designated<br />
in Opinion 85, <strong>Smithsonian</strong> Miscellaneous Collections,<br />
73(3): 17, 1925, Sphaerozius nitidus Stimpson,<br />
1858) have to be replaced by the generic<br />
name Panopeus H. Milne Edwards, the well known<br />
and widely distributed genus of mud crabs now<br />
known as Panopeus would have to be given a<br />
different generic name. This genus is known from<br />
the eastern Atlantic (south coast of Portugal to<br />
Angola), both coasts of America (Massachusetts<br />
to Brazil and Lower California to Chile), and<br />
Hawaii; it consists of about 12 species, most of<br />
which live in the littoral zone and several are very<br />
common. The name Panopeus has been very consistently<br />
used for it. The only other name available<br />
for it is Eupanopeus Rathbun (1898:273; typespecies,<br />
by original designation, Panopeus herbstii<br />
H. Milne Edwards, 1834), a name which has not<br />
been used since 1908.<br />
In order to prevent the considerable confusion<br />
that a strict application of the Code would cause,<br />
Holthuis (1979b) has applied to the International<br />
Commission on Zoological Nomenclature to use<br />
their plenary power to set aside all type-selections<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
for the genus Panopeus. Having done so, he also<br />
asked the Commission to select as the type-species<br />
of that genus the species Panopeus herbstii H. Milne<br />
Edwards, 1834. In the meantime, as prescribed<br />
by the Code, the name Panopeus will be used by<br />
us in the sense in which it is currently employed<br />
by zoologists.<br />
As pointed out above, Panopeus herbstii is a<br />
composite species, its syntypes consisting of (1)<br />
the North American material that H. Milne Edwards<br />
had before him when he described the<br />
species, (2) the North American material described<br />
and figured by Say (1817) under the<br />
name Cancer panope Herbst, and (3) the type-specimen<br />
of Cancer panope figured by Herbst (1801, pi.<br />
54: fig. 5). The material under (1) presumably is<br />
the species now generally known as Panopeus herbstii,<br />
that under (2) is a mixture of Panopeus herbstii<br />
and Neopanope texana sayi (Smith, 1869) (cf. Rathbun,<br />
1930:335, 369), and that under (3) is Sphaerozius<br />
scaber (Fabricius, 1798). In order to legalize<br />
the current use of the name Panopeus herbstii for<br />
the common large East American mud crab,<br />
Holthuis (1979b: 159) selected as the lectotype of<br />
Panopeus herbstii the specimen from oyster beds of<br />
the east coast of the United States, that Say<br />
(1817, pi. 4: fig. 3) figured.<br />
By these actions it will remain possible to use<br />
both the generic name Panopeus H. Milne Edwards,<br />
1834, and the specific epithet herbstii H.<br />
Milne Edwards, 1834, in the accustomed sense.<br />
* Panopeus africanus A. Milne Edwards, 1867<br />
Panopeus africanus.—Frade, 1950:11, 26.—Capart, 1951:148,<br />
fig. 54, pi. 3: fig. 3.—Monod, 1956:325, figs. 406-415.—<br />
Rossignol, 1957:82, 83.—Longhurst, 1957:374, 375, 380,<br />
382; 1958:88.—Gauld, 1960:70.—Forest and Gantes,<br />
1960:352.—Rossignol, 1962:118.—Guinot and Ribeiro,<br />
1962:61.—Forest and Guinot, 1966:84.—Zariquiey Alvarez,<br />
1968:404, fig. 134b [Spain; references].—Uschakov,<br />
1970:443, 444, 447, 455 [listed].—Hartmann-Schroder<br />
and Hartmann, 1974:15.—Powell, 1979:127.<br />
Panopeus sp.—Monod, 1956:335, figs. 435-438.<br />
Eupanopeus africanus.—Bott, 1964:30.<br />
MATERIAL EXAMINED.—Pillsbury Material: Nigeria:<br />
Sta 1, Lagos harbor, shore, 4
NUMBER 306 147<br />
Other Material: Liberia: Rock Spring, Monrovia, O. F.<br />
Cook, G. N. Collins, 1$ (W). Free Port area, Monrovia,<br />
oyster cultch, 22 Apr 1953, G. C. Miller, 16* (W). Locality<br />
same, 24 Apr 1953, 1$ ov (W).<br />
Dahomey: Lagoon of Lac Nokoue near Zogbo, W of<br />
Cotonou, 29 Mar 1964, H. Hoestlandt, 1? (L). Point XI<br />
Lagoon near Contonou, 29 Mar 1964, H. Hoestlandt, 16*<br />
(L). Zogbo Lagoon near Contonou, 10 Apr 1964, H.<br />
Hoestlandt, 1$ (L).<br />
Nigeria: Harbor of Lagos, 13 Jun 1963, A. R. Longhurst,<br />
16* (L). S bank of mouth of Escravos River near Ajudaibo,<br />
Niger delta, 05°34.5'N, O5°11.75'E, 30 Jul 1975, C. B.<br />
Powell, 1 large 6* (L). W of Forcados town, near confluence<br />
of Odimodi Creek and Forcados River, 05°22'N, 05°26'E,<br />
28 Feb 1976, C. B. Powell, 14 specimens (L). Niger delta,<br />
between Brass and Port Harcourt, May-Aug 1960, H. J. G.<br />
Beets, 16* (L).<br />
Gabon: No specific locality, Duparquet, syntype of Panopeus<br />
afncanus, 1 dry c* (W, USNM 20263).<br />
Congo: Pointe-Noire, intertidal, Mar 1965, A. Crosnier,<br />
26*, 4$ (1 ov) (W).<br />
Zaire: Banana, mouth of Congo River, American Museum<br />
Congo Expedition 1909-1915, Jul-Aug 1915, H. Lang,<br />
46*, 5$ (1 ov) (W). Data same, Jul 1915, 56*, 39 (1 ov) (W).<br />
Angola: Luanda, W coast; American Museum Congo<br />
Expedition 1909-1915, 21 Sep 1915, H. Lang, 1$, 2juv (W).<br />
Between Luanda and Cuanza, 23 km from Luanda, 20 Jun<br />
1967, G. Hartmann, 1$ (L). Lobito, 1899, P. Kamerman, 16*<br />
(L).<br />
DESCRIPTION.—Capart, 1951:148.<br />
Figures: Capart, 1951, fig. 54, pi. 3: fig. 3;<br />
Monod, 1956, figs. 406-415.<br />
MalePleopod:Capart, 1951, pi. 3: fig. 3 (Congo);<br />
Monod, 1956, figs. 410-414 (Senegal, Sierra<br />
Leone, Ivory Coast).<br />
Color: "Coloration gris fonce a brun; extremite<br />
des pinces noir et blanc" (Capart, 1951:148).<br />
Rossignol (1957:83) gave a more detailed color<br />
description: "marron plus ou moins foncee.<br />
Chelipedes: face externe et bord superieur de la<br />
main, de raeme que le pouce et le doigt, tete de<br />
negre (bouts de doigts jaunatres). Face inferieure<br />
et bord inferieure de la main jaunatres."<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths ranging from 6 to 43 mm; the carapace<br />
widths of ovigerous females range from 17<br />
to 30.5 mm.<br />
REMARKS.—We suspect that Monod's Panopeus<br />
sp., a damaged male, 12 X 16 mm, from the<br />
Ivindo River, near Ogooue, a locality in a river<br />
some distance from the sea in Gabon, actually<br />
can be identified with this species. One of the<br />
characters used by Monod to distinguish this<br />
specimen is the laterally directed fifth anterolateral<br />
tooth on the carapace, which is also somewhat<br />
upturned; it is curved upward in some of<br />
our specimens. Rathbun (1921:439) noted that<br />
"there is considerable variation in the shape of<br />
the lateral teeth of the carapace ... ," and she<br />
figured one specimen (1921, fig. 19a) in which<br />
the posterior teeth are directed almost laterally.<br />
The front of Monod's specimen is damaged. The<br />
deep sinus between the first and second tooth<br />
shown by Monod is also figured by Rathbun for<br />
three different specimens.<br />
The male pleopod of Monod's specimen, which<br />
seems to terminate in a club-shaped tip rather<br />
than in the trilobed apex typical of Panopeus, may<br />
be abnormal or damaged; the specimen is quite<br />
small, so it may just be undeveloped. A survey of<br />
the pleopods of species of Panopeus in the <strong>Smithsonian</strong><br />
<strong>Institution</strong>, in which males of all species<br />
but P. convexus A. Milne Edwards, 1880, are represented,<br />
reveals that all of the other species in<br />
the genus have the typically trilobed male<br />
pleopod.<br />
Monod (1956:329) commented on the absence<br />
of the red spot on the inner surface of the ischium<br />
of the third maxilliped in the specimen from<br />
Ivindo. That spot, characteristic of P. ajricanus, is<br />
not always present in species in which it is known<br />
to occur (A. B. Williams, pers. comm.).<br />
The occurrence of Monod's specimen from Gabon<br />
in a river is consistent with the known habitat<br />
preference of P. africanus for lagoons and estuaries.<br />
Monod (1956:329) noted the occurrence of a<br />
red spot on the inner surface of the ischium of the<br />
third maxilliped in this species, as well as in<br />
Eurypanopeus blanchardi (p. 130). That spot also is<br />
characteristic of the western Atlantic P. herbstii H.<br />
Milne Edwards, 1834. In that species, Williams<br />
(1965:197) noted that in a sample of almost 600<br />
specimens from Beaufort, North Carolina, that<br />
spot was present in 100% of the males but only in<br />
55% of the females. The significance of the spot<br />
is unknown.
148<br />
BIOLOGY.—Panopeus africanus is a characteristic<br />
inhabitant of estuaries and lagoons along the<br />
<strong>West</strong> <strong>African</strong> coast; it may also occur littorally<br />
and sublittorally. Longhurst (1958) found it in<br />
Sierra Leone in estuaries as well as offshore to a<br />
depth of 140 m. It is rarely found far from the<br />
intertidal zone.<br />
Longhurst (1957) found this species in the<br />
stomach contents of the following fishes in the<br />
Sierra Leone River: Arius latiscutatus Giinther,<br />
Pomadasys jubelini (Cuvier), Diagramma macrolepis<br />
(Boulenger), Galeoides decadactylus (Bloch), and Cynoglossus<br />
senegalensis (Kaup).<br />
Apparently this species spawns all year. Ovigerous<br />
females have been collected in January,<br />
March, April, July, and November (Capart,<br />
1951; Monod, 1956; Rossignol, 1957; p. 147<br />
herein).<br />
DISTRIBUTION.—Eastern Atlantic, from S Portugal<br />
and SW Spain southward to Angola, including<br />
the offshore islands of Fernando Poo,<br />
Principe, and Sao Tome; possibly introduced to<br />
Durban Bay, South Africa (Barnard, 1954, 1955).<br />
Shallow water, usually in estuaries and lagoons.<br />
Monod (1956) summarized earlier records and<br />
reported many specimens from localities between<br />
Mauritania and Angola. Since 1956 it has been<br />
reported from the following localities.<br />
Morocco: Oued Bou Regreg (Forest and Gantes, 1960).<br />
Guinea: Conakry, 0-2.5 m; lie Roume, lies de Los, shore<br />
(Uschakov, 1970).<br />
Sierra Leone: Sierra Leone River (Longhurst, 1957). Off<br />
Sierra Leone, 0-140 m (Longhurst, 1958).<br />
Ghana: Ada; off Accra, 8 m (Gauld, 1960).<br />
Nigeria: Elechi Creek, Port Harcourt, 04°47'15"N,<br />
06°48'45"E (Powell, 1979).<br />
Principe: No specific locality (Frade, 1950; Forest and<br />
Guinot, 1966). Rio Papagaio, shore (Forest and Guinot,<br />
1966).<br />
Sao Tome: No specific locality (Frade, 1950).<br />
Congo: Lagoon of Loango (Rossignol, 1957). Wof Pointe-<br />
Noirc (Rossignol, 1962).<br />
Angola: Mangais, Lobito (Bott, 1964). Baia de Luanda,<br />
shore; Baia do Lobito, shore; Praia da Rocha, near Benguela,<br />
shore (Guinot and Ribeiro, 1962). Between Cacuaco and<br />
Lobito-Benguela (Hartmann-Schroder and Hartmann,<br />
1974).<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Genus Paractaea Guinot, 1969<br />
ParactaeaGuinot, 1969d:241 [type-species: Xantho rufopunctatus<br />
H. Milne Edwards, 1834, by original designation; gender:<br />
feminine]; 1971:1071 [list of species: Actaea margaritaria A.<br />
Milne Edwards, 1867, not included]; 1976:249 (revision).<br />
REMARKS.—The eastern Atlantic Actaea margaritaria<br />
A. Milne Edwards, 1867, was listed under<br />
neither Actaea (p. 1070) nor Paractaea (p. 1071) by<br />
Guinot (1971), who noted (1969d:244):<br />
Enfin, il convient de dire un mot de quelques autres<br />
especes susceptibles d'entrer, apres revision, dans le genre<br />
Paractaea gen. nov. En effet, un certain nombre d'especes<br />
jusqu'a present rattachees a Actaea et qui ne peuvent etre<br />
conservees dans ce genre tel que nous l'avons delimite, offrent<br />
les memes caracteres essentiels que les Paractaea. II s'agit de<br />
Actaea margaritaria A. Milne Edwards, 1867, espece est-atlantique.<br />
In 1976 Guinot assigned margaritaria to Paractaea.<br />
* Paractaea margaritaria<br />
(A. Milne Edwards, 1867)<br />
Actaea margaritaria.—Capart, 1951:159.—Gauld, 1960:70.—<br />
Serene, 1961:197 [listed].—Rossignol, 1962:117.—Chace,<br />
1966:637.—Forest and Guinot, 1966:77.—Guinot, 1969d:<br />
224, 251 [discussion].<br />
Actaea (Actaea) margaritaria.—Monod, 1956:294, figs. 357-<br />
360.—Guinot and Ribeiro, 1962:56.—Ribeiro, 1964:9.<br />
Paractaea margaritaria.—Guinot, 1976:251.<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 70, 33 m, branched Foraminifera, 1$ (L).<br />
Annobon: Sta 275, 9-69 m, rubble of coralline algae,<br />
26* (W). Sta 282, 18-37 m, nodular coralline algae, 1$, 3 juv<br />
(W). Sta 283, 51-55 m, nodular coralline algae, 3(5, 2? (L).<br />
DESCRIPTION.—Capart, 1951:159.<br />
Figures: Monod, 1956, figs. 357-360; Guinot,<br />
1976, pi. 16: fig. 6.<br />
Male Pleopod: Monod, 1956, figs. 358-360<br />
(Ghana).<br />
Color: "Rouge intense, une tache jaune en arriere<br />
des yeux" (Capart, 1951:159).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 4 to 9 mm.
NUMBER 306 149<br />
BIOLOGY.—Paractaea margaritaria is a sublittoral<br />
species, occurring on rough bottom in depths<br />
between 4-5 and 91 m. It has habitat preferences<br />
similar to those of P. rufopunctata africana; the two<br />
species were taken together at three stations by<br />
the Calypso (Forest and Guinot, 1966) and at two<br />
stations by the Pillsbury. Our material was collected<br />
on bottom with branched Foraminifera or<br />
the characteristic coralline algae found off Annobon.<br />
The Calypso specimens were taken in calcareous<br />
algae at four stations and on rocks and<br />
coral at two stations. Guinot (1976) reported one<br />
specimen from sand and gorgonians off Togo.<br />
Off <strong>West</strong> Africa, ovigerous females have been<br />
collected in June and November (Guinot and<br />
Ribeiro, 1962; Ribeiro, 1964; Forest and Guinot,<br />
1966). Chace (1966) recorded an ovigerous female<br />
from Saint Helena in April.<br />
DISTRIBUTION.—<strong>West</strong> coast of Africa and Saint<br />
Helena (Chace, 1966). Off <strong>West</strong> Africa it is<br />
known from the Cape Verde Islands, Ghana,<br />
Gabon, and the offshore islands of the Gulf of<br />
Guinea: Annobon, Sao Tome, and Principe; it<br />
has not previously been recorded from the <strong>African</strong><br />
mainland as far north as Liberia. It is a sublittoral<br />
species, occurring in depths between 4-5 and<br />
91 m. Monod (1956) reported material from<br />
Ghana and Annobon; since 1956 it has been<br />
recorded from the following:<br />
Cape Verde Islands: No specific locality (Guinot, 1976).<br />
Porto da Praia, Sao Tiago (as La Praya) (Guinot, 1976). Sao<br />
Vicente (as Cap St. Vincent) (Guinot, 1976). Baia do Porto<br />
Grande, Sao Vicente, 3.5-11 m (Guinot and Ribeiro, 1962;<br />
Ribeiro, 1964).<br />
Ghana: Off Accra, 15 m (Gauld, 1960).<br />
Togo: No specific locality (Guinot, 1976).<br />
Principe: 01°43'N, 07°28'55"E, 37 m (Forest and Guinot,<br />
1966).<br />
Sao Tome: 00°20'N, 06°46'E, 10 m; Ponta Diogo Vaz,<br />
W coast, 30 m; off Diogo Nunes, 4-5 m (all Forest and<br />
Guinot, 1966).<br />
Annobon: No specific locality, 12 m (Guinot, 1976).<br />
01°27.5'S, O5°36.5'E, 35 m, and Isla Tortuga, NW coast,<br />
15-40 m (Forest and Guinot, 1966).<br />
Gabon: No specific locality, in 50 m (Guinot, 1976). Off<br />
Libreville, 45-57 m (Rossignol, 1962).<br />
Congo: Baie de Pointe-Noire, 13 m (Guinot, 1976).<br />
Paractaea monodi Guinot, 1969<br />
Actaca (Actaea) rufopunctota.—Monod, 1956:293 [part] [not<br />
Actaea rufopunctata H. Milne Edwards, 1834].<br />
Actaea rufopunctata.—Guinot, 1969d:250 [discussion] [not Actaea<br />
rufopunctata H. Milne Edwards, 1834].<br />
Paractaea monodi Guinot, 1969d:259, fig. 33; 1971:1072<br />
[listed]; 1976:250 [listed], fig. 7.—Turkay, 1976b:61<br />
[listed], 67.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Azores: No other data, paratypes, Id, 19<br />
(W).<br />
Madeira: S coast near Ponta da Oliveira, 32°39'N, 16°-<br />
49'W, 0-20 m, diving, 15 Mar 1976, Onversaagd Sta 82, 1
150<br />
Azores, Madeira, Ilhas Desertas, the Canary Islands,<br />
the Cape Verde Islands, and the Mediterranean;<br />
for comments on material from Mediterranean<br />
localities see Guinot (1969d:260, 261).<br />
Guinot (1969d) summarized the literature. We<br />
have found only the following records published<br />
since then:<br />
Madeira: Funchal harbor; between Ponta da Garajau<br />
and Ponta da Oliveira, 5-6 m (Tiirkay, 1976b).<br />
Canary Islands: Arrecife, E of Lanzarote (Guinot, 1976).<br />
Egypt: W of Abou Kir; Abou Kir bay region: Rosetta to<br />
Port Said (Ramadan and Dowidar, 1976).<br />
*Paractaea rufopunctata africana Guinot, 1976<br />
Actaea rufopwclata.—Bouvier, 1906:4%.—Balss, 1914:102.—<br />
Crosnier, 1964:31.—Forest and Guinot, 1966:76.<br />
Actaea (Actaea) rufopunctata.—Monod, 1956:293 [part].<br />
Paractaea rufopunctata forme africana Guinot, 1969d:251, fig.<br />
26; 1971:1071 [listed].<br />
Paractaea rufopunctata africana Guinot, 1976:250, pi. 16: fig. 5.<br />
MATERIAL EXAMINED.—Pillsbury Material: Annobon:<br />
Sta 275, 9-69 m, rubble of coralline algae, 36, 1$ (L, W).<br />
Sta 282, 18-37 m, nodular coralline algae, Ic3, 1? (L).<br />
DESCRIPTION.—Guinot, 1969d:246 (P. rufopunctata<br />
rufopunctata), 251 (P. rufopunctata forme africana).<br />
Figures: Guinot, 1969d, fig. 26; 1976, pi. 16:<br />
fig. 5.<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 13 to 18 mm.<br />
REMARKS.—In her revision of Actaea and Paractaea,<br />
Guinot (1969d) recognized Paractaea rufopunctata<br />
rufopunctata as an Indo-<strong>West</strong> Pacific taxon<br />
and introduced the infrasubspecific name, africana,<br />
with no standing in nomenclature, for the<br />
tropical eastern Atlantic population; she later<br />
validated the name in 1976. In her discussion of<br />
the problem she noted (1969d:251) that "il y aura<br />
lieu de revenir sur le statut des representants<br />
ouest-africains de rufopunctata, en d'autres termes<br />
de decider si ceux-ci meritent bien de constituer<br />
une forme distincte de la rufopunctata typique."<br />
Unfortunately, the Pillsbury material is not adequate<br />
for us to elucidate the problem.<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
BIOLOGY.—Off <strong>West</strong> Africa this species occurs<br />
from the intertidal zone to a depth of at least<br />
45 m; there is one Pillsbury record of 9-69 m. It<br />
was taken at 10 stations by the Calypso (Forest<br />
and Guinot, 1966): three were on or near (0-6 m)<br />
shore, two were from depths of 6-8 m, and five<br />
were from depths ranging from 15 (15-40 m) to<br />
45 m. The species apparently prefers rough bottom;<br />
the coralline algae habitat common around<br />
the offshore islands of the Gulf of Guinea has<br />
yielded several specimens from both the Calypso<br />
and the Pillsbury collections. This species also was<br />
found in the following habitats by the Calypso<br />
(Forest and Guinot, 1966): sand and rocks; rocks<br />
and coral; rocks, calcareous algae, and coral.<br />
Crosnier (1964) found the species in sponges, coral<br />
and marl on rocky bottom with gorgonians off<br />
Cameroon.<br />
Off <strong>West</strong> Africa, ovigerous females have been<br />
recorded in June (Forest and Guinot, 1966).<br />
DISTRIBUTION.—<strong>West</strong> Africa. Guinot (1969d)<br />
recognized a forme africana for the population<br />
occurring off tropical <strong>West</strong> Africa, from the offshore<br />
islands of the Gulf of Guinea: Sao Tome,<br />
Annobon, and Principe, which she elevated to<br />
subspecific status in 1976. She erred, however, in<br />
considering the <strong>West</strong> <strong>African</strong> population to be<br />
exclusively insular (1969d:251); Crosnier (1964)<br />
has reported the species from the continental shelf<br />
off Cameroon. Guinot (1969d:259, fig. 33) assigned<br />
material from the Cape Verde Islands to<br />
Paractaea monodi Guinot (p. 149). <strong>West</strong> <strong>African</strong><br />
records for P. rufopunctata africana are as follows:<br />
Cameroon: No specific locality (Crosnier, 1964).<br />
Principe: 01°35'N, 07°28'E, 45 m; 01°43'N, O7°28'55"E,<br />
37 m; and Ponta da Mina, beach (Forest and Guinot, 1966).<br />
Sao Tome: No specific locality (Bouvier, 1906; Guinot,<br />
1969d). Ponta Diogo Vaz, W coast, 0-6 and 30 m; in front<br />
of Sao Tome, 8 m; W of Ponta Diogo Nunes, shore; in front<br />
of Ponta Oquedelrei, 6 m (all Forest and Guinot, 1966).<br />
Annobon: No specific locality (Balss, 1914). 01°25'12"S,<br />
05°36'05"E, 20 m, and NW side, Isla Tortuga, 15-40 m<br />
(Forest and Guinot, 1966; Guinot, 1976).<br />
The identity of the material from Ascension<br />
Island assigned to this species by Benedict (1893)
NUMBER 306 151<br />
and Rathbun (1930) remains to be determined.<br />
That material will be studied in a review of the<br />
decapods of Ascension now underway by Manning<br />
in collaboration with Fenner A. Chace, Jr.<br />
Genus Paraxanthias Odhner, 1925<br />
Paraxanthias Odhner, 1925:85 [type-species: Xanthodes notatus<br />
Dana, 1852, by original designation; gender: masculine].—Guinot,<br />
1971:1069 [list of species].<br />
Paraxanthias eriphioides<br />
(A. Milne Edwards, 1867)<br />
Paraxanthias eriphioides.—Monod, 1956:304, figs. 371-375<br />
[Cape Verde Islands; references].—Guinot and Ribeiro,<br />
1962:57 [Cape Verde Islands].—Ribeiro, 1964:10 [Cape<br />
Verde Islands].—Guinot, 1968a:718 [discussion]; 1971:<br />
1069 [listed].<br />
Paraxanthias Periphioides.—Guinot, 1968a: 720, figs. 48, 53<br />
[Cape Verde Islands].<br />
DISTRIBUTION.—Eastern Atlantic, from the<br />
Azores and the Cape Verde Islands; sublittoral,<br />
10-85 m.<br />
Genus Pilumnopeus A. Milne Edwards, 1863<br />
Pilumnopeus A. Milne Edwards, 1863:289 [a genus without<br />
included nominal species; type-species: Pilumnopeus crassimanus<br />
A. Milne Edwards, 1867, a subjective junior synonym<br />
of Ozius serratifrons Kinahan, 1858, by subsequent<br />
designation by Balss, 1933b: 33, 34; gender: masculine;<br />
name 1643 on Official List, there dated 1867 in error].—<br />
Takeda and Miyake, 1969:119 [definition; list of Indo-<br />
<strong>West</strong> Pacific species].<br />
* Pilumnopeus africanus (De Man, 1902)<br />
Heteropanope (Pilumnopeus) ajricana.—Monod, 1956:270, figs.<br />
319-325.<br />
Pilumnopeus africanus.—Uschakov, 1970:455 [listed].<br />
MATERIAL EXAMINED.—Pillsbury Material: Nigeria:<br />
Sta 224, Lagos, shore, sand beach, 1$ (W).<br />
Other Material: Liberia: Free Port area, Monrovia, oyster<br />
cultch, 13 Mar 1953, G. C. Miller, 1
152<br />
MATERIAL EXAMINED.—Ptllsbury Material: None.<br />
Other Material: Nigeria, between Brass and Port Harcourt,<br />
Niger delta, May-Aug 1960, H. J. G. Beets, 16*, 19 (L).<br />
DESCRIPTION.—Capart, 1951:150.<br />
Figures: Capart, 1951, fig. 55, pi. 3: fig. 1;<br />
Monod, 1956, figs. 314-318.<br />
Male Pleopod: Capart, 1951, pi. 3: fig. 1 (Congo);<br />
Monod, 1956, figs. 316-318 (Ghana).<br />
Color: According to Capart (1951:150) this species<br />
is "brun-rouge, les pinces plus claires, les<br />
doigts noirs a leur extremite."<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 4 to 11 mm.<br />
BIOLOGY.—Pilumnopeus caparti is an inshore, estuarine<br />
species; most of the material reported in<br />
the literature was collected at or near the mouths<br />
of rivers. Capart's (1951) material was taken in<br />
6-8 m on brown, black mud, and one of Monod's<br />
specimens was taken from a beacon at a river<br />
mouth. Longhurst (1957) found this species in the<br />
stomach contents of Arius latiscutatus Giinther in<br />
the Sierra Leone River and in 1958 characterized<br />
P. caparti as an estuarine, shallow shelf species; it<br />
was found on rocks, muddy sand, shelly mud,<br />
and sand. It also was reported by Longhurst in<br />
the latter paper from one shelf station in 72 m;<br />
that record appears to be questionable.<br />
Ovigerous females have been collected in August<br />
(Capart, 1951).<br />
DISTRIBUTION.—<strong>West</strong> Africa, from scattered localities<br />
between Sierra Leone and the Congo,<br />
usually in estuaries. Records in the literature<br />
include the following:<br />
Sierra Leone: No specific locality, in estuaries (Longhurst,<br />
1958). Freetown (Monod, 1956). Sierra Leone River<br />
(Longhurst, 1957).<br />
Ghana: U.A.C. beach, Ada, River Volta (type-locality)<br />
(Monod, 1956; Gauld, 1960).<br />
Cameroon: Mouth of Wouri (River) (Monod, 1956).<br />
Congo: Mouth of Songololo River, Pointe-Noire (Rossignol,<br />
1962).<br />
Zaire: Crique de Banana, 06°01'S, 12°23'3O"E, 6-8 m<br />
(Capart, 1951).<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
on Official List].—Takeda and Miyake, 1968:2 [list of<br />
Indo-<strong>West</strong> Pacific species].<br />
Pilumnus hirtellus (Linnaeus, 1761)<br />
Cancer hirtellus Linnaeus, 1761:493.<br />
Pilumnus hirtellus.—Forest and Gantes, 1960:352 [Morocco].—Maurin,<br />
1968a: 107 [Mediterranean].—Zariquiey<br />
Alvarez, 1968:392, figs. 2g, 128c,d, 129f [Spain;<br />
references].—Christiansen, 1969:75, fig. 30, map 24 [Scandinavia].—Turkay,<br />
1976b:61 [listed], 69 [Madeira].<br />
SYNONYM.—Pilumnus hirtellus ponticus Czerniavsky,<br />
1868.<br />
DISTRIBUTION.—Eastern Atlantic, from Norway<br />
and British Isles southward to NW Morocco,<br />
Madeira, the Cape Verde Islands, Mediterranean;<br />
sublittoral, to a depth of 15-20 m.<br />
Pilumnus inermis A. Milne Edwards and<br />
Bouvier, 1894<br />
Pilumnus inermis.—Monod, 1956:247, figs. 291-295.—Longhurst,<br />
1958:88.—Gauld, 1960:70.—Rossignol, 1962:117.<br />
—Crosnier, 1964:31.—Forest and Guinot, 1966:70.—Zariquiey<br />
Alvarez, 1968:391 [Portugal; references].—Uschakov,<br />
1970:455 [listed].—Turkay, 1976a:25 [listed], 38,<br />
fig. 26 [Portugal in part]; 1976b:61 [listed], 69.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Azores: Ilha do Faial, Talisman, syntype of<br />
Pilumnus hirtellus var. inermis, 1 damaged $ (W, USNM<br />
22951).<br />
Madeira: S of Madeira, 32°39'N, 16°49'W, 125-150 m,<br />
shells and shell agglomerates, triangular dredge, 16 Mar<br />
1976, Onversaagd Sta 94, 16*, 2$, 1 juv (L).<br />
Senegal: Around Dakar, 28 Jan 1941, Th. Monod, 16*<br />
(W).<br />
DESCRIPTION.—A. Milne Edwards and Bouvier,<br />
1894:40.<br />
Figures: Monod, 1956, figs. 291-295.<br />
Male Pleopod: Monod, 1956, figs. 293-295 (Senegal);<br />
Turkay, 1976a, fig. 26 (Morocco).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 4.5 to 13 mm. The damaged<br />
female syntype has a carapace width of 11.6 mm.<br />
Genus Pilumnus Leach, 1815<br />
BIOLOGY.—Pilumnus inermis is generally a sub-<br />
Pilumnus Leach, 1815a:321 [type-species: Cancer hirtellus Linlittoral species; Monod (1956:249) recorded one<br />
naeus, 1761, by monotypy; gender: masculine; name 348 specimen taken at a beach, but the majority of
NUMBER 306 153<br />
the records in the literature are from depths below<br />
4-5 m. Of the eight <strong>West</strong> <strong>African</strong> records in<br />
Monod (1956) for which depth is given, seven are<br />
from depths between 15 and 30 m. Indeed, literature<br />
records for this species suggest that it lives<br />
in shallower water in the Gulf of Guinea than it<br />
does in the northern part of its range. Nunes-<br />
Ruivo (1961) found the species in 350 m off<br />
Portugal, and Tiirkay (1976a) reported it in<br />
depths between 120 and 375 m off Morocco and<br />
(1976b) recorded it from ca 150 m, 220 m, and<br />
110-440 m off Madeira. Forest and Guinot (1966:<br />
71) pointed out that of 14 stations at which this<br />
species was taken by the Calypso in the Gulf of<br />
Guinea, 13 were in depths between 4-5 and<br />
21 m.<br />
This species, like most Pilumnus, prefers a rough<br />
bottom. Longhurst (1958) reported it from shelly<br />
sand off Sierra Leone, and Crosnier (1964) took<br />
it in sponges, coral and marl on rocky bottom<br />
with gorgonians off Cameroon. Rossignol (1962)<br />
reported it from muddy sand with Antedon off<br />
Pointe-Noire. The Calypso collected it from a variety<br />
of bottoms off Sao Tome and Principe islands<br />
in the Gulf of Guinea, usually on coralline<br />
algae or a mixture of coralline algae and other<br />
substrates (Forest and Guinot, 1966).<br />
Off <strong>West</strong> Africa, ovigerous females have been<br />
collected in February, March, April, June, and<br />
July (Monod, 1956; Forest and Guinot, 1966).<br />
Nunes-Ruivo (1961) reported ovigerous females<br />
taken off Portugal in August.<br />
DISTRIBUTION.—Eastern Atlantic, from Portugal,<br />
the Azores, Madeira, the Cape Verde Islands,<br />
and the <strong>African</strong> coast from Morocco and Cabo<br />
Bojador, Spanish Sahara, southward to the<br />
Congo and possibly Gabon (that record based on<br />
material collected by Heurtel, whose localities are<br />
unreliable, as pointed out by Monod, 1956), including<br />
the offshore islands of the Gulf of Guinea,<br />
Principe and Sao Tome; sublittoral, in 4-5 m to<br />
400 m. Monod (1956) reported material from the<br />
Azores, Madeira, Spanish Sahara, Senegal,<br />
Guinea, Guinea-Bissau, Sierra Leone, Ghana,<br />
Principe, Sao Tome, and Gabon; since then the<br />
species has been recorded from the following:<br />
Madeira: No specific locality; Funchal harbor, ca. 150 m,<br />
220 m, and 110-440 m (Tiirkay, 1976b).<br />
Morocco: 33°19'N, 09°00'W, 120-180 m; 33°05.5'N,<br />
09°18'W, 160-250 m; 31°35'N, 10°05'W, 150-160 m; 31°-<br />
01'N, 10°16'W, 360-375 m (all Tiirkay, 1976a).<br />
Guinea: No specific locality (Uschakov, 1970).<br />
Sierra Leone: No specific locality, in 44-76 m (Longhurst,<br />
1958).<br />
Ghana: Off Accra, 15 m; Prampram; Dixcove (all Gauld,<br />
1960).<br />
Cameroon: No specific locality (Crosnier, 1964).<br />
Principe: 01°42'30"N, 07°28'E, 21 m; 01°36'50"N, 07°-<br />
22'10"E, 19 m; 01°43'10"N, 07°28'20"E, 73 m; between<br />
Ponta da Mina and Ilheu Santana, 8-10 m and 10-12 m; in<br />
front of [Cais de] Santana, 11 m; in front of Praia Pequena,<br />
5-6 m (all Forest and Guinot, 1966).<br />
Sao Tome: Off Sao Tome, 5 m; 00°20'N, 06°46'E, 10 m;<br />
00°25'15"N, 06°43'05"E, 8-30 m; Baia de Ana de Chaves,<br />
5 m; in front of Ponta Oquedelrei, 6 m; in front of Praia<br />
Lagarto, 5-6 m; off Ponta Diogo Nunes, 4-5 m (all Forest<br />
and Guinot, 1966).<br />
Congo: Pointe-Noire, 8-9 m (Rossignol, 1962).<br />
* Pilumnus perrieri A. Milne Edwards and<br />
Bouvier, 1898<br />
Pilumnus perrieri.—Capart, 1951:143 [discussion].—Monod,<br />
1956:244, figs. 288-290.—Rossignol, 1962:117.—Forest<br />
and Guinot, 1966:70.—Uschakov, 1970:455 [listed].<br />
MATERIAL EXAMINED.—Pillsbury Material: Ghana:<br />
Sta 23, 42 m, foliate brown to orange bryozoans, 1? (L). Sta<br />
27, 33 m, 1(3, 1$ ov (W).<br />
Nigeria: Sta 248, 33 m, 1
154<br />
taken on mud; mud with Area; mud, calcareous<br />
algae, and shell; rocks; and on calcareous algae<br />
(five stations). Rossignol (1962:117) noted that<br />
the species is "assez commun dans le coralligene<br />
et les fonds rocheux."<br />
Ovigerous females have been collected in<br />
March and May (Monod, 1956; Pillsbury).<br />
DISTRIBUTION.—<strong>West</strong> Africa, from scattered localities<br />
between the Cape Verde Islands and Senegal<br />
to Gabon; sublittoral, in depths between 20<br />
and 91 m. Monod (1956) summarized the earlier<br />
literature and reported material from Senegal<br />
and Guinea; since 1956 the species has been<br />
reported from the following:<br />
Guinea: No specific locality (Uschakov, 1970).<br />
Nigeria: Off the mouths of the Niger River, O4°O3'N,<br />
06°12'E, 32 m (Forest and Guinot, 1966).<br />
Principe: Between Ilheu Caroco and Ponta do Pico Negro,<br />
40 m (Rossignol, 1962). 01°38'25"N, 07°22'05"E, 31 m;<br />
01°43'10"N, 07°28'20"E, 73 m; 01°43'N, 07°28'55"E, 37 m;<br />
in front of Baia de Santo Antonio, 50 m (Forest and Guinot,<br />
1966).<br />
Annobon: O1°27.5'S, 05°36.5'E, 35 m; N of San Antonio,<br />
23 m (Forest and Guinot, 1966).<br />
Rio Muni: W Corisco Bay, 40 m (Rossignol, 1962).<br />
Gabon: W of Libreville, 45-57 m; W of Nyanga, 03°S,<br />
65-70 m (Rossignol, 1962).<br />
Pilumnus spinifer H. Milne Edwards, 1834<br />
?Pilumnus hirtellus.—Capart, 1951:140, fig. 50 [Spanish Sahara]<br />
[not Pilumnus hirtellus Linnaeus, 1761)].<br />
Pilumnus spinifer.—Monod, 1956:251, figs. 296, 297 [Azores,<br />
Mauritania; references].—Zariquiey Alvarez, 1968:391,<br />
fig. 129a-e [Spain; references].—Christiansen, 1969:77,<br />
fig. 31, map 25 [Scandinavia].—Tiirkay, 1976b:61 [listed],<br />
69 [Madeira].<br />
REMARKS.—There is a dry syntype of this species<br />
(USNM 20262), a male, carapace width<br />
30.8 mm, from the Mediterranean, in the collections<br />
of the <strong>Smithsonian</strong> <strong>Institution</strong>.<br />
DISTRIBUTION.—Eastern Atlantic, from Sweden,<br />
Portugal, Azores, Mediterranean, NW coast<br />
of Africa to Mauritania; sublittoral, to about<br />
100 m.<br />
* Pilumnus stebbingi Capart, 1951<br />
Pilumnus stebbingi Capart, 1951:144, fig. 52, pi. 3: fig. 6.—<br />
Monod, 1956:241, 632, figs. 279-287.—Longhurst, 1958:<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
88.—Rossignol, 1962:116.—Guinot and Ribeiro, 1962:<br />
52.—Forest and Guinot, 1966:69.<br />
MATERIAL EXAMINED.—Pillsbury Material: Ghana:<br />
Sta 24, 35-37 m, dark red bryozoans, \S (L).<br />
Annobon: Sta 283, 51-55 m, nodular coralline algae, 26cJ,<br />
399 (7 ov), 15 juv (L, W). Sta 284, 73 m, black basaltic rocks,<br />
2$, 2juv(W).<br />
DESCRIPTION.—Capart, 1951:144.<br />
Figures: Capart, 1951, fig. 52, pi. 3: fig. 6;<br />
Monod, 1956, figs. 279-287.<br />
Male Pleopod: Capart, 1951, pi. 3: fig. 6 (Angola);<br />
Monod, 1956, figs. 284-287 (Senegal,<br />
Guinea).<br />
Color: Monod (1956:241) made the following<br />
observations on specimens from Guinea: "Epines<br />
rouges sur les pattes, qui sont annelees de rouge."<br />
Rossignol (1962:116) added: "Caracteristique par<br />
son tomentum epais, feutre, de couleur grise a<br />
gris-noir avec, sur les pinces, des tubercules epineux<br />
rouges."<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 3 to 13 mm; the carapace widths<br />
of ovigerous females is 10 to 12 mm.<br />
REMARKS.—The ornamentation of the chelae<br />
in our specimens is very variable: some have only<br />
pink tubercles, others pink and white tubercles;<br />
in some specimens the entire outer surface of the<br />
chela is covered with tubercles, in others the<br />
tubercles are restricted to the upper half of the<br />
surface.<br />
BIOLOGY.—Pilumnus stebbingi is a shelf rather<br />
than shore species, inhabiting moderate depths<br />
on relatively rough bottom. All but three of the<br />
20 depth records in the literature are from depths<br />
between 30 and 73 m; the exceptions are 25-<br />
30 m (Capart, 1951), 10-25 m (Longhurst, 1958),<br />
and 23 m (Forest and Guinot, 1966). Capart's<br />
(1951) material was collected on rocks; rocks,<br />
gravel, and coral; and sandy, rocky green mud.<br />
In addition to several records of the species from<br />
the coralline algae off Principe and Annobon, the<br />
same habitat in which the Pillsbury collected a<br />
relatively large series off the latter island, Forest<br />
and Guinot (1966) reported the species from mud,<br />
shells, gorgonians and ascidians; mud and com-
NUMBER 306 155<br />
pacted sand (sable construit); mud, shells, and<br />
Cidaris; and, either mud and shell or sand, rocks<br />
and coral and calcareous algae. Longhurst (1958)<br />
found the species on muddy shell or shelly sand.<br />
Ovigerous females have been collected in May,<br />
June, and September (Monod, 1956; Forest and<br />
Guinot, 1966; Pillsbury).<br />
DISTRIBUTION.—Off <strong>West</strong> Africa, from scattered<br />
localities between Spanish Sahara and Angola,<br />
including the offshore islands of Principe<br />
and Annobon; sublittoral, in depths between 10-<br />
25 m and 73 m. Capart (1951) reported material<br />
from Gabon and Angola, and Monod (1956)<br />
added records from Senegal, Guinea, Sierra<br />
Leone, Annobon, and Principe. Records in the<br />
literature since 1956 include the following:<br />
Spanish Sahara: 21°05'N, 17°14'W, 43-45 m (Forest and<br />
Guinot, 1966).<br />
Senegal: 12°55.5'N, 17°33'W, 65-75 m (Forest and<br />
Guinot, 1966).<br />
Guinea-Bissau: 10°19'N, 16°34'W, 60-73 m (Forest and<br />
Guinot, 1966).<br />
Sierra Leone: No specific locality, in 10-25 m (Longhurst,<br />
1958).<br />
Principe: 01°35'N, 07°28'E, 45 m; 01°38'25"N, 07°22'-<br />
05"E, 31 m; 01°38'35"N, 07°21'35"E, 35 m; 01°43'10"N,<br />
07°28'20"E, 73 m; 01°43'N, 07°28'25"E, 37 m (all Forest<br />
and Guinot, 1966).<br />
Annobon: 01°27.5'S, 05°36.5'E, 35 m; N of San Antonio,<br />
23 m (both Forest and Guinot, 1966).<br />
Gabon: W of Nyanga, 65-70 m (Rossignol, 1962).<br />
Cabinda: W of Cabinda, 55 m (Rossignol, 1962).<br />
Angola: Luanda (Guinot and Ribeiro, 1966).<br />
Genus Platychelonion Crosnier and Guinot,<br />
1969<br />
Platychelonion Crosnier and Guinot, 1969:725 [type-species:<br />
Platychelonion planissimum Crosnier and Guinot, 1969, by<br />
monotypy; gender: neuter.].—Guinot, 1971:1078 [transferred<br />
with a question mark to Geryonidae].<br />
Platychelonion planissimum Crosnier and<br />
Guinot, 1969<br />
Platychelonion planissimum Crosnier and Guinot, 1969:725, figs.<br />
1-9 [Congo].<br />
DISTRIBUTION.—Known only from the typelocality,<br />
Pointe-Noire, Congo, in 10-20 m.<br />
Genus Platypodiella Guinot, 1967<br />
Platypodiella Guinot, 1967d:562 [type-species: Cancerspectabilis<br />
Herbst, 1794, by original designation; gender: feminine];<br />
1971:1074 [list of species].<br />
Platypodiella picta (A. Milne Edwards, 1869)<br />
Cancer geographicus.—Monod, 1933b:548 [footnote, nomen<br />
nudum].<br />
Platypodia picta.—Monod, 1956:299, figs. 363-367 [Senegal,<br />
Sierra Leone, Ghana, Congo; references].—Rossignol,<br />
1957:83, fig. 3 [Congo].—Longhurst, 1958:88 [Sierra<br />
Leone].—Gauld and Buchanan, 1959:128 [Ghana].—<br />
Gauld, 1960:70 [Ghana].—Guinot and Ribeiro, 1962:57<br />
[Cape Verde Islands].—Rossignol, 1962:117 [Congo].—<br />
Ribeiro, 1964:10 [Cape Verde Islands].—Forest and<br />
Guinot, 1966:79 [Sao Tome].—Guinot, 1967d:562 [listed],<br />
563 [discussion; transferred to Platypodiella].<br />
Platypodiella picta.—Guinot, 1971:1074 [listed].<br />
DISTRIBUTION.—<strong>West</strong> Africa, from the Canary<br />
Islands, the Cape Verde Islands, several mainland<br />
localities between Senegal and the Congo, and<br />
the offshore islands of Annobon and Sao Tome<br />
in the Gulf of Guinea; shallow water, intertidal<br />
and sublittoral.<br />
Genus Pseudomedaeus Guinot, 1968<br />
Pseudomedaeus Guinot, 1968a:7l8 [discussion], 726 [type-species:<br />
Medaeus africanus Monod, 1956, by original designation;<br />
gender: masculine]; 1971:1069 [list of species].<br />
* Pseudomedaeus africanus (Monod, 1956)<br />
?Paraxanthios eriphioides.—Capart, 1951:161, fig. 61 [fide<br />
Guinot and Ribeiro, 1962:58] [not Paraxanthias eriphioides<br />
(A. Milne Edwards, 1867)].<br />
Xanthias tuberculidens.—Capart, 1951, pi. 3: fig. 13 [fide<br />
Guinot and Ribeiro, 1962:58] [not Xanthias tuberculidens<br />
Rathbun, 1911].<br />
Medaeus africanus Monod, 1956:306, fig. 380.—Longhurst,<br />
1958:88.—Gauld,1960:70.—Rossignol,1962:118.—Guinot<br />
and Ribeiro, 1962:58, fig. 25.—Crosnier, 1964:34.—Forest<br />
and Guinot, 1966:80.—Crosnier, 1967:328, figs. 5, 11, 12,<br />
15.—Guinot, 1968a:718 [discussion], 726, 727 [listed].<br />
Pseudomedaeus africanus.—Guinot, 1968a:726 [discussion], 727<br />
[listed], fig. 57; 1971:1069 [listed].—Williams, 1978:553,<br />
fig. 4a.
156<br />
MATERIAL EXAMINED.—Pillsbury Material: Ivory<br />
Coast: Sta 64, 68 m, 2c5 (L, W).<br />
Gerommo Material: Gabon: Sta 211, 100 m, Id (W).<br />
Other Material: Dahomey: OfT Dahomey, 55 m, Oct 1963,<br />
A. Crosnier, 2$ (W).<br />
DESCRIPTION.—Monod, 1956:306.<br />
Figures: Capart, 1951, fig. 61, pi. 3: fig. 13;<br />
Monod, 1956, fig. 380; Guinot and Ribeiro, 1962,<br />
fig. 25; Crosnier, 1967, figs. 5, 11, 12, 15.<br />
Male Pleopod: Capart, 1951, pi. 3: fig. 13 (Angola);<br />
Guinot and Ribeiro, 1962, fig. 25 (Spanish<br />
Sahara); Williams, 1978, fig. 4a (Gabon).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 5 to 21 mm; the largest was taken<br />
by the Geronimo.<br />
BIOLOGY.—Pseudomedaeus africanus is an offshore<br />
species, living on the shelf and upper slope in<br />
depths between 34 and 200 m. Crosnier (1964)<br />
characterized it as a cold water species, living in<br />
depths greater than 50 m off Cameroon. Longhurst<br />
(1958) found it on shelly mud in 34 to<br />
100 m off Sierra Leone, and Gauld (1960) reported<br />
it from colonies of Dendrophyllia in 100 m<br />
off Ghana. A large series of more than 70 specimens<br />
was collected by the Calypso on mud shells,<br />
gorgonians and ascidians in 43-45 m off Spanish<br />
Sahara (Forest and Guinot, 1966).<br />
Ovigerous females have been collected in<br />
March and May (Monod, 1956; Forest and<br />
Guinot, 1966).<br />
DISTRIBUTION.—<strong>West</strong> Africa, from scattered localities<br />
between Spanish Sahara and Angola, in<br />
depths between 34 and 200 m. Records since 1956<br />
include the following:<br />
Spanish Sahara: 21°05'N, 17°14'W, 43-45 m (Forest and<br />
Guinot, 1966).<br />
Senegal: Dakar (Guinot, 1968a).<br />
Sierra Leone: No specific locality, in 34-100 m (Longhurst,<br />
1958).<br />
Ghana: OfT Accra, 100 m (Gauld, 1960).<br />
Dahomey: No specific locality, 55 m (Williams, 1978).<br />
Cameroon: No specific locality, deeper than 50 m (Crosnier,<br />
1964).<br />
Gabon: No specific locality (Crosnier, 1967). W of Libreville,<br />
45-57 m, and W of Nyanga, Pointe Panga, 03°10'S<br />
(Rossignol, 1962). 02°32'S, 09°05'E, 101 m (Williams, 1978).<br />
Congo: No specific locality (Crosnier, 1967). WSW of<br />
Pointe-Noire, 120 m (Rossignol, 1962).<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Angola: Ilha de Luanda, 105 m (Guinot and Ribeiro,<br />
1962).<br />
Genus Xantho Leach, 1814<br />
Xantho Leach, 1814:430 [type-species: Cancer incisus Leach,<br />
1814, by monotypy; gender: masculine; name 1016 on<br />
Official List].—Guinot, 1971:1067 [list of species].<br />
Salax Gistel, 1848:xi [substitute name for Xantho Leach, 1814;<br />
type-species: Cancer incisus Leach, 1814; gender; masculine].<br />
Xantho incisus (Leach, 1814)<br />
Xanlho floridus.—Chapman and Santler, 1955:374 [Azores]<br />
[not Cancer Jloridus Linnaeus, 1767].<br />
Xantho (Xantho) incisa.—Monod, 1956:274 [Cape Verde Islands,<br />
Sao Tome(?); references].<br />
Xantho incisus.—Figueira, 1960:9 [Azores].—Forest and<br />
Games, 1960:352 [Morocco].<br />
Xantho incisus incisus.—Zariquiey Alvarez, 1968:398 [Spain;<br />
references].—Guinot, 1968a: 703, figs. 17, 26 [discussion];<br />
1971:1067 [listed].—Tiirkay, 1976b:61 [listed], 68 [Madeira,<br />
Ilhas Desertas].<br />
REMARKS.—There is material of this species<br />
from the Azores and the Canary Islands, but not<br />
from tropical <strong>West</strong> Africa, in the collection of the<br />
<strong>Smithsonian</strong> <strong>Institution</strong>.<br />
In the Mediterranean this species is replaced<br />
by Xantho granulicarpus Forest, 1953 (Zariquiey<br />
Alvarez, 1968:398).<br />
DISTRIBUTION.—Eastern Atlantic, from the<br />
North Sea southward to Morocco, including the<br />
Azores, Madeira, Ilhas Desertas, Canary Islands,<br />
and Cape Verde Islands; possibly also from Sao<br />
Tome and Principe islands in the Gulf of Guinea<br />
(see Monod, 1956:274). It usually lives in shallow<br />
water, littorally to 30-40 m, but also has been<br />
recorded from as deep as about 100 m.<br />
Xantho pilipes A. Milne Edwards, 1867<br />
Xantho (Xantho) pilipes.—Monod, 1956:275, figs. 326-329<br />
[Mauritania, Senegal; references].—Guinot and Ribeiro,<br />
1962:53 [Angola].<br />
Xantho pilipes.—Zariquiey Alvarez, 1968:395, fig. 130b<br />
[Spain; references].—Guinot, 1968a: 704 [discussion].—<br />
Christiansen, 1969:79, fig. 32, map 26 [Scandinavia].—
NUMBER 306 157<br />
Guinot, 1971:1068 [listed]. —Tiirkay, 1975a:71 [listed], 74,<br />
figs. 4, 5 [Spanish Sahara].<br />
DISTRIBUTION.—W coast of Norway, Shetland<br />
Isles, and England southward to Angola, Mediterranean;<br />
shallow water, intertidal to about<br />
40 m.<br />
Xantho sexdentatus (Miers, 1881)<br />
Xantho sexdentatus.—Capart, 1951:157.—Guinot, 1968a:704<br />
[discussion]; 1971:1068 [listed].<br />
Xantho (Xantho) sexdentata.—Monod, 1956:277, figs. 330-334.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Senegal: Les Almadies, Dakar, 0-0.5 m,<br />
under stones, 9 Jun 1964, F. M. Bayer, R. B. Manning, L. B.<br />
Holthuis, 16, 1 juv (L).<br />
DESCRIPTION.—Miers, 1881a:211.<br />
Figures: Monod, 1956, figs. 330-334.<br />
Male Pleopod: Monod, 1956, figs. 332-334 (Senegal).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 6 and 8 mm.<br />
BIOLOGY.—Xantho sexdentatus is a shallow water<br />
species, occurring from the intertidal zone to a<br />
depth of 23 m. Apparently it is a characteristic<br />
inhabitant of rocky shores of Senegal (Sourie,<br />
1954a).<br />
Ovigerous females have been collected in January,<br />
February, July, and October (Monod,<br />
1956).<br />
DISTRIBUTION.—<strong>West</strong> Africa, where it is known<br />
only from the coasts of Spanish Sahara, Mauritania,<br />
and Senegal, from the intertidal zone to a<br />
depth of 23 m. Monod (1956) summarized earlier<br />
records. We have found no references to this<br />
species published since 1956, other than the remarks<br />
of Guinot (1968a) as part of her revision of<br />
the xanthids.<br />
Genus Xanthodius Stimpson, 1859<br />
Xanthodius Stimpson, 1859:52 [type-species: Xanthodius stemberghii<br />
Stimpson, 1859, by monotypy; gender: masculine;<br />
name 379 on Official List].—Guinot, 1971:1068 [list of<br />
species; Xantho denticulata White not included].<br />
* Xanthodius denticulatus (White, 1848)<br />
Xantho (Xantho) denticulata.—Monod, 1956:280, figs. 335-<br />
339.<br />
Xantho denticulata.— Gauld, 1960:70.<br />
Xantho (Xantho) denticulatus.—Forest and Guinot, 1966:74.<br />
Xantho denticulatus.—Guinot, 1968a:700 [discussion].<br />
Xanthodius denticulatus.—Guinot, 1968a:712 [discussion].<br />
[Xantho] denticulatus.—Guinot, 1971:1068 [listed; "Parfois<br />
aussi, range dans Xanthodius"].<br />
MATERIAL EXAMINED.—Pillsbury Material: Fernando<br />
Poo: Sta 258, shore, 1$ (L).<br />
Annobon: Sta 271, shore, sand beach: 1$ (L). Sta 275, 9-<br />
69 m, rubble of coralline algae, 19, 1 juv (W).<br />
DESCRIPTION.—Monod, 1956:281.<br />
Figures: Monod, 1956, figs. 335-339.<br />
Male Pleopod: Monod, 1956, figs. 337-339<br />
(Ghana).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 5 to 11 mm.<br />
REMARKS.—We assign this species, which was<br />
not assigned to any genus in Guinot's preliminary<br />
revision (1968a, 1971), to Xanthodius. Guinot<br />
(1971:1068) noted that it and three other xanthids,<br />
including the <strong>West</strong> <strong>African</strong> Cycloxanthops<br />
occidentalis, were "au voisinage de Macromedaeus<br />
[Ward, 1942], et presentant egalement des affinites<br />
avec Leptodius et peut-etre aussi avec Cycloxanthops,<br />
trois-quatre especes apparentees." Apparently<br />
this problem is under study by Guinot.<br />
BIOLOGY.—Xanthodius denticulatus is a shallow<br />
water species, living from the intertidal zone to a<br />
depth of 21 m or more; it was taken in depths of<br />
11 m or less in 11 of 12 of the stations occupied<br />
by the Calypso (Forest and Guinot, 1966). One of<br />
the collections taken by the Pillsbury was in 9-<br />
69 m, so it may also occur somewhat deeper than<br />
21 m. It appears to be fairly abundant sublittorally<br />
off Sao Tome, where it was taken at 9<br />
stations by the Calypso (Forest and Guinot, 1966).<br />
The Pillsbury specimens were taken from shore<br />
stations and in the offshore bed of coralline algae<br />
off Annobon. The following bottom types were<br />
noted for the Calypso collections: calcareous algae<br />
(5 stations); rocks or sand and rocks (4 stations);<br />
and algae and calcareous algae (1 station).<br />
Off <strong>West</strong> Africa ovigerous females have been
158<br />
collected in February and June (Monod, 1956;<br />
Forest and Guinot, 1966).<br />
DISTRIBUTION.—Atlantic. <strong>West</strong>ern Atlantic<br />
from Bermuda and the Bahamas southward to<br />
Brazil. Eastern Atlantic, from Ghana on the<br />
mainland and the offshore islands of the Gulf of<br />
Guinea: Fernando Poo, Principe, Sao Tome, and<br />
Annobon; shallow water, intertidal to at least<br />
21 m, possibly as deep as 69 m. Monod (1956)<br />
was the first to record the species from <strong>West</strong><br />
Africa; he reported material from Ghana, Sao<br />
Tome, and Annobon. Records in the literature<br />
since 1956 include the following:<br />
Ghana: Prampram and Winneba, intertidal (Gauld,<br />
1960).<br />
Principe: 01°42'30"N, 07°28'E, 21 m; Ponta da Mina,<br />
beach (Forest and Guinot, 1966).<br />
Sao Tome: No specific locality, shore; 00°20'N, 06°46'E,<br />
10 m; off Sao Tome 8 m; in front of Ponta Oquedelrei, 6 m;<br />
Ilheu das Cabras, 0-2 m; in front of the harbormaster's<br />
office, shore; in front of Praia Lagarto, 5-6 m; off Ponta<br />
Diogo Nunes, 4-5 m; in front of Ponta Sao Sebastiao, 11 m<br />
(all Forest and Guinot, 1966).<br />
Annobon: N of San Antonio, 9 m (Forest and Guinot,<br />
1966).<br />
Xanthodius inaequalis faba (Dana, 1852)<br />
Aciaeodes faba Dana, 1852b: 195; atlas, 1855 pi. 11: fig. 1.<br />
Chlorodius (Leptodius) convexus A. Milne Edwards, 1869:410.<br />
Xantho (Leptodius) inaequalis convexa.—Monod, 1956:284 [part<br />
of synonymy], 290, figs. 341a, 349, 350.—Guinot and<br />
Ribeiro, 1962:55.—Ribeiro, 1964:8.<br />
Xanthodius (Leptodius) inaequalis convexus.—Guinot, 1968a: 712<br />
[listed].<br />
Xanthodius inaequalis convexus.—Guinot, 1968a: 714 [discussion],<br />
fig. 31; 1971:1068 [listed].<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Cape Verde Islands, Sao Vicente, 26 Jul<br />
1883, Talisman, \6, 1? (L).<br />
DESCRIPTION.—A. Milne Edwards and Bouvier,<br />
1900:97.<br />
Figures: A. Milne Edwards and Bouvier, 1900,<br />
pi. 17: figs. 1-6.<br />
Male Pleopod: Monod, 1956, figs. 349, 350 (Cape<br />
Verde Islands).<br />
Color: "Coloration vert olivatre, yeux grenat,<br />
pattes ambulatoires lavees de violet" (Monod,<br />
1956:290).<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
REMARKS.—At least two authors, Odhner<br />
(1925:37, footnote, and 80) and Monod (1933b:<br />
511-513), have pointed out that Chlorodius convexus<br />
A. Milne Edwards, 1869, from the Cape Verde<br />
Islands, is conspecific with Actaeodes faba Dana,<br />
1852, from the same locality; Dana's name has<br />
priority and must be used for the taxon from the<br />
Cape Verde Islands.<br />
Monod (1956:284, synonymy) noted that A.<br />
Milne Edwards (1869) included material from<br />
Gabon as well as from the Cape Verde Islands in<br />
his original account of Chlorodius convexus; we can<br />
find only localities from the Cape Verde Islands<br />
in his account (Sao Vicente, Santa Luzia, Maio,<br />
Santiago).<br />
BIOLOGY.—Like the nominate subspecies, X.<br />
inaequalis faba is a littoral species inhabiting rocky<br />
shores. Monod (1956) recorded one specimen<br />
taken in pools with coral, and Guinot and Ribeiro<br />
(1962) and Ribeiro (1964) found it in rocks, on<br />
calcareous algae, and in worm tubes on rocks.<br />
Ovigerous females have been collected in May,<br />
August, September, and October (Monod, 1956;<br />
Guinot and Ribeiro, 1962; Ribeiro, 1964).<br />
DISTRIBUTION.—Known only from the Cape<br />
Verde Islands, where it occurs in the littoral zone.<br />
The nominate subspecies, Xanthodius inaequalis inaequalis<br />
occurs on the <strong>African</strong> mainland and the<br />
offshore islands of the Gulf of Guinea. Monod<br />
(1956) summarized the literature. Records since<br />
his paper were published include the following:<br />
Cape Verde Islands: No specific locality (Guinot, 1968a).<br />
Baia das Gatas, Praia da Matiota, and Baia da Calheta, Sao<br />
Vicente; Baia de Sal Rei, Boavista; Pedra Lume, Sal; and<br />
Baia do Tarrafal and Porto da Praia, Sao Tiago (all Guinot<br />
and Ribeiro, 1962; Ribeiro, 1964).<br />
* Xanthodius inaequalis inaequalis<br />
(Olivier, 1791)<br />
Xantho faba.—Capart, 1951:155, fig. 58, pi. 3: fig. 14 [not<br />
Actaeodes faba Dana, 1852].<br />
Xantho (Leptodius) inaequalis punctata.—Monod, 1956:284 [part<br />
of synonymy], 286, figs. 340, 341b, 342-348.<br />
Xantho inaequalis.—Gauld and Buchanan, 1959:127.—Gauld,<br />
1960:70.—Bassindale, 1961:491, fig. 5.<br />
Xantho (Leptodius) inaequalis inaequalis.—Guinot and Ribeiro,<br />
1962:54.—Forest and Guinot, 1966:75.—Guinot, 1968a:<br />
712 [listed].
NUMBER 306 159<br />
Leptodius angolensis Bott, 1964:30, fig. 3, pi. 1: figs. 4, 5.—<br />
Guinot, 1968a:712 [discussion].<br />
Xanthodius inaequalis inaequalis.—Guinot, 1968a:7l4 [discussion];<br />
1971:1068 [listed].<br />
SYNONYM.—Leptodius punctatus Miers, 1881.<br />
MATERIAL EXAMINED.—Pillsbury Material: Fernando<br />
Poo: Sta 257, shore, 26, 1$ (L). Sta 258, shore, 15
160<br />
EASTERN ATLANTIC SPECIES.—Seven, six of<br />
which occur off tropical <strong>West</strong> Africa. Several of<br />
the names used by Monod (1956) have been<br />
changed, as follows:<br />
Name in Monod Current Name<br />
Pilumnoplax atlantica<br />
Pilumnoplax oxyacantha<br />
Carcinoplax bamardi<br />
Goneplax angulata<br />
Typh locarcinodes<br />
Machaerus atlanticus*<br />
Machaerus oxyacantha*<br />
Carcinoplax bamardi*<br />
Goneplax rhomboides<br />
Typhlocarcinodes inUgrifrons*<br />
integrifrons<br />
Acidops cessaci Acidops cessacii<br />
Hexapus (Thaumastoplax) Thaumastoplax anomalipes<br />
anomalipes (Hexapodidae)<br />
(Goneplacidae)<br />
Hexapus (Pseudohexapus) Pseudohexapus platydactylus<br />
platydactylus (Hexapodidae)<br />
(Goneplacidae)<br />
Hexapus (Parahexapus) Parahexapus africanus<br />
africanus (Hexapodidae)<br />
(Goneplacidae)<br />
Hexapus {Hexapus) Hexapinus buchanani<br />
buchanant (Hexapodidae)<br />
(Goneplacidae)<br />
The four species assigned by Monod to Hexapus<br />
(Goneplacidae: Hexapodinae) are herein assigned<br />
to the Hexapodidae (pp. 166-181).<br />
The extralimital species is Eucrate crenata de<br />
Haan, 1835. An Indo-<strong>West</strong> Pacific immigrant<br />
into the eastern Mediterranean (Holthuis and<br />
Gottlieb, 1958; Ramadan and Dowidar, 1976).<br />
Subfamily CARCINOPLACINAE H. Milne<br />
Edwards, 1852<br />
Genus Carcinoplax H. Milne Edwards, 1852<br />
Curtonotus de Haan, 1833:4 [a subgenus established without<br />
included nominal species; an invalid junior homonym of<br />
Curtonotus Stephens, 1827 (Coleoptera)].<br />
Carcinoplax H. Milne Edwards, 1852:164 [substitute name<br />
for Curtonotus de Haan, 1833; type-species: Cancer (Curtonotus)<br />
longimanus de Haan, 1833, by subsequent designation<br />
by Glaessner, 1929:111; gender: feminine].—Guinot,<br />
1969b:524 [revision]; 1971:1081 [list of species].<br />
• Carcinoplax bamardi Capart, 1951<br />
Carcinoplax bamardi Capart, 1951:170, fig. 65, p!. 3: figs. 5,<br />
12.—Monod, 1956:351, figs. 456-461.—Forest, 1963:627,<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
628.—Maurin, 1968b:484.—Guinot, 1969b:526 [discussion];<br />
1971:1081 [listed].<br />
MATERIAL EXAMINED.—Pillsbury Material: Ivory<br />
Coast: Sta 44, 403-586 m, hard dark gray mud, 16* (L). Sta<br />
51, 329-494 m, 26, 19, 1 juv (L).<br />
Nigeria: Sta 255, 264-269 m, 16\ 19 ov (L). Sta 256, 409-<br />
485 m, 1$ (L).<br />
Gerommo Material: Gabon: Sta 179, 293 m, 2? (W). Sta<br />
198, 300 m, 2c5, 2$ (W). Sta 203, 200 m, 19 (W).<br />
DESCRIPTION.—Capart, 1951:170.<br />
Figures: Capart, 1951, fig. 65, pi. 3: figs. 5, 12;<br />
Monod, 1956, figs. 456-461.<br />
Male Pleopod: Capart, 1951, pi. 3: figs. 5, 12<br />
(Angola); Monod, 1956, figs. 460, 461 (Senegal).<br />
Color: Capart (1951:172) gave the following<br />
notes on color: "Coloration rose bistre, des taches<br />
noires a l'extremite des doigts des pinces."<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 15 to 28 mm; the carapace width<br />
of the single ovigerous female is 23 mm.<br />
BIOLOGY.—Carcinoplax bamardi is a deep water<br />
species, living in depths between 200 m and at<br />
least 500 m; there is one record for the species in<br />
depths ranging from 403 to 586 m. Of 16 available<br />
depth records for the species, there are only<br />
five from depths of less than 300 m, none shallower<br />
than 200 m. It has been collected on mud,<br />
sandy mud, or hard dark gray mud.<br />
Ovigerous females have been collected in<br />
March and May (Capart, 1951; Pillsbury).<br />
DISTRIBUTION.—<strong>West</strong> Africa, from scattered localities<br />
between Spanish Sahara and Angola, in<br />
depths between 200 m and at least 500 m (403-<br />
586 m). The species has not previously been<br />
recorded from off Nigeria or Gabon, although<br />
these records are well within its known range.<br />
Monod (1956), who was the second to record the<br />
species, reported material from Senegal; records<br />
in the literature since 1956 include the following:<br />
Spanish Sahara: Off Villa Cisneros, 300-500 m (Maurin,<br />
1968b).<br />
Ivory Coast: 04°54'N, 03°23'W, 380-400 m (Forest,<br />
1963).<br />
Ghana: 04°16'S, 02°09'30"W, 380-400 m (Forest, 1963).
NUMBER 306 161<br />
Subfamily EURYPLACINAE Stimpson, 1871<br />
Genus Machaerus Leach, 1818<br />
Machaerus Leach, 1818, in 1817-1818:413 [a genus established<br />
without included nominal species; type-species: Pilumnoplax<br />
oxyacantha Monod, 1956, by present designation;<br />
gender: masculine].<br />
DEFINITION.—Carapace smooth, convex anteroposteriorly,<br />
broader than long, with 3 or 4<br />
pairs of anterolateral teeth, outer orbital tooth<br />
distinct from first anterolateral tpoth. Frontal<br />
margin straight, short, less than x k carapace<br />
width, transversely grooved, with median notch.<br />
Supraorbital margin with 2 closed incisions. Basal<br />
antennal segment not fused to front, lacking lateral<br />
projection into orbit, flagellum in orbit. Male<br />
abdomen with 7 somites, third covering width of<br />
sternum, terminal somite elongate triangular.<br />
Chelipeds stout, slightly unequal. Walking legs<br />
slender, compressed, elongate, distal 3 segments<br />
setose dorsally and ventrally. Male first pleopod<br />
stout, curved, tapering distally, spinulose subapically.<br />
REMARKS.—Leach (1818, in 1817-1818:413)<br />
introduced the generic name Machaerus with the<br />
following remarks: il Machaerus; a new genus allied<br />
to Gonoplax, but differing in having short peduncles<br />
to its eyes, which are inserted into the same<br />
part of the shell as in that genus." The name<br />
apparently has been overlooked by most carcinologists;<br />
it was mentioned by Monod (1956:351,<br />
footnote). We believe that two <strong>West</strong> <strong>African</strong> goneplacid<br />
crabs placed in Pilumnoplax by Monod<br />
(1956), P. atlantica Miers, 1881, and P. oxyacantha<br />
Monod, 1956, but subsequently excluded from<br />
Pilumnoplax and the related Neopilumnoplax, should<br />
be assigned to Machaerus.<br />
The genus Pilumnoplax Stimpson, 1858, in<br />
which the two species here assigned to Machaerus<br />
were placed by Monod (1956), has as its typespecies<br />
Pilumnoplax sulcatifrons Stimpson, 1858<br />
(designated by Rathbun, 1918:21). Tesch (1918:<br />
154, 155) assigned P. sulcatifrons to Eucrate de<br />
Haan, 1835, making Pilumnoplax Stimpson a subjective<br />
junior synonym of Eucrate de Haan. As<br />
noted by Guinot (1969c:688) carcinologists, including<br />
Balss (1957:1656) and Glaessner (1969:<br />
R527) have continued to employ Pilumnoplax<br />
sensu Miers, 1886.<br />
Serene recognized that a new genus was required<br />
for Pilumnoplax sensu Miers, and in his<br />
checklist of the Indo-<strong>West</strong> Pacific Brachyura,<br />
Serene (1968:90) first introduced Neopilumnoplax<br />
as a manuscript name; it is a nomen nudum.<br />
Serene considered that five Indo-<strong>West</strong> Pacific<br />
species belonged to Neopilumnoplax: Pilumnus heterochir<br />
Studer, 1882, Pilumnoplax sinclairi Alcock,<br />
1899, Pilumnoplax abyssicola Miers, 1886, Pilumnoplax<br />
cooki Rathbun, 1911, and, with a questionmark,<br />
Pilumnoplax acanthomerus Rathbun, 1911.<br />
Neopilumnoplax Serene, 1969, was named in<br />
Guinot (1969c:689, footnote); its type-species is<br />
Pilumnus heterochir Studer, 1882. Neopilumnoplax<br />
was "established to include the species [of Pilumnoplax]<br />
described after Stimpson (1858)" (Serene,<br />
in Guinot, 1969c:689, footnote). Guinot noted<br />
that Serene included several species in this genus.<br />
She, however, restricted it to Pilumnus heterochir,<br />
Pilumnoplax americana Rathbun, 1898, and, possibly,<br />
to Pilumnoplax sinclairi Alcock, 1899 (erroneously<br />
spelled sainclairi throughout Guinot). Of<br />
the five species listed in Neopilumnoplax by Serene<br />
in 1968, two, P. heterochir and P. sinclairi, were<br />
retained in Neopilumnoplax by Guinot (1969c:689,<br />
footnote), two, Pilumnoplax abyssicola Miers, 1886,<br />
and P. cooki Rathbun, 1911, were transferred to<br />
Carcinoplax, and one, Pilumnoplax acanthomerus<br />
Rathbun, 1911, was not assigned to any genus.<br />
Guinot (1969c:689) made the following comment<br />
about that species: "C'est un Crabe enigmatique<br />
dont nous nous demandons s'il s'agit d'une<br />
Goneplacidae."<br />
Guinot (1969b:517) studied the two <strong>West</strong> <strong>African</strong><br />
species here assigned to Machaerus and noted<br />
that "les deux especes ouest-africaines decrites<br />
dans le genre Pilumnoplax Stimpson a savoir P.<br />
atlantica Miers, 1881 (p. 259; cf. MONOD, 1956, p.<br />
341, fig. 442-449) et P. oxyacantha Monod, 1956<br />
(p. 346, fig. 450-455), offrent tous les caracteres<br />
communs a Euryplax-Eucrate-Frevillea et doivent<br />
etre rattaches aux Euryplacinae." She went on to
162<br />
discuss differences between these two species and<br />
those genera and concluded that "pour le moment,<br />
nous laisserons atlantica et oxyacantha sans<br />
attribution generique: en effet, ni Eucrate, ni Hettroplax,<br />
ni Euryplax, ni Frevillea ne peuvent vraiment<br />
convenir."<br />
Machaerus appears to show closest affinities with<br />
Neopilumnoplax Serene, differing primarily in having<br />
the outer orbital tooth distinct from the first<br />
anterolateral tooth and a much slenderer male<br />
abdomen, the terminal somite of which resembles<br />
a slender isosceles triangle, much longer than<br />
broad.<br />
•Machaerus atlanticus (Miers, 1881),<br />
new combination<br />
Pilumnoplox atlantica.—Monod, 1956:341, figs. 442-449.—Buchanan,<br />
1958:24.—Longhurst, 1958:88.—Gauld, 1960:<br />
70.—Forest and Guinot, 1966:85.—Crosnier, 1969:531.—<br />
Guinot, 1969b:517, 518 [discussion]; 1969c:688 [discussion].—Uschakov,<br />
1970:455 [listed].<br />
"Pilumnoplax" atlantica.—Guinot, 1969b:507, 508 [discussion],<br />
figs. 36, 53 [atlantica Stimpson in legends].<br />
[Pilumnoplax] atlantica.—Guinot, 1971:1081 [listed].<br />
MATERIAL EXAMINED.—Pillsbury Material: Ivory<br />
Coast: Sta 46, 38-42 m, mud with dense Jullienella, 1
NUMBER 306 163<br />
Sierra Leone, and Ghana. Since 1956 the species MEASUREMENTS.—Our specimens have cara-<br />
has been recorded from the following:<br />
pace widths of 7 to 27 mm; the single ovigerous<br />
Senegal: Joal, 10-11 m (Guinot, 1969b).<br />
Guinea: No specific locality (Uschakov, 1970). 09°40'N,<br />
14°05'W, 18 m (Forest and Guinot, 1966).<br />
female has a carapace width of 26 mm.<br />
BIOLOGY.—Machaerus oxyacantha, like M. atlanticus,<br />
is a sublittoral, shallow shelf species that<br />
Sierra Leone: OfT Sierra Leone, 6-30 m (Longhurst, prefers various types of mud bottoms including<br />
1958).<br />
sandy mud and mud with Foraminifera or shells,<br />
Ghana: OfT Accra, 8-20 fm (15-37 m) (Buchanan, 1958);<br />
10 and 36 m (Gauld, 1960). 04°37'N, 00°50'W, 90-100 m<br />
(Forest and Guinot, 1966).<br />
Nigeria: Off the mouths of the Niger River, 04°03'N,<br />
06°12'E, 32 m (Forest and Guinot, 1966).<br />
Congo: Pointe-Noire, 5-10 m (Crosnier, 1969).<br />
in depths between 7-8 m and 73 m. Of 50 depth<br />
records available, only one (7-8 m) is shallower<br />
than 10 m and only five are deeper than 50 m;<br />
88% of the depth records fall between 13 m (13-<br />
26 m) and 50 m. The species may occur in<br />
brackish water but it apparently prefers normal<br />
*Machaerus oxyacantha (Monod, 1956),<br />
new combination<br />
sea water: It was collected by the Pillsbury in<br />
brackish water off Nigeria, and Monod (1956)<br />
reported one specimen from Ghana taken in a<br />
Pilumnoplax atlantica.—Capart, 1951:166, fig. 63 [not Pilum- river seine.<br />
noplax atlantica Miers, 1881].<br />
Ovigerous females have been recorded in Jan-<br />
Pilumnoplax oxyacantha Monod, 1956:346, figs. 450-455.— uary, April, May, and December, suggesting that<br />
Gauld, 1960:70.—Rossignol, 1962:118.—Guinot and Ri- the species may spawn all year (Monod, 1956).<br />
beiro, 1962:63.—Crosnier, 1964:38, fig. on pi. A.—Forest<br />
DISTRIBUTION.—<strong>West</strong> Africa, from Mauritania<br />
and Guinot, 1966:85.—LeLoeuff and Intes, 1968:31, table<br />
1.—Guinot, 1969b:517, 518 [discussion]; 1969c:688 [dis-<br />
to Angola, possibly including Principe Island in<br />
cussion].<br />
the Gulf of Guinea; sublittoral, from 7-8 m to<br />
"Pilumnoplax" oxyacantha.—Guinot, 1969b:507, 508 [discus- 73 m. Monod (1956) recorded material from Sension],<br />
figs. 38, 42, 54, 55.<br />
egal and Ghana. Records in the literature since<br />
[Pilumnoplax} oxyacantha.—Guinot, 1971:1081 [listed]. 1956 include the following:<br />
MATERIAL EXAMINED.—Pillsbury Material: Ivory<br />
Coast: Sta 46, 38-4^ m, mud with denseJullienella, 46 (W).<br />
Sta 47, 37 m, bottom with Jullienella, \6, 1 juv (L). Sta 48,<br />
22 m, Id (L).<br />
Ghana: Sta 16, 46 m, mud with Foraminifera, shells,<br />
several specimens (L). Sta 28, 49-53 m, 1$, 1 carapace (W).<br />
Nigeria: Sta 241, 59-63 m, mud and shell, 3
164 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
rhomboides Linnaeus, 1758, by monotypy; gender: feminine;<br />
name 313 on Official List].—Guinot, 1969b:520<br />
[comments on genus]; 1971:1081 [list of species].<br />
Goneplat Leach, 1814:393, 430 [invalid original spelling for<br />
Goneplax Leach, 1814; name 482 on Official Index].<br />
Gonoplax Leach, 1816:409, 413 [invalid spelling of Goneplax<br />
Leach, 1814; name 483 on Official Index].<br />
Frevillea A. Milne Edwards, 1880:15 [type-species: Frevillea<br />
barbata A. Milne Edwards, 1880, by subsequent designation<br />
by Rathbun, 1918:25; gender: feminine].<br />
Goneplax rhomboides (Linnaeus, 1758)<br />
Cancer rhomboides Linnaeus, 1758:626.<br />
Goneplax angulata.—Capart, 1951:168, fig. 64, pi. 3: figs. 4, 9<br />
[England], 10.—Barnard, 1954:126 [discussion].—Monod,<br />
1956:354, figs. 462-465.—Peres, 1964:27, 28, 29.—<br />
Maurin, 1968a: 19 [Spain]; 1968b:482, 489.<br />
Goneplax rhomboides.—Capart, 1951, pi. 3: fig. 11 [Nice].—<br />
Forest and Games, 1960:353.—Guinot and Ribeiro, 1962:<br />
63.—Forest and Guinot, 1966:86.—Zariquiey Alvarez,<br />
1968:414, figs, le, 138a,b [Spain; references].—Crosnier,<br />
1970:1215 [listed], 1216.—Tiirkay, 1976a:25 [listed], 38,<br />
fig. 28 [Portugal, in part]; 1976b:61 [listed], 71.<br />
Goneplax.—Maurin, 1968a: 14 [Spain].<br />
Goneplax rhomboides.—Le Loeuff and Intes, 1968, table 1.<br />
SYNONYMS.—Cancer angulatus Pennant, 1777;<br />
Ocypode bispinosa Lamarck, 1801; Ocypode longimana<br />
Latreille, 1803.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Undaunted Material: Angola: Sta 94, 90 m, 1
NUMBER 306 165<br />
33°19.3'N, 08°39'W, 85 m; 33°17.2'N, 08°34.5'W, 65 m;<br />
31°01'N, 10° 16'W, 360-375 m; and 3O°2O'N, 10°03'W, 255-<br />
355 m (Turkay, 1976a). Off Agadir, 60-130 m; ofTEssaouira,<br />
32°08'N, 09°02'W, 33 m; and 31°37'N, 09°54'W, 70 m<br />
(Forest and Games, 1960).<br />
Spanish Sahara: Off Cabo Garnet, 400-700 m (Maurin,<br />
1968b).<br />
Mauritania: Off Tamzak (as Tamxat), 200-400 m<br />
(Maurin, 1968b).<br />
Ivory Coast: Off Grand-Bassam, 200 m (Le Loeuff and<br />
Intes, 1968).<br />
Ghana: O4°37'N, 00°50'W, 90-100 m (Forest and Guinot,<br />
1966).<br />
Angola: Baia dos Tigres, 110 m (Guinot and Ribeiro,<br />
1962). 16°27'S, 11°35'E, 90 m; and 16°37'S, 11°22'E, 126<br />
m (Crosnier, 1970).<br />
Subfamily RHIZOPINAE Stimpson, 1858<br />
Genus Acidops Stimpson, 1871<br />
Acidops Stimpson, 1871b: 110 [p. 137 on separate] [type-species:<br />
Acidops fimbriatus Stimpson, 1871, by monotypy; gender:<br />
masculine].<br />
Epimelus A. Milne Edwards, 1878:227 [p. 8 on separate]<br />
[type-species: Epimelus cessacii A. Milne Edwards, 1878, by<br />
monotypy; gender: masculine; name 147 on Official List].<br />
Acidops cessacii (A. Milne Edwards, 1878)<br />
Acidops cessaci.—Monod, 1956:359, figs. 469, 470 [Senegal].—Guinot<br />
and Ribeiro, 1962:64, figs. 27, 28 [Cape<br />
Verde Islands].—Ribeiro, 1964:13 [Cape Verde Islands].—Forest<br />
and Guinot, 1966:88 [Sao Tome, Annobon].<br />
Epimelus cessaci.—Garth, 1968:314 [discussion].<br />
DISTRIBUTION.—<strong>West</strong> Africa, from the Cape<br />
Verde Islands, Senegal, and Sao Tome and Annobon<br />
islands in the Gulf of Guinea; intertidal to<br />
10-30 m.<br />
Subfamily TYPHLOCARCINOPINAE Rathbun,<br />
1909<br />
Genus Typhlocarcinodes Alcock, 1900<br />
Typhlocarcinodes Alcock, 1900:326 [a genus established without<br />
included nominal species; type-species: Typhlocarcinus<br />
integrifrons Miers, 1881, by subsequent designation by<br />
Tesch, 1918:227; gender: masculine].<br />
REMARKS.—Alcock (1900) established this genus<br />
for an Indian species that he did not name<br />
and that he compared with the <strong>West</strong> <strong>African</strong><br />
Typhlocarcinus integrifrons Miers, 1881. The genus<br />
was established without any included nominal<br />
species. Tesch (1918:226, 227) considered that<br />
Typhlocarcinodes included four species: T. integrifrons<br />
(Miers, 1881), Caecopilumnus hirsutus Borradaile,<br />
1903, Typhlocarcinops piroculata Rathbun,<br />
1911, and Typhlocarcinodes crassipes Tesch, 1918;<br />
he designated T. integrifrons as the type-species.<br />
According to Article 69(a)(ii) of the International<br />
Code of Zoological Nomenclature, "if no nominal species<br />
were included at the time the genus was<br />
established, the nominal species-group taxa that<br />
were first subsequently and expressly referred to<br />
it are to be treated as the only original included<br />
species." Thus Tesch's designation of Typhlocarcinus<br />
integrifrons Miers as the type-species of Typhlocarcinodes<br />
is valid.<br />
Monod (1956:365) questioned whether the<br />
Indo-<strong>West</strong> Pacific species were congeneric with<br />
T. integrifrons. Tesch (1918:227) and Serene<br />
(1964b:237) pointed out differences between T.<br />
integrifrons and Indo-<strong>West</strong> Pacific species, and<br />
Takeda (1973:52) accepted the possibility that<br />
two genera might be involved.<br />
Those authors who consider the three Indo-<br />
<strong>West</strong> Pacific species assigned to this genus, Typhlocarcinodes<br />
hirsutus (Borradaile, 1902), T. piroculatus<br />
(Rathbun, 1911), and T. crassipes Tesch,<br />
1918, to be generically distinct from the eastern<br />
Atlantic T. integrifrons (Miers, 1881) should refer<br />
those species to Caecopilumnus Borradaile (1902b:<br />
267; type-species: Caecopilumnus hirsutus Borradaile,<br />
1902, by monotypy; gender: masculine).<br />
* Typhlocarcinodes integrifrons (Miers, 1881)<br />
Typhlocarcinodes integrifrons.—Monod, 1956:356, 632, figs.<br />
466-468—Longhurst, 1958:88.—Forest and Guinot,<br />
1966:87, fig. 8.—Crosnier, 1967:334.<br />
MATERIAL EXAMINED.—PUlsbury Material. Ghana:<br />
Sta 24, 35-37 m, dark red bryozoans, 16", 19 (L).<br />
DESCRIPTION.—Miers, 1881a:260.
166<br />
Figures: Monod, 1956, figs. 466-469; Forest and<br />
Guinot, 1966, fig. 8.<br />
Male Pleopod: Forest and Guinot, 1966, fig. 8<br />
(Principe).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 6 to 9 mm.<br />
BIOLOGY.— Typhlocarcinodes integrifrons is a sublittoral<br />
species, known to occur in depths between<br />
12 and 90-100 m. It may prefer shallower water,<br />
as suggested by the only depth records available,<br />
viz. 16-44, 18, 30, 30-34, 35-37, 48, 51, and 90-<br />
100 m.<br />
Bottom types on which it is known to occur<br />
include: dark red bryozoans, 35-37 m (Pillsbury);<br />
shelly mud (2 specimens) and shelly sand (4<br />
specimens) in 16-44 m (Longhurst, 1958); sand<br />
and mud in 12 m, mud and shells in 30 m, and<br />
rocks and shells in 90-100 m (Forest and Guinot,<br />
1966); and coarse sand in 30 m and green mud in<br />
48 m (Crosnier, 1967).<br />
Ovigerous females have been collected in May<br />
(Forest and Guinot, 1966).<br />
DISTRIBUTION.—<strong>West</strong> Africa, from localities between<br />
Senegal and Dahomey as well as Principe<br />
Island in the Gulf of Guinea; sublittoral, from<br />
12 m to 90-100 m. Records in the literature<br />
include the following:<br />
Senegal: Goree (Miers, 1881a; Monod, 1956).<br />
Guinea: 1 mi [ 1.6 km] SE of lie Blanche, off Conakry,<br />
18 m (Monod, 1956).<br />
Sierra Leone: No specific locality, in 44 m (Monod, 1956);<br />
in 16-44 m (Longhurst, 1958).<br />
Ivory Coast: 04°36'N, 06°50^, 48 m (Crosnier, 1967).<br />
Ghana: 04°37'N, OO^O'W, 90-100 m (Forest and Guinot,<br />
1966).<br />
Dahomey: 06°04'N, 02°26'E, 30 m (Crosnier, 1967).<br />
Principe: 01°39'35"'N, 07°26'53"E, 12 m, and 01°37'N,<br />
O7°22'E, 30 m (Forest and Guinot, 1966).<br />
Family HEXAPODIDAE Miers, 1886<br />
HEXAPODINAE Miers, 1886:275.<br />
EASTERN ATLANTIC GENERA.—Four, Hexapinus,<br />
new genus, Parahexapus, Pseudohexapus, and Thaumastoplax,<br />
each of which is represented by a tropical<br />
<strong>West</strong> <strong>African</strong> species.<br />
EASTERN ATLANTIC SPECIES.—Four, all occur-<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
ring off <strong>West</strong> Africa. This family was not represented<br />
in the Pillsbury collections. The following<br />
name changes have occurred since Monod (1956):<br />
Name in Monod Current Name<br />
Hexapus (Thaumastoplax) Thaumastoplax anomalipes<br />
anomalipes<br />
Hexapus (Pseudohexapus) Pseudohexapus platydactylus<br />
platydactylus<br />
Hexapus (Parahexapus) Parahexapus afneanus<br />
africanus<br />
Hexapus (Hexapus) Hexapinus buchanani<br />
buchanani<br />
REMARKS.—The subfamily Hexapodinae was<br />
established within the Family Pinnotheridae by<br />
Miers (1886), who included four genera: Hexapus<br />
de Haan, 1833; Amorphopus Bell, 1859; Thaumastoplax<br />
Miers, 1881; and Asthenognathus Stimpson,<br />
1858. The latter genus is now considered to be a<br />
pinnotherid (Schmitt, McCain and Davidson,<br />
1973:128). Ortmann (1894:690) retained the<br />
Hexapodinae within the Pinnotheridae. Alcock<br />
(1900:287) recognized the affinities of the Hexapodinae<br />
with the Goneplacidae and included it<br />
as one of five subfamilies of the latter. He was<br />
followed in this by Borradaile (1907) and subsequent<br />
authors.<br />
We believe that the complete suppression of<br />
the last pair of pereiopods in members of this<br />
subfamily is a fundamental character which warrants<br />
the recognition of the Hexapodidae as a<br />
family distinct form but related to the Goneplacidae.<br />
We treat it as a family here.<br />
The following genera and species have been<br />
named in the family: Hexapus was erected by de<br />
Haan (1833:5; 1835:63) for Cancer sexpes Fabricius,<br />
1798; it is apparent that several different<br />
species have erroneously been identified with H.<br />
sexpes by subsequent authors. Other species described<br />
in Hexapus include: H. williamsi Glassell,<br />
1938, eastern Pacific; H. stebbingi Barnard, 1947,<br />
South Africa; H. buchanani Monod, 1956, <strong>West</strong><br />
Africa; H. granuliferus Campbell and Stephenson,<br />
1970, Australia; and H. estuarinus Sankarankutty,<br />
1975, India. Each of these latter species are assigned<br />
to different genera below.
NUMBER 306 167<br />
Amorphopus (Bell, 1859:27) was established for<br />
a single species from an unknown locality, A.<br />
cylindraceus Bell. According to Bell, it differs from<br />
Hexapus in having a rudimentary pair of fifth legs,<br />
a feature shared with no other member of the<br />
family. Rathbun (1897b: 163) pointed out that<br />
this name was preoccupied twice [according to<br />
Neave, Nomenclator Zoologicus, the oldest name is<br />
Amorphopus Audinet-Serville, 1838 (Insecta)] and<br />
substituted Paeduma for it. Rathbun's name apparently<br />
has been overlooked by nearly all zoologists<br />
dealing with this group; Gordon (1971:108)<br />
pointed out that Amorphopus was preoccupied.<br />
Amorphopus Bell has been recognized by Alcock<br />
(1900:293), Stebbing (1910:315) (who suggested<br />
that "there seems little justification for the separation<br />
either of Amorphopus or Thaumastoplax from<br />
Hexapus."), Tesch (1918:238), Balss (1957:1658),<br />
and Serene (1968:93). We recognize Paeduma<br />
Rathbun (p. 173).<br />
Thaumastoplax Miers (1881a:261) was established<br />
for a single species from Senegal, 71 anomalipes<br />
Miers; it was distinguished from Hexapus in<br />
having a slender third maxilliped with the propodus<br />
dilated distally. Rathbun (1910:113) described<br />
two additional species from Thailand, T.<br />
orientalis and T. chuenensis. A fourth species, T.<br />
spiralis Barnard (1950:301), was described from<br />
South Africa. Although Monod (1956:362, 363)<br />
treated Thaumastoplax as a subgenus of Hexapus,<br />
Tesch (1918:238), Balss (1957:1658), Glaessner<br />
(1969:R527) and Sakai (1976:555) all considered<br />
it to be a distinct genus. We also consider Thaumastoplax<br />
to be a valid genus; species described in<br />
it subsequent to 1881 are assigned to other genera<br />
below.<br />
Lambdophallus Alcock (1900:329) was based on<br />
an Indian species, L. sexpes Alcock, in which the<br />
apices of the male pleopods extended laterally<br />
well beyond the abdomen, lying in deep lateral,<br />
transverse grooves in the sternum. A second species,<br />
Lambdophallus anfractus, was described on material<br />
from Thailand by Rathbun (1909:113).<br />
Stephensen (1945:182) considered Lambdophallus<br />
to be a synonym of Hexapus and synonymized<br />
both L. sexpes and L. anfractus with H. sexpes.<br />
Monod (1956:362) and Sakai (1976:554) both<br />
considered Lambdophallus to be a subgenus of Hexapus;<br />
Tesch (1918:239) and Balss (1957:1658) both<br />
considered it to be a distinct genus, as we do.<br />
Hexaplax Doflein (1904:122) was established for<br />
a deep water species from India, H. megalops<br />
Doflein. It is characterized by long slender walking<br />
legs and very large, hammer-shaped eyes. It<br />
has been considered a distinct genus by all subsequent<br />
authors (see Sakai, 1976:556, for synonymy).<br />
Parahexapus was erected by Balss (1922:77) for<br />
P. africanus from Cabinda; one of its distinguishing<br />
features is its slender third maxilliped. Balss<br />
(1957:1658) retained Parahexapus as a distinct genus,<br />
but Monod (1956:362) considered it to be a<br />
subgenus of Hexapus. We believe that it is generically<br />
distinct.<br />
Pseudohexapus was recognized by Monod (1956:<br />
362, 365) as a subgenus of Hexapus. It included<br />
but one species from <strong>West</strong> Africa, Hexapus {Pseudohexapus)<br />
platydactylus, characterized by broadened,<br />
flattened dactyli on the walking legs. We<br />
recognize Pseudohexapus here.<br />
Monod (1956), who studied four species representing<br />
four different subgenera, reviewed the<br />
literature and pointed out numerous discrepancies<br />
in accounts in the literature purporting to<br />
deal with the same species. He pointed out that<br />
Stephensen's (1945:182) H. sexpes probably was<br />
not the same species reported by de Haan and<br />
provided diagnoses of five subgenera within Hexapus:<br />
Hexapus, Thaumastoplax, Parahexapus, Pseudohexapus,<br />
and Lambdophallus.<br />
<strong>West</strong> <strong>African</strong> material available to us includes<br />
representatives of only two species, Thaumastoplax<br />
anomalipes and Parahexapus africanus. The collections<br />
of the <strong>Smithsonian</strong> <strong>Institution</strong>, however,<br />
include specimens identified with several additional<br />
species from Indo-<strong>West</strong> Pacific localities:<br />
Hexapus sexpes (Fabricius) from Japan and the<br />
Philippine Islands, Thaumastoplax orientalis (Rathbun)<br />
from Japan, paratypes of Lambdophallus anfractus<br />
Rathbun from Thailand, and Hexapus williamsi<br />
Glassell from Mexico. Examination of this<br />
relatively rich material, which proved to be nee-
Igg SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
essary in order to determine the status of the slender or distally dilated, and the presence or<br />
species known from <strong>West</strong> Africa, has suggested to absence of a flagellum on the exopod; eye shape;<br />
us that all of the subgenera recognized by Monod relative length of walking legs; shape and configshould<br />
be recognized as distinct genera, that sev- uration of the male abdomen; the shape and<br />
eral new genera should be recognized, and that extent of sternal grooves in the male, and strucseveral<br />
name changes are required for known tureof the male pleopod. These characters appear<br />
species. All existing genera and the new genera to us to afford reliable features for differentiation<br />
that we believe should be recognized are charac- of genera.<br />
terized below. Gordon (1971:108) was correct in stating that<br />
Monod's (1956:361) excellent review of the "the taxonomy of the Hexapodinae poses some<br />
subgenera of Hexapus stressed the importance of problems which can only be solved by a careful<br />
several characters that appear to have been over- re-examination of all of the available material."<br />
looked in the past; several of these have not been We believe that all records in the literature redescribed<br />
for all species. They include: the shape quire verification. Descriptions of several new<br />
of the third maxillipeds, whether slender or species and a redescription of the type of Cancer<br />
broad, and especially the shape of the propodus, sexpes Fabricius are in preparation by Manning.<br />
Key to Genera of Hexapodidae<br />
1. Eye hammer-shaped, cornea very large [carpus, propodus, and dactylus<br />
of third maxilliped slender; walking legs long, slender, merus of longest<br />
pair longer than carapace] Hexaplax<br />
Eye small, reduced, cornea scarcely or not at all broader than stalk ... 2<br />
2. Eyes fixed. Walking legs (Figure 34a) long, slender, merus of longest pair<br />
longer than carapace. Male with long, narrow transverse grooves in<br />
sternum extending laterally from abdominal fossa beyond bases of third<br />
maxillipeds (Figure 34b) (carpus, propodus, and dactylus of third<br />
maxilliped slender) ". . . Lambdophallus<br />
Eyes movable. Walking legs short, merus of longest pair shorter than<br />
carapace. Male lacking long, narrow transverse grooves in sternum<br />
extending laterally from abdominal fossa beyond bases of third<br />
maxillipeds 3<br />
3. Propodus of third maxilliped strongly dilated distally (Figure 31b, 39a) . 4<br />
Propodus of third maxilliped not strongly dilated distally, slender, like<br />
carpus and dactylus (Figures 31a, 33a) 6<br />
4. Male pleopod stout, strongly twisted into a tight spiral (Figure<br />
37*) Spiroplax, new genus<br />
Male pleopod variously shaped, not twisted into a tight spiral 5<br />
5. Third to fifth abdominal somites fused in male. Male pleopod (Figure<br />
39c) slender, sinuous, apex directed anteriorly Thaumastoplax<br />
Third and fourth and fourth and fifth abdominal somites fused in male.<br />
Male pleopod (Figure 35b) slender, strongly recurved posteriorly, apex<br />
directed posteriorly or recurved anteriorly, shaped like a figure<br />
° Paeduma<br />
6. Male abdomen with second through sixth somites fused<br />
Stevea, new genus
NUMBER 306 169<br />
Male abdomen with third to fifth somites fused 7<br />
7. Dactylus of each walking leg enlarged, triangular, strongly flattened<br />
dorsoventrally (male pleopod stout, with proximally directed subapical<br />
spines) Pseudohexapus<br />
Dactyli of walking legs not enlarged or strongly flattened dorsoventrally<br />
8<br />
8. Terminal somite of male abdomen (Figure 40a,d) trilobed or trefoil in<br />
shape, with distinct lateral angular projections (sixth abdominal somite<br />
divided longitudinally in type-species; male pleopod stout, bent laterally,<br />
apex directed anteriorly) Tritoplax, new genus<br />
Terminal somite of male abdomen (Figures 3\b, 32c) broadly rounded,<br />
not trilobed 9<br />
9. Third maxilliped (Figure 36a) slender, almost pediform, ischium width<br />
about % merus length. Male pleopod (Figure 36c) slender, sinuous,<br />
tapering abruptly to almost filiform apex, with strong, recurved spines<br />
near midlength (sixth and seventh somites of male abdomen (Figure<br />
36b) very slender, width half that of first fused somite) . Parahexapus<br />
Third maxilliped broad, ischium width usually subequal to or greater<br />
than merus length 10<br />
10. Sternum in male (Figure 336) with broad, deep grooves extending anterolaterally<br />
from end of abdominal fossa to bases of third maxillipeds.<br />
Male pleopods bent laterally, apices lying in sternal grooves . Hexapus<br />
Sternum in male (Figure 32c) lacking broad, deep grooves extending<br />
anterolaterally from end of abdominal fossa to bases of third maxillipeds,<br />
with at most triangular anterolateral projections on abdominal<br />
fossa, not extending laterally beyond border of first fused somite. Male<br />
pleopods not bent, apices concealed under abdomen or visible on either<br />
side of terminal somite Hexapinus, new genus<br />
Genus Hexapinus, new genus<br />
FIGURES 31, 32c,d<br />
TYPE-SPECIES.—Hexapus latipes de Haan, 1835,<br />
pi. D.<br />
ETYMOLOGY.—The masculine name Hexapinus<br />
is derived from the name Hexapus and the Latin<br />
suffix-j>iMJ, pertaining to.<br />
DIAGNOSIS.—Eyes very small, cornea subglobular.<br />
Third maxilliped (Figure 31a) with broad<br />
ischium and merus, inner margin of ischium concave<br />
proximally, straight or convex distally, carpus,<br />
propodus, and dactylus slender, cylindrical,<br />
dactylus longer than propodus; exopod with flagellum.<br />
Walking legs short, stout, merus of longest<br />
pair shorter than carapace. Male pleopods (Figure<br />
3\c,d) stout, sinous but not strongly angled<br />
distally, apices concealed under abdomen or<br />
scarcely visible next to seventh somite. Male abdomen<br />
(Figure 3\b) with third to fifth somites<br />
fused, terminal somite rounded apically, smooth<br />
laterally. Female abdomen of 7 somites.<br />
INCLUDED SPECIES.—Hexapus latipes de Haan,<br />
1835, from Japan; Hexapus buchanani Monod,<br />
1956, from <strong>West</strong> Africa; Hexapus granuliferus<br />
Campbell and Stephenson, 1970, from Australia;<br />
and one or more species identified with Hexapus<br />
sexpes by various authors: A. Milne Edwards,<br />
1873b:253, pi. 12: fig. 1 (New Caledonia); De<br />
Man, 1888:322, pi. 13: fig. 3 (Amboina); Zehntner,<br />
1894:159 (Amboina); Nobili, 1906a: 146 (Persian<br />
Gulf); Tesch, 1918:240, pi. 17: fig. 1 (Indonesia);<br />
Balss, 1938:74 (Marshall Islands); Stephensen,<br />
1945:182, fig. 53 (Persian Gulf); Guinot-
a<br />
170<br />
FIGURE 31.—Hexapinus buchanam (Monod): a, third maxilliped;<br />
b, male abdomen; c, male pleopod; d, apex of male<br />
pleopod (all from Monod, 1956, figs. 498-500).<br />
Dumortier and Dumortier, 1960:130, fig. 9 (no<br />
locality); Griffin, 1972:85 (Australia).<br />
Hexapus sexpes sensu A. Milne Edwards, 1873(b)<br />
(Figure 32a,6) from New Caledonia differs from<br />
our material of Hexapinus latipes from Japan in<br />
having a narrow cornea and the surface of the<br />
carapace covered with setae. It may well represent<br />
a new genus. Unfortunately, the female holotype<br />
could not be located at the Museum national<br />
d'Histoire naturelle, Paris, in October 1977 (J.<br />
Forest, pers. comm.).<br />
We suspect that two distinct Japanese species<br />
have been referred to H. sexpes by various authors.<br />
De Haan (1835) reported that his material measured<br />
3X5 lines (1 line = 2.12 mm) or 6 X 11<br />
mm; thus it was a relatively small species. Sakai<br />
(1976) noted that some specimens from Japan<br />
measured as large as 18 X 29.2 mm. Available<br />
material of both forms in the collection of the<br />
<strong>Smithsonian</strong> <strong>Institution</strong> exhibit specific differences.<br />
An examination of the Japanese specimen that<br />
de Haan (1835:35, 63, pi. 11: fig. 5) described<br />
and figured as Hexapus sexpes (Fabricius, 1798)<br />
and which still is preserved dry in the collection<br />
of the Rijksmuseum van Natuurlijke Historie,<br />
Leiden (a male, cl 6.7 mm, cb 10.0 mm, Crust.<br />
D. 31783), shows that it is different from Fabricius'<br />
type of Cancer sexpes and should be referred<br />
to the genus Hexapinus. For it the specific epithet<br />
latipes de Haan is available. On plate D of Fauna<br />
Japonica, namely, de Haan (1835) used the name<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Hexapus latipes for a figure of the mouthparts of<br />
his Japanese Hexapus, which he named H. sexpes<br />
in his text and on pi. 11: fig. 5. That the word<br />
latipes is not a typographical error is shown by the<br />
fact that the piece of cardboard on which de<br />
Haan glued the mouthparts of this specimen, and<br />
which is also present in the collection of the<br />
Leiden Museum, bears the following inscription<br />
in de Haan's handwriting: "Hexapus latipes<br />
Faun. Jap." Evidently de Haan at first considered<br />
the species new and gave it the name Hexapus<br />
latipes, and when he later changed his mind and<br />
identified the species with Cancer sexpes, the name<br />
latipes had already been engraved on plate D and<br />
could not be changed. It is peculiar that de Haan<br />
under Hexapus sexpes nowhere referred to plate D<br />
and did not correct the name latipes in the<br />
erratum.<br />
The specific epithet latipes, published in the<br />
combination Hexapus latipes with a figure, is an<br />
available name and must be used for the present<br />
Japanese Hexapinus.<br />
Hexapinus latipes has been discussed under the<br />
name Hexapus sexpes by Tesch (1918), Sakai (1934,<br />
1936, 1939, 1976), and Odawara (1965).<br />
Although de Haan identified his species with<br />
Cancer sexpes, he noted that in his specimens the<br />
FIGURE 32.—Hexapus sexpes sensu A. Milne Edwards: a, dorsal<br />
view; b, front (from A. Milne Edwards, 1873b, pi. 12: fig. 1,<br />
la). Hexapinus granuliferus (Campbell and Stephenson): c,<br />
abdomen of male; d, gonopod (apex broken?) (from Campbell<br />
and Stephenson, 1970, fig. 49g,h).
NUMBER 306 171<br />
front was truncate and deflexed and further noted<br />
(1835:64) that "Fabricius dicit frontem emarginatam."<br />
Hexapinus differs from the other hexapodid genera<br />
with short legs, broad maxillipeds, and the<br />
third to fifth abdominal somites fused in males,<br />
Hexapus and Tritoplax, as follows: In Hexapus sternal<br />
grooves extend anterolaterally from the abdominal<br />
fossa almost to the level of the bases of<br />
the third maxillipeds, and the male pleopods are<br />
bent laterally and are situated in these grooves;<br />
in Hexapinus these grooves are restricted to the<br />
end of the abdominal fossa and the male pleopods<br />
are more or less straight and are concealed under<br />
the abdomen (Figure "52c,d)- In Tritoplax the terminal<br />
somite of the male abdomen is trilobed or<br />
trefoil in shape, whereas in Hexapinus it is evenly<br />
rounded.<br />
We are not certain that Hexapus granuliferus<br />
Campbell and Stephenson, 1970, should be assigned<br />
here, although superficially it resembles<br />
Hexapinus latipes. Their figure (4%) of the male<br />
pleopod (Figure 32
172<br />
tylus longer than propodus; exopod with flagejlum.<br />
Walking legs short, merus of longest leg<br />
(third) shorter than carapace. Male pleopods<br />
(Figure 33c) not concealed under abdomen, lying<br />
in deep, oblique grooves (Figure 33b) on anterior<br />
part of sternum, apices setose, directed anterolaterally.<br />
Male abdomen (Figure 336) with third to<br />
fifth somites fused, terminal somites rounded, not<br />
trilobed. Female abdomen with 7 free somites.<br />
INCLUDED SPECIES.—Cancer sexpes Fabricius,<br />
1798, from southern India; Lambdophallus anfractus<br />
Rathbun, 1909, from Thailand and Japan; and<br />
Hexapus estuarinus Sankarankutty, 1975, from<br />
India.<br />
REMARKS.—De Haan (1835:35, 63, pi. 11: fig.<br />
5, pi. D) included in the genus Hexapus (which he<br />
had established in 1833 without nominal species)<br />
two nominal species: Cancer sexpes Fabricius, 1798,<br />
and his new Hexapus latipes. On plate D, de Haan<br />
used the name Hexapus latipes for the specimen,<br />
which in his text (pages 35 and 63, and pi. 11:<br />
fig. 5), he had indicated with the name Hexapus<br />
sexpes (Fabricius). Hexapus latipes is an available<br />
name, and the epithet latipes has to be used for<br />
the species of Hexapinus to which de Haan's specimen<br />
belongs. This specimen, namely, is specifically<br />
(and generically) distinct from the typespecimen<br />
of Cancer sexpes Fabricius, 1798. As Cancer<br />
sexpes is the type-species of the genus Hexapus<br />
it should be known as Hexapus sexpes (Fabricius,<br />
FIGURE 33 — Hexapus anfractus (Rathbun): a, third maxilliped;<br />
b, sternum of male; c, male pleopod. (All from Rathbun,<br />
1910, fig. 36.)<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
1798); for de Haan's Japanese species the name<br />
Hexapinus latipes (de Haan, 1835) is correct.<br />
De Haan (1835:64) did note differences between<br />
his Japanese specimen, of which he described<br />
the front as deflexed and truncate, and<br />
the description of Cancer sexpes in which "Fabricius<br />
dicit frontem emarginatam."<br />
Fabricius (1798:334) gave the following description<br />
of Cancer sexpes:<br />
C. thorace laevi integerrimo, fronte emarginata, pedibus<br />
senis.<br />
Habitat in India orientali Dom. Daldorff.<br />
Parvus thorax laevis in medio lineolis duabus impressis.<br />
Frons deflexa, emarginata. Pedes omnino sex inermes. Chelae<br />
breves, laeves altera maiore.<br />
The type of Cancer sexpes Fabricius, in the Universitetets<br />
Zoologiske Museum, Copenhagen (see<br />
Zimsen, 1964:650), was examined by Manning in<br />
Copenhagen in 1977. It is a male, cl 6.2 mm, cb<br />
9.5 mm, preserved dry, and in very good condition.<br />
As mentioned by Fabricius, the front is<br />
faintly emarginate. It was compared with the<br />
holotype of Lambdophallus anfractus Rathbun, and<br />
differs from the latter in having the carapace less<br />
granular, the fingers of the chela longer than the<br />
palm, and the walking legs are slenderer and less<br />
setose. There is no doubt, however, that the two<br />
species are congeneric. Both share the deep,<br />
oblique sternal grooves, figured well by Rathbun<br />
(1910, fig. 36a; see Figure 33b herein), with similar<br />
male pleopods. The abdomen of the male in both<br />
species has the third to the fifth somites fused. A<br />
redescription of the type-specimen of Hexapus<br />
sexpes is in preparation.<br />
Hexapus estuarinus Sankarankutty, 1975, from<br />
the Cochin region, SW India, may well prove to<br />
be synonymous with Hexapus sexpes. In both species<br />
the chelae are unequal and the second leg is<br />
the longest of the walking legs.<br />
This genus has deep sternal grooves like Lambdophallus<br />
sensu stricto, but differs in having very<br />
short pereiopods, as in most other genera in the<br />
family. The sternal grooves are deep and well<br />
defined but extend anterolaterally almost to the<br />
base of the maxillipeds rather than extending<br />
laterally as in Lambdophallus (Figure 34b). In H.
NUMBER 306 173<br />
anfractus the third legs are the longest, the male<br />
abdomen extends almost to the level of the insertion<br />
of the third maxillipeds, and the carapace is<br />
relatively broad (width =1.5 times length).<br />
We do not agree with Campbell and Stephenson<br />
(1970:291) that Hexapus anfractus (Rathbun)<br />
can be identified with Hexapinus latipes from Japan.<br />
The sternal grooves in our material of H.<br />
anfractus are much better developed than they are<br />
in our Japanese material of H. latipes, in which<br />
the grooves are represented only by an emargination<br />
in the lateral margins of the sternal abdominal<br />
trench. We refer Japanese specimens previously<br />
assigned to H. sexpes to a new genus, Hexapinus.<br />
In L. anfractus the grooves extend anterolaterally<br />
from the terminal segment of the abdomen<br />
to the bases of the articulations of the third<br />
maxillipeds. The size and extent of development<br />
of these grooves are completely different in the<br />
two forms, and different from that found in Lambdophallus,<br />
sensu stricto, in which they extend laterally<br />
from the last abdominal somite.<br />
Genus Lambdophallus Alcock, 1900<br />
FIGURE 34<br />
Lambdophallus Alcock, 1900:329 [type-species: Lambdophallus<br />
sexpes Alcock, 1900, by monotypy; gender: masculine].<br />
DIAGNOSIS.—Eyes small, immovable, cornea<br />
slightly broader than stalk. Third rnaxilliped<br />
(Figure 34A) broad, ischium slightly convex distally,<br />
carpus, propodus, and dactylus slender, subcylindrical,<br />
dactylus as long as propodus; flagellum<br />
of exopod not described. Pterygostomian<br />
region with row of oblique striae. Walking legs<br />
(Figure 34a) elongate, merus of longest pair longer<br />
than carapace. Transverse grooves on sternum<br />
(Figure 346) for reception of male pleopods present.<br />
Male pleopods (Figure 34b,c) with apices not<br />
concealed under abdomen, each bent at right<br />
angle, distal part of each lying in deep transverse<br />
groove on sternum, apices naked, with subterminal<br />
setae, directed laterally. Male abdomen<br />
(Figure 34b,c) with 5 segments, third to fifth fused,<br />
terminal segment rounded apically, smooth laterally.<br />
REMARKS.—This genus contains only the typespecies,<br />
L. sexpes Alcock. The fixed eyes, long<br />
pereiopods, and long, L-shaped male pleopods<br />
extending laterally into transverse grooves in the<br />
anterior part of the sternum are diagnostic for the<br />
genus.<br />
In L. sexpes the third and fourth legs are subequal<br />
in length, the male abdomen falls short of<br />
the level of insertion of the third maxillipeds, and<br />
the carapace is relatively broad (width = 1.6<br />
times length).<br />
Another species subsequently described in this<br />
genus, L. anfractus Rathbun, 1909, from Thailand,<br />
is transferred to Hexapus (p. 171).<br />
Genus Paeduma Rathbun, 1897<br />
FIGURE 35<br />
Amorphopus Bell, 1859:27 [invalid junior homonym of Amorphopus<br />
Audinet-Serville, 1838 (Insecta); type-species: Amorphopus<br />
cylindraceus Bell, 1859, by monotypy; gender: masculine].<br />
Paeduma Rathbun, 1897b: 163 [substitute name for Amorphopus<br />
Bell, 1859; type-species: Amorphopus cylindraceus Bell,<br />
1859; gender: neuter].<br />
DIAGNOSIS.—Eyes small, movable, cornea<br />
about as broad as stalk. Third maxillipeds slender,<br />
ischium convex mesially, propodus dilated<br />
distally, as broad as merus, longer than carpus or<br />
dactylus; exopod with flagellum. Pterygostomian<br />
region lacking oblique striae. Walking legs short,<br />
merus of longest pair (second) shorter than carapace.<br />
Sternal grooves absent. Apices of male pleopods<br />
(Figure 3bb) concealed under abdomen,<br />
pleopods slender, recurved posteriorly or doubly<br />
recurved into a figure 8, apices apparently naked<br />
or nearly so. Male abdomen (Figure 35a) with<br />
third and fourth and fifth and sixth somites fused,<br />
terminal somite short, subtriangular, rounded apically,<br />
not trilobed.<br />
INCLUDED SPECIES.—Amorphopus cylindraceus<br />
Bell, 1859, locality unknown, and Thaumastoplax<br />
orientalis Rathbun, 1909 and T. chuenensis Rathbun,<br />
1909, both from Thailand.
174<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 34.—Lambdophallus sexpes Alcock: a, dorsal view; b, ventral view; c, male abdomen and<br />
apices of pleopods. (From "Illustrations of the Zoology of H. M. Indian Marine Surveying<br />
Steamer Investigator" Crustacea, 10 (1902), pi. 62: fig. 1-lb.)<br />
REMARKS.—The genus Amorphopus was recognized<br />
by Bell (1859:27) for a single species, A.<br />
cylindraceus, from an unknown locality. Bell's<br />
name Amorphopus is preoccupied, as pointed out<br />
by Rathbun (1897b: 163), who substituted Paeduma<br />
for it. As we noted above, with the exception<br />
of Gordon (1971:108) and Sankarankutty (1975:<br />
4), subsequent authors apparently have overlooked<br />
Paeduma.<br />
Paeduma has remained unidentifiable since its<br />
introduction by Bell; no subsequent workers on<br />
the group have been able to identify any species<br />
with it. As Gordon (1971:108) pointed out, one of<br />
the characters mentioned by Bell, the fusion of<br />
segments of the abdomen, is unusual (i.e., the<br />
third and fourth segments and the fifth and sixth<br />
segments are fused, rather than the third, fourth,<br />
and fifth, as we have found in all other genera of<br />
the family, save two: in Stevea, new genus, the<br />
second through the sixth segments are fused and<br />
in Hexaplax some segments are indistinctly fused).<br />
FIGURE 35.—Paeduma orientate<br />
(Rathbun), male: a, abdomen;<br />
b, pleopod. (Both from Rathbun,<br />
1910, fig. 33.)<br />
However, in her expanded account of Tkaumastoplaxorientalis<br />
Rathbun, 1909, Rathbun (1910:347)<br />
reported an identical configuration of the abdomen<br />
in that species, and Sakai (1939:579) mentioned<br />
it in material from Japan identified with<br />
T. orientalis. Further, Bell (1859:28) also had<br />
noted: "the abdomen very long and narrow, the<br />
first and second joint transversely linear, the third<br />
and fourth united and forming a triangle truncated<br />
anteriorly at the articulation of... the fifth<br />
and sixth joints united, and which with the seventh<br />
form a very narrow and linear piece extending<br />
forwards to the posterior margin of the oral<br />
aperture." This is precisely the condition figured<br />
by Rathbun (1910, fig. 33) (Figure 35a herein) for<br />
T. orientalis. In Rathbun's species, the fused third<br />
and fourth somites taper abruptly, forming a<br />
truncated triangle anteriorly, and the slender terminal<br />
sections of the abdomen extend forward to<br />
a point between the maxillipeds. Inasmuch as all<br />
other hexapodid genera, with the exception of<br />
Stevea and possibly Hexaplax, have the third to<br />
fifth somites of the male abdomen fused, we<br />
believe that Rathbun's species, T. orientalis, must<br />
be assigned to Paeduma. This genus also now<br />
includes P. chuenense (Rathbun, 1909), based on a<br />
juvenile male, and a species from Japan, identified<br />
by earlier workers with T. orientalis, which we<br />
believe represents a new species, the description<br />
of which is in preparation. Paeduma cyclindraceum<br />
(Bell), with its unequal chelipeds, remains un-
NUMBER 306 175<br />
identifiable; the holotype, long thought to be lost,<br />
has been rediscovered in the Bell collection at the<br />
University Museum, Oxford, England, by Angelo<br />
A. DiMauro, Jr., of the University of Connecticut.<br />
His study of the Bell collection should allow us to<br />
be able to identify Bell's species.<br />
Paeduma resembles Thaumastoplax in having the<br />
propodus of the third maxillipeds dilated distally,<br />
in having the second walking legs strongest, and,<br />
in P. orientale at least, in lacking the oblique striae<br />
on the pterygostomian region. Paeduma differs<br />
from other hexapodid genera in having the third<br />
and fourth and the fifth and sixth somites of the<br />
abdomen fused in males.<br />
Genus Parahexapus Balss, 1922<br />
Parahexapus Balss, 1922:77 [type-species: Parahexapus africanus,<br />
Balss, 1922, by monotypy; gender: masculine].<br />
Hexapus {Parahexapus).—Monod, 1956:362, 370.<br />
DIAGNOSIS.—Eyes small, movable, cornea<br />
slightly broader than stalk. Third maxillipeds<br />
(Figure 36a) slender, pediform, ischium convex<br />
mesially, carpus, propodus and dactylus slender,<br />
cylindrical, dactylus longer than propodus; exopod<br />
with flagellum. Pterygostomian region with<br />
transverse granulate ridge. Walking legs short,<br />
merus of longest pair shorter than carapace. Sternal<br />
grooves absent. Apices of male pleopods concealed<br />
by abdomen, pleopods (Figure 36V) sinuous,<br />
slender, abruptly tapering distally to slender<br />
apex, directed anteriorly, with large spines<br />
near midlength directed proximally. Male abdomen<br />
(Figure 36b) with third to fifth somites fused,<br />
slender, subterminal and terminal somites long,<br />
latter rounded distally, not trilobed.<br />
INCLUDED SPECIES.—Parahexapus africanus Balss,<br />
1922.<br />
REMARKS.—Parahexapus shares slender third<br />
maxillipeds with Thaumastoplax, Spiroplax, and<br />
Paeduma; but in Parahexapus the propodus of the<br />
third maxilliped is very slender, not dilated distally.<br />
The first male pleopod (Figure 36V) of<br />
Parahexapus africanus differs from those of known<br />
species in the other three genera in having strong<br />
spines near midlength and in abruptly tapering<br />
distally.<br />
In P. africanus the interantennular septum is<br />
scarcely discernible in anterior view, the male<br />
abdomen extends anteriorly between the bases of<br />
the third maxillipeds, and the carapace is rather<br />
broad (width = 1.5-1.9 times length). Although<br />
the sutures in the abdomen of the single male<br />
available to us are very indistinct, it appears that<br />
the third to fifth somites are fused and the sixth<br />
somite is elongated, as figured by Crosnier (1967,<br />
fig. 33) (Figure 36A herein), as in Thaumastoplax<br />
anomalipes.<br />
Parahexapus africanus Balss, 1922<br />
FIGURE 36a-c<br />
Hexapus (Parahexapus) africanus.—Monod, 1956:370, figs. 494-<br />
496.—Rossignol, 1962:118.—Crosnier, 1967:337, figs. 30,<br />
33.<br />
Hexapus africanus.—Longhurst, 1958:88.—Gauld, 1960:71.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Congo: Baie de Pointe-Noire, A. Crosnier,<br />
26, 1$ (W).<br />
DESCRIPTION.—Monod, 1956:370.<br />
Figures: Monod, 1956, figs. 494-496; Crosnier,<br />
1967, figs. 30, 33.<br />
Male pleopod: Crosnier, 1967, fig. 30 (Congo).<br />
MEASUREMENTS.—The carapaces of our males<br />
measure 3.4 X 5.6 and 2.8 X 4.9 mm; that of the<br />
female measures 3.0 X 4.6 mm. Monod (1956)<br />
reported ovigerous females measuring 4X7 mm<br />
and 5 X 7.5 mm, and Crosnier (1967) reported<br />
males measuring 3.2 X 5 mm and 3.4 X 5.6 mm<br />
and a female as large as 3.4 X 5.6 mm. Balss<br />
(1922) reported that the female holotype measured<br />
4 X 7.5 mm.<br />
BIOLOGY.—Parahexapus africanus is a sublittoral<br />
species, which has been collected in depths between<br />
5 m and about 25 m (16-27 m; 22-25 m).<br />
Longhurst (1958) found the species off Sierra<br />
Leone in 16-27 m on shelly sand and sandy mud.<br />
Crosnier (1967) found it on sandy mud in 22-<br />
25 m off the mouth of the Congo River.
176 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 36.—Parahexapus africanus Balss: a, third maxilliped<br />
(from Monod, 1956, fig. 4%); b, abdomen of male; c, male<br />
pleopod (b,c, from Crosnier, 1967, figs. 30, 33). Pseudohexapus<br />
platydactylus Monod: d, abdomen of male; e, third maxilliped;<br />
/, male pleopod (d-f, from Monod, 1956, figs. 481, 484, 488).<br />
Ovigerous females have been collected in February<br />
(Monod, 1956).<br />
DISTRIBUTION.—<strong>West</strong> Africa, where it has been<br />
recorded from localities on the mainland between<br />
Sierra Leone and Cabinda, in depths between<br />
5 m and about 25 m (16-27 m is the deepest<br />
record). Records in the literature include the<br />
following:<br />
Sierra Leone: No specific locality, in 16-27 m (Longhurst,<br />
1958).<br />
Ghana: Off Accra, 5 m (Monod, 1956; Gauld, 1960).<br />
Congo: Baie de Pointe-Noire (Rossignol, 1962; Crosnier,<br />
1967). Off the mouth of the Congo River, 05°56'S, 12°O7'E,<br />
22-25 m (Crosnier, 1967).<br />
Cabinda: Landana, 6 m (Balss, 1922).<br />
Genus Pseudohexapus Monod, 1956<br />
Hexapus {Pseudohexapus) Monod, 1956:362, 365 [type-species:<br />
Hexapus (Pseudohexapus) platydactylus Monod, 1956, by<br />
monotypy; gender: masculine].<br />
DIAGNOSIS.—Eyes small, movable, cornea as<br />
broad as stalk. Third maxilliped (Figure 36e)<br />
broad, ischium short, mesial margin concave, projecting<br />
distally, carpus, propodus and dactylus<br />
slender, cylindrical, propodus longer than dactylus;<br />
exopod without flagellum. Pterygostomian<br />
region lacking oblique striae, with hairy ridge.<br />
Walking legs short, merus of longest pair (fourth)<br />
shorter than carapace. Dactyli of walking legs<br />
flattened dorsoventrally. Sternal grooves absent<br />
(?). Apices of male pleopods completely concealed<br />
by abdomen; pleopods (Figure 36/) stout, apices<br />
curved mesially, with terminal spinules and few<br />
subterminal setae. Male abdomen (Figure 36d)<br />
with 5 free somites, third to fifth fused, terminal<br />
somite elongate, rounded apically, not trilobed.<br />
INCLUDED SPECIES.—Pseudohexapus platydactylus<br />
Monod, 1956.<br />
REMARKS.—In Pseudohexapus the carapace is<br />
subquadrate (width = 1.2-1.3 times length), the<br />
pterygostomian region lacks a row of oblique<br />
striae, the propodus of the third maxilliped is<br />
slender and the exopod lacks a flagellum, the<br />
sternal grooves are absent, and the male pleopod<br />
is stout with its apex turned mesially. The flattened<br />
dactyli of the walking legs apparently are<br />
unique within the family.<br />
Pseudohexapus platydactylus Monod, 1956<br />
FIGURE 36d-f<br />
Hexapus (Pseudohexapus) platydactylus Monod, 1956:365, figs.<br />
478-493 [Senegal, Gambia, Ghana].—Gordon, 1971:107,<br />
fig. 2 [Ghana].<br />
Hexapus platydactylus.—Gauld, 1960:71 [Ghana].—Longhurst,<br />
1958:88 [Sierra Leone].<br />
DISTRIBUTION.—<strong>West</strong> Africa, from Senegal,<br />
Gambia, Sierra Leone, and Ghana, sublittoral, in<br />
5-15 m.<br />
Genus Spiroplax, new genus<br />
FIGURE 37<br />
TYPE-SPECIES.—Thaumastoplax spiralis Barnard,<br />
1950:301, fig. 56h-l, South Africa.
NUMBER 306<br />
FIGURE 37.—Spiroplax spiralis (Barnard): a, carapace; b, third<br />
maxilliped; c, sternum; d, abdomen of male; e, male pleopod.<br />
(All from Barnard, 1950, fig. 56h-l.)<br />
ETYMOLOGY.—From the Greek, speira (coil,<br />
twist), in combination with the noun plax (a<br />
plate), alluding to the form of the male pleopod;<br />
the gender of the name is feminine.<br />
DIAGNOSIS.—Eyes movable, cornea slightly<br />
broader than stalk. Third maxilliped (Figure 37£)<br />
slender, ischium with obtuse mesial projection,<br />
propodus shorter than dactylus, with mesial expansion<br />
distally; exopod flagellum not described.<br />
Pterygostomian region not described. Walking<br />
legs not described. Sternal grooves not described.<br />
Male pleopods (Figure 37*) apparently concealed<br />
under abdomen, strongly calcified, stout, spiral,<br />
apices setose. Male abdomen (Figure 37a 1 ) with<br />
third to fifth somites fused, very broad, terminal<br />
somite triangular, not trilobed.<br />
REMARKS.—The dilated propodus of the third<br />
maxillipeds suggests affinities with both Thaumastoplax<br />
and Paeduma, but in Spiroplax the carapace<br />
is much more oval (Figure 37a) (width = 1.3<br />
times length), the male abdomen is much<br />
broader, and the stout, spiral male pleopods are<br />
very different from the slender, almost filamentous<br />
male pleopods of representatives of the other<br />
two genera. Spiroplax includes only the typespecies.<br />
Genus Stevea, new genus<br />
FIGURE 38<br />
177<br />
TYPE-SPECIES.—Hexapus williamsi Glassell,<br />
1938:445, pi. 35, Guatemala (type locality) and<br />
Gulf of Tehuantepec, W coast of Mexico ($,<br />
9.4 X 14.4 mm, W).<br />
ETYMOLOGY.—The genus is named for Steve A.<br />
Glassell, the first carcinologist to record a hexapodid<br />
crab from the Americas; the gender of the<br />
generic name is feminine.<br />
DIAGNOSIS.—Carapace suboval (Figure 38a).<br />
Eyes small, movable, cornea subglobular, narrower<br />
than stalk. Third maxilliped (Figure 38c)<br />
broad, ischium flattened or slightly concave distally,<br />
carpus, propodus and dactylus slender, subcylindrical,<br />
dactylus longer than propodus; exopod<br />
with flagellum. Pterygostomian region with<br />
row of oblique striae. Walking legs short, merus<br />
of longest pair (third) shorter than carapace. Sternal<br />
grooves absent. Apices of male pleopods completely<br />
concealed by abdomen; structure of male<br />
pleopod not described. Male abdomen (Figure<br />
3&/) with three free somites, second through sixth<br />
fused, terminal somite triangular, not trilobed.<br />
REMARKS.—In the only species assigned to this<br />
genus, Stevea williamsi (Glasssell, 1938) (incorrectly<br />
spelled Hexapus williamsoni, in Stephensen,<br />
1945:182), there is a distinct median lobe on the<br />
front, visible in anterior view, the interantennular<br />
septum is poorly developed, the male abdomen<br />
extends beyond the bases of the third maxillipeds,<br />
and the carapace is relatively broad (width =1.5<br />
times length).<br />
The presence of only three free somites in the<br />
male abdomen will distinguish this genus from<br />
all others in the family.<br />
Genus Thaumastoplax Miers, 1881<br />
Thaumastoplax Miers, 188la:261 [type-species: Thaumastoplax<br />
anomalipes Miers, 1881, by monotypy; gender: feminine].<br />
DIAGNOSIS.—Eyes small, movable, cornea<br />
rounded, only slightly broader than stalk. Third<br />
maxilliped (Figure 39a) slender, ischium convex
178<br />
a<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 38.—Stevea williamsi (Glassell): a, dorsal view; b, front; c, third maxilliped; d, ventral<br />
view. (All from Glassell, 1938, pi. 35.)<br />
mesially, propodus strongly dilated distally, almost<br />
as broad as merus, longer than carpus or<br />
dactylus; exopod with flagellum. Pterygostomian<br />
region lacking row of oblique striae. Walking legs<br />
short, merus of longest pair (second) shorter than<br />
carapace. Sternal grooves absent. Male pleopods<br />
(Figure 39c) slender, sinuous, apices directed anteriorly,<br />
completely concealed under abdomen,<br />
armed with minute spinules. Male abdomen (Figure<br />
396) with third to fifth somites fused, terminal<br />
somite elongate, sinuous laterally, rounded apically,<br />
not trilobed.<br />
INCLUDED SPECIES.—Thaumastoplax anomalipes<br />
Miers, 1881, from <strong>West</strong> Africa. Two other species<br />
assigned to this genus, T. orientalis Rathbun, 1909,<br />
and T. chuenesis Rathbun, 1909, are here transferred<br />
to Paeduma (p. 173). The fourth species<br />
assigned to this genus, T. spiralis Barnard, 1950,<br />
is transferred to Spiroplax (p. 176).<br />
REMARKS.—Several features appear to be diagnostic<br />
for this genus, now known to include<br />
only a single species from tropical <strong>West</strong> Africa.<br />
The third maxilliped is slender and the propodus<br />
is strongly dilated distally, being almost as broad<br />
as the merus; the pterygostomian region is not<br />
ornamented with oblique striae; the second pair<br />
of walking legs is much the strongest of the three;<br />
the interantennular septum is very low, scarcely<br />
visible in frontal view; and the male pleopod is<br />
slender, sinuous, and directed anteriorly.<br />
Two other genera, Paeduma and Spiroplax, share<br />
the maxilliped with the distally dilated propodus,<br />
but both can be differentiated readily from Thaumastoplax.<br />
In Paeduma the third and fourth and<br />
fifth and sixth, rather than the third to fifth,<br />
abdominal somites in the male are fused, and the<br />
male pleopods are strongly recurved posteriorly
NUMBER 306 179<br />
or into a figure 8. In Spiroplax the male pleopods<br />
are much stouter and are twisted into a spiral.<br />
Thaumastoplax anomalipes Miers, 1881<br />
FIGURE 39<br />
Thaumastoplax anomalipes Miers, 188la:261, pi. 14: fig. 2.<br />
Hexapus (Thaumastoplax) anomalipes.—Monod, 1956:363, figs.<br />
471_477._Rossignol, 1962:118.—Gordon, 1971:108 [discussion].<br />
Hexapus anomalipes.—Longhurst, 1958:88.—Gauld, 1960:71.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Congo: Baie de Pointe-Noire, 7-10 m, A.<br />
Crosnier, 1
180<br />
posteriorly on carpus and propodus; inner surface<br />
of carpus and propodus also with some strong<br />
tubercles; dactylus slender, curved ventrajly,<br />
shorter than propodus; latter broad, length only<br />
1.6 times greatest depth; propodus shorter than<br />
carpus, merus about 0.8 times as long as carapace,<br />
0.75 times as long as carpus and propodus combined.<br />
Third walking leg slightly stouter than<br />
first; dactylus straight or slightly curved ventrally,<br />
more than half as long as propodus; latter 1.4<br />
times as long as broad, shorter than carpus; merus<br />
slender, 2.7 times as long as broad, 0.6 times as<br />
long as carapace, as long as carpus and propodus<br />
combined.<br />
Male abdomen narrow, composed of 5 free<br />
somites, third to fifth fused, about as long as sixth<br />
somite. Latter appearing slender, about 1.3 times<br />
as long as broad. Terminal somite as long as<br />
broad, 0.6 times as long as fused somites, lateral<br />
margins sinuous, apex rounded. Abdominal<br />
trench in male extending anteriorly to base of<br />
third maxillipeds. Abdomen of female with 7 free<br />
somites.<br />
Male pleopod as figured by Monod (1956, fig.<br />
476), slender, sinuous, elongate, almost naked,<br />
completely concealed under abdomen.<br />
Figures: Monod, 1956, figs. 471-477.<br />
Male Pleopod: Monod, 1956, fig 476 (Senegal).<br />
MEASUREMENTS.—Our male specimen measures<br />
3.9 X 5.7 mm, the female 4.8 X 8.0 mm.<br />
The largest specimen studied by Monod was an<br />
ovigerous female 11X19 mm.<br />
BIOLOGY.—Thaumastoplax anomalipes is a sublittoral<br />
species, living in depths between about 3 m<br />
and 34 m. The specimens recorded by Monod<br />
(1956) were taken on mud-sand in 19 m, on shelly<br />
sand in 8 m, and on muddy sand with gorgonians<br />
in 10 m. Longhurst (1958) found the species in<br />
estuaries as well as offshore in depths between 6<br />
and 34 m; in the estuary the species occurred on<br />
muddy sand and shelly mud, whereas offshore it<br />
was taken on shelly sand and shelly mud.<br />
DISTRIBUTION.— <strong>West</strong> coast of Africa, from a<br />
few localities between Senegal and Rio Muni;<br />
sublittoral, in depths between ca 3 m and 34 m.<br />
Monod (1956) recorded material from Senegal<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
and Guinea; since 1956 the species has been<br />
recorded from the following:<br />
Sierra Leone: No specific locality, in 6-34 m (Longhurst,<br />
1958).<br />
Ghana: Off Accra, ca. 3 m (Gauld, 1960).<br />
Congo: Baie de Pointe-Noire (Rossignol, 1962).<br />
Genus Tritoplax, new genus<br />
FIGURE 40<br />
TYPE-SPECIES.—Hexapus stebbingi Barnard,<br />
1947.<br />
ETYMOLOGY.—The feminine name is derived<br />
from the Greek, tritos (a third) and plax (a plate),<br />
alluding to the shape of the terminal somite of<br />
the male abdomen.<br />
DIAGNOSIS.—Eyes very small, cornea subglobular.<br />
Third maxilliped with broad ischium and<br />
merus, ischium concave proximally, straight distally,<br />
carpus, propodus, and dactylus slender,<br />
subcylindrical; exopod of flagellum not described.<br />
Pterygostomian region apparently lacking<br />
oblique striae. Walking legs short, merus of longest<br />
pair shorter than carapace. Sternal grooves<br />
restricted to trilobed anterior terminus of abdominal<br />
fossa, grooves not extending anterolaterally<br />
to bases of third maxillipeds. Male pleopods (Figure<br />
406) slender, sinuous, bent laterally, naked<br />
apex recurved anteriorly. Male abdomen (Figure<br />
40a) with third to fifth somites fused, sixth longitudinally<br />
divided in type-species, terminal<br />
somite distinctly trilobed, with angled lateral<br />
projections.<br />
INCLUDED SPECIES.—Hexapus stebbingi Barnard,<br />
1947 (see Barnard, 1950:299), from South Africa.<br />
An unnamed species identified by Stephensen<br />
(1945, fig. 53B; Figure AQd herein) as Hexapus<br />
sexpes, which also has a distinctly trilobed abdomen,<br />
although the sixth somite is not longitudinally<br />
divided, may also belong here.<br />
REMARKS.—The trilobed male abdomen and<br />
the longitudinally divided sixth abdominal somite<br />
in the male are both quite characteristic and<br />
will serve to distinguish this genus from others in<br />
the family. The trilobed shape of the anterior end
NUMBER 306 181<br />
of the abdominal fossa is similar to that found in<br />
species of Hexapinus, but the terminal somite of<br />
the male abdomen and the shape of the male<br />
pleopod are different in the two genera.<br />
FIGURE 40.—Tritoplax stebbingi (Barnard): a, abdomen of<br />
male; b, male pleopod; c, apex; d, abdomen and apex of<br />
pleopod of male (a-c, from Barnard, 1950, fig. 56*^/"; d, from<br />
Stephensen, 1945, fig. 53b).<br />
Family PINNOTHERIDAE de Haan, 1833<br />
PINNOTHERIDEA de Haan, 1833:2, 5 [corrected to Pinnotheridae<br />
by Bell, 1845:119].<br />
ASTHENOGNATHIDAE Stimpson, 1858b: 107.<br />
XENOPHTHALMIDAE Stimpson, 1858b: 107.<br />
DISSODACTYLIDAE Smith, 1870:172.<br />
PINNOTHERELINAE Alcock, 1900:294, 335.<br />
ANOMALOFRONTINAE Rathbun, 1931:84.<br />
EASTERN ATLANTIC GENERA.—Two, Asthenognathus<br />
and Pinnotheres, both represented by species<br />
living off tropical <strong>West</strong> Africa.<br />
EASTERN ATLANTIC SPECIES.—Fourteen, ten of<br />
vvhich are named; some are of doubtful taxonomic<br />
standing.<br />
Schmitt, McCain, and Davidson (1973) listed<br />
2 genera and 10 species of Pinnotheridae from<br />
<strong>West</strong> Africa: Asthenognathus atlanticus Monod, Pinnotheres<br />
pinnotheres (Linnaeus), Pinnotheres pisum<br />
(Linnaeus), Pinnotheres leloeuffi Crosnier, Pinnotheres<br />
mccainae Schmitt, Pinnotheres sp. A, B, C, D, of<br />
Monod (1956) and Pinnotheres sp. of Guinot and<br />
Ribeiro (1962). Pinnotheres sp. C of Monod is<br />
described herein as a new species, Pinnotheres conicola.<br />
Pinnotheres tellinae, which also is described<br />
here as new, includes in part Monod's species A.<br />
The only <strong>West</strong> <strong>African</strong> records of Pinnotheres<br />
pinnotheres are those by Balss (1922). Unfortunately,<br />
Balss did not give any details of his material<br />
that would confirm the correctness of his<br />
identification, and it seems likely that his specimens,<br />
which were collected in Cameroon and<br />
Gabon, actually belong to one or more of the true<br />
<strong>West</strong> <strong>African</strong> species that were described since<br />
1922. Until Balss' material can be reexamined,<br />
Pinnotheres pinnotheres has to be included in the list<br />
of <strong>West</strong> <strong>African</strong> species, be it with great reserve.<br />
Also the otherwise European species, P. pisum<br />
(Linnaeus), has been recorded from Morocco and<br />
Mauritania. At present, therefore, six named species<br />
of Pinnotheres are known from <strong>West</strong> Africa (P.<br />
leloeuffi, P. mccainae, P. conicola, P. tellinae, P. pisum,<br />
P. pinnotheres; the last doubtfully), and four unnamed<br />
species (species A (part), B, and D of<br />
Monod, 1956, and Pinnotheres sp. of Guinot and<br />
Ribeiro, 1962).<br />
The pinnotherids were not represented in the<br />
Pillsbury collections. The following species of pinnotherids<br />
are extralimital:<br />
Pinnotheres ascidicola Hesse, 1872. Atlantic coast<br />
of France; in ascidians.<br />
Pinnotheres marioni Gourret, 1887. Mediterranean<br />
(Bay of Marseilles); in ascidians.<br />
Pinnotheres pectunculi Hesse, 1872. Atlantic coast<br />
of Brittany (France); in Glycimeris (Lamellibranchia).<br />
Subfamily ASTHENOGNATHINAE Stimpson,<br />
1858<br />
Genus Asthenognathus Stimpson, 1858<br />
Asthenognathus Stimpson, 1858b: 107 [p. 53 in separate; typespecies:<br />
Asthenognathus inaequipes Stimpson, 1858, by<br />
monotypy; gender: masculine; name 287 on Official List].<br />
Asthenognathus atlanticus Monod, 1933<br />
Asthenognathus atlanticus.—Monod, 1956:383, figs. 541-545<br />
[Mauritania, Senegal, Ghana].—Gauld, 1960:71
182 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
[Ghana].—Guinot and Ribeiro, 1962:65 [Angola].—Rossignol,<br />
1962:119 [Congo].—Monod, 1963: fig. 39 [no material].—Zariquiey<br />
Alvarez, 1968:410, fig. 137 [Spain;<br />
references].—Schmitt, McCain, and Davidson, 1973:128<br />
[synonymy].—Noel and Amouroux, 1977:135 [Mediterranean].<br />
DISTRIBUTION.—Eastern Atlantic, from Normandy,<br />
NW France to Angola and Mediterranean;<br />
on echinoids and commensal with polychaetes;<br />
sublittoral, in depths between 8 and 70<br />
m.<br />
Subfamily PINNOTHERINAE de Haan, 1833<br />
Genus Pinnotheres Bosc, 1802<br />
Pinnotheres Bosc, 1802:59, 239 [type-species: Cancerpisum Linnaeus,<br />
1767, by selection by Latreille, 1810:422; gender:<br />
masculine; name 352 on Official List].<br />
Holothunophilus Nauck, 1880:66 [type-species: Holothuriophilus<br />
trapeziformis Nauck, 1880, by original designation; gender;<br />
masculine; name 319 on Official List].<br />
Arcotheres Burger, 1895:361 [type-species: Pinnotherespalaensis<br />
Burger, 1895, by selection by Rathbun, 1918:62; gender:<br />
masculine].<br />
Zaops Rathbun, 1900b:588, 590 [type-species: Pinnotheres depressum<br />
Say, 1817, a subjective junior synonym of Pinnotheres<br />
ostreum Say, 1817, by original designation and monotypy;<br />
gender: masculine].<br />
Pinnozoea Aikawa, 1933:130, 246 [type-species: Cancer pisum<br />
Linnaeus, 1767, by selection by Schmitt, McCain, and<br />
Davidson, 1973:37; gender: feminine].<br />
Pinnotheres conicola, new species<br />
FIGURES 41, 42<br />
Pinnoteres sp. C—Monod, 1956:380, figs. 508, 509, 526-538<br />
[not figs. 524, 525 = Pinnotheres sp. B].—Longhurst, 1958:<br />
88.<br />
Pinnotheres sp. C.—Silas and Alagarswami, 1967:1214 [listed].—Schmitt,<br />
McCain, and Davidson, 1973:92 [synonymy].<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Cameroon: Kribi, in large Conus obtained<br />
by native fishermen who fished with a canoe ("pirogue")<br />
near the shore, 10 Mar 1964, B. de Wilde-Duyfjes, holotype,<br />
Id (L).<br />
DESCRIPTION.—Carapace (Figure 41) firm, almost<br />
circular in outline, about 1.1 to 1.3 times<br />
wider than long. Surface of carapace naked,<br />
smooth, except for coarsely pitted branchial regions,<br />
slightly convex dorsally, falling off more<br />
steeply laterally; anterolateral margin broadly<br />
and evenly rounded. Lateral surface with long,<br />
soft hairs, which, with similar hairs on pereiopods,<br />
enclose naked upper surface in hirsute ring. Front<br />
almost square, slightly produced in middle, in<br />
dorsal view anterior margin scarcely produced<br />
beyond circular outline of carapace. Eyes small,<br />
short, with reduced cornea; orbit subcircular.<br />
Third maxilliped (Figure 42a) placed<br />
obliquely, almost transversely. Merus operculiform,<br />
very wide, 2/3 as wide as long, greatest<br />
width in anterior half, anterointernal angle<br />
evenly rounded. Carpus short. Propodus oval.<br />
Dactylus very short, narrow, inserted on posterior<br />
half of lower margin of propodus, falling short of<br />
apex of propodus. Carpus and propodus together<br />
less than half as long as merus.<br />
Chelipeds (Figure 42b-d) well developed, fingers<br />
about 6/7 as long as palm. Dactylus with<br />
single large tooth on basal half of cutting edge,<br />
distal half with some small granular denticles.<br />
Cutting edge of fixed finger also with large tooth,<br />
placed before that of dactylus, continued posteriorly<br />
as crenulated ridge. Base of fingers, upper<br />
surface of dactylus, lower surface of fixed finger,<br />
and cutting edges densely covered with soft, slender,<br />
plumose hairs. Palm with upper half of outer<br />
surface pilose, lower half naked; inner surface<br />
completely covered with long, smooth hairs except<br />
for naked spot anteriorly near base of fingers.<br />
Carpus short, rounded, upper surface naked, remainder<br />
covered by long hairs. Merus with upper<br />
and outer part naked, remainder clothed with<br />
long hairs. Ischium with distinct lobe on anterointernal<br />
angle of upper margin.<br />
Walking legs (Figure 42e-h) subsimilar, all<br />
rather robust, none longer or slenderer than others.<br />
Dactylus of each walking leg similar to that<br />
of other legs, about 2/3 as long as propodus<br />
(measured dorsally), short, simple, terminating in
NUMBER 306<br />
narrow, slightly curved tip. Propodus, carpus,<br />
and merus each high, with thick coat of hairs<br />
dorsally and ventrally but with lateral surfaces<br />
bare. Propodus about twice as long as high, merus<br />
somewhat more slender.<br />
Male abdomen (Figure 42*) elongate, with<br />
fifth and sixth somites fused, line of separation<br />
scarcely distinct but present; Monod (1956:378,<br />
fig. 508) showed 7 free somites on male abdomen.<br />
Telson triangularly rounded.<br />
Male Pleopod: Monod, 1956:382, figs. 537, 538.<br />
MEASUREMENTS.—The holotype has a carapace<br />
length of 10 mm and a carapace width of 11.5<br />
mm. The two male specimens reported upon by<br />
Monod had cl 9 and 9.5 mm, and cb 11 and 10<br />
mm, respectively; his ovigerous female had cl<br />
15 mm, cb 18 mm, and a juvenile female cl 8 and<br />
cb 9 mm.<br />
REMARKS.—In Monod's (1956) fundamental<br />
work on the crabs of <strong>West</strong> Africa, his figures 508<br />
and 509 are labeled Pinnotheres sp. B, while figures<br />
524 and 525 are labeled Pinnotheres sp. C. However,<br />
a comparison of these figures with the measurements<br />
of these specimens given in the text<br />
and with the details shown in his figures 510 to<br />
523 and 526 to 538, indicate that the explanation<br />
said to be of his figures 524 and 525 actually is<br />
that of his figures 508 and 509 and vice-versa.<br />
Judging by the enlargement indicated, the male<br />
and female of Monod's figures 508 and 509 have<br />
FIGURE 41.—Pinnotheres conicola, new species: a, male; b, female.<br />
(From Monod, 1956, figs. 524, 525.)<br />
183<br />
cb 11 and 14 and cl 9 and 11 mm, respectively,<br />
which agrees with the measurements given for<br />
Pinnotheres sp. C from Conakry; the male and<br />
female of Monod's figures 524 and 525 have,<br />
respectively, cb 20 and 20 mm, and cl 16 and<br />
18 mm, which agree well with the specimen of<br />
Pinnotheres sp. B from Goree. There can, therefore,<br />
be little doubt that Monod's figures 508, 509,<br />
526-538 show Pinnotheres sp. C, and figures 510-<br />
525 show Pinnotheres sp. B. Monod's (1956:382,<br />
figs. 539, 540) Pinnotheres sp. D shows a great<br />
resemblance to the present species, and might<br />
well be specifically the same. As Monod did not<br />
provide a description of his species D, this problem<br />
can only be solved by an examination of the<br />
material in question.<br />
The short stubby legs fringed with heavy rows<br />
of setae distinguish this species immediately from<br />
most other named species of this genus known<br />
from the eastern Atlantic: Pinnotheres pinnotheres<br />
(Linnaeus), P. pisum (Linnaeus), and P. mccainae<br />
Schmitt. From P. leloeuffi Crosnier it can be distinguished<br />
by the less wide carapace, by the<br />
dorsal and ventral fringes of hair on all the segments<br />
of the legs except the dactylus, the wider<br />
merus of the third maxilliped and the shape of<br />
the male abdomen. Actually it seems closest to<br />
Monod's (1956) Pinnotheres sp. D with which it<br />
may prove to be identical. In its pubescence the<br />
new species shows some resemblance to Pinnotheres
184<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 42.—Pinnotheres conicola, new species: a, third maxilliped; b-d, chelipeds; e-h, second to<br />
fifth pereiopods; i, abdomen of holotype, male, cl. 10.0 mm, Kribi (a-h from Monod, 1956, figs.<br />
526-532).<br />
barbatus Desbonne from the <strong>West</strong> Indies, and also<br />
resembles that species in the robust walking legs,<br />
which are of similar size and shape and which<br />
have all dactyli short and falcate. It differs<br />
strongly, however, in the shape of the third maxilliped<br />
and the male abdomen. In P. barbatus the<br />
dactylus of the third maxilliped is implanted at<br />
the tip of the propodus, and the male abdomen<br />
has all somites free. Furthermore the carapace of<br />
the American species is softer and shows a large<br />
pit on either side of the middle (see Rathbun,<br />
1918:88, pi. 19: figs. 8-11).<br />
TYPE-LOCALITY.—Kribi, Cameroon.<br />
DISPOSITION OF TYPE.—The holotype (Crust.<br />
D. 28792) is in the collection of the Rijksmuseum<br />
van Natuurlijke Historie, Leiden.<br />
The specimens reported by Monod (1956) from<br />
Conakry are probably in the collection of the<br />
Institut Fondamental d'Afrique Noire, Dakar,<br />
Senegal. The specimen reported by Longhurst<br />
(1958) may be in the collection of the British<br />
Museum (Natural History), London.<br />
ETYMOLOGY.—The specific epithet is derived<br />
from the Latin generic name Conus, referring to<br />
the association of the species with a species of<br />
that genus.<br />
BIOLOGY.—All specimens of this species known<br />
so far were found to live commensally with gastropod<br />
mollusks of the genus Conns; only the host<br />
of the Sierra Leone specimens is identified to<br />
species, viz. Conus papillionaceus Hwass. The Cameroon<br />
specimen was said to be from a large<br />
Conus. The depths at which the specimens were<br />
found is not accurately known: Longhurst (1958)<br />
gave the depth at which his specimens were taken<br />
as "shallow" and as "0-40 m." The Cameroon<br />
specimen evidently also was taken in shallow<br />
water as it was obtained from native fishermen<br />
fishing near the coast in a canoe.<br />
One of the specimens reported from Sierra<br />
Leone was an ovigerous female; the date of collection<br />
was not recorded.<br />
DISTRIBUTION.—<strong>West</strong> Africa, where it has been<br />
recorded from Guinea, Sierra Leone, and now
NUMBER 306 185<br />
Cameroon; our specimen represents a considerable<br />
range extension to the south. Records in the<br />
literature include the following:<br />
Guinea: Conakry (Monod, 1956).<br />
Sierra Leone: No specific locality (Monod, 1956;<br />
Longhurst, 1958).<br />
Pinnotheres leloeuffl Crosnier, 1969<br />
Pinnotheres leloeuffi Crosnier, 1969:531, figs. 1-10, 17 [Ivory<br />
Coast].—Schmitt, McCain, and Davidson, 1973:52 [synonymy].<br />
DISTRIBUTION.—<strong>West</strong> Africa, from Vridi, Ivory<br />
Coast, in 20 m. Host unknown.<br />
Pinnotheres mccainae Schmitt, 1973<br />
FIGURE 43<br />
Pinnotheres rouxi Rossignol, 1957:84, fig. 4.—Silas and Alagarswami,<br />
1967:1208 [listed].—Schmitt, McCain, and<br />
Davidson, 1973:2, 57 [synonymy]. [Preoccupied by Pinnotheres<br />
rouxi H. Milne Edwards, 1837.]<br />
Pinnoteres rouxi.—Rossignol, 1962:118.<br />
Pinnotheres mccainae Schmitt, in Schmitt, McCain, and Davidson,<br />
1973:2,5, 11,57.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Congo: Pointe-Noire, from mantle cavity<br />
of Donax rugosus Linnaeus, 25 Apr 1955, A. G. Humes, 10$<br />
ov (L, W).<br />
DESCRIPTION.—Carapace quadrangular anteriorly,<br />
semicircular posteriorly. Anterolateral angles<br />
rounded, about as wide as front, each occupying<br />
about 1/3 of length of anterior margin of<br />
carapace. Lateral margins of carapace diverging<br />
posteriorly toward posterolateral angles. Carapace<br />
with surface naked, somewhat convex, integument<br />
thin. Front slightly convex in dorsal<br />
view, extending anteriorly to level of anterolateral<br />
angles; front bent strongly downward, coarsely<br />
pitted, at each anterolateral angle.<br />
Third maxilliped (Figure 43a,b) placed very<br />
obliquely, almost transversely. Dactylus narrow,<br />
elongated, articulated just before middle of lower<br />
margin of propodus, extending almost to end of<br />
propodus. Merus about twice as long as palp and<br />
about twice as long as broad. Exopod almost<br />
entirely covered by merus, basal segment rather<br />
broad, more than twice as long as wide. Distal 2<br />
segments of exopod narrow, together somewhat<br />
more than half as long as first segment.<br />
Cheliped of female (Figure 43c) elongate, chela<br />
more than 3 times as long as high. Fingers about<br />
3/4 as long as palm, tips sharp, curved. Basal<br />
part of cutting edges of both fingers with distinct<br />
tooth, that on fixed finger proximal to that on<br />
dactylus. Palm twice as long as high. Chela with<br />
fringe of long hairs on lower part of inner surface<br />
extending over all of fixed finger and distal part<br />
of palm. Dactylus with hairs scattered over surface.<br />
Row of hairs present on upper inner surface<br />
of carpus, merus with scattered hairs on inner<br />
surface.<br />
Pereiopods (Figure 43d-g) similar, of same<br />
length, on both sides of body. Third leg longest,<br />
about twice as long as fifth, \ x h times as long as<br />
second or fourth. Dactyli of second and fifth legs<br />
shortest, that of fourth slightly longer, that of<br />
third about \ l h. times length of shortest dactyli.<br />
In all legs, dactylus terminating in hook-shaped<br />
claw. Dactylus of second leg slightly more than<br />
1/3 as long as propodus, half as long as carpus;<br />
merus twice as long as carpus. Dactylus of third<br />
leg 2/5 as long as propodus, half as long as carpus;<br />
merus less than twice as long as carpus. Dactylus<br />
of fourth leg 2/5 as long as propodus, 2/3 as long<br />
as carpus, 1/3 as long as merus. Fifth pereiopod<br />
shortest, stoutest; dactylus 2/5 as long as propodus,<br />
2/3 as long as carpus, latter 3/4 as long as<br />
merus. Pereiopods not markedly hairy, only some<br />
scattered hairs present, densest proximally.<br />
Abdomen of ovigerous females distinctly wider<br />
than carapace.<br />
Eggs numerous, small, spherical, diameter<br />
about 0.2 mm.<br />
Figures: Rossignol (1957) provided figures of<br />
the general shape of an ovigerous female, a rather<br />
poor figure of the third maxilliped, and one of<br />
the chela. Some additional sketches are provided<br />
here (Figure 43).<br />
Male Pleopod: The shape of the male pleopod is<br />
unknown. The single type-specimen is a female,<br />
and our material consists entirely of females. Rossignol<br />
(1962) evidently had males at his disposal,<br />
as he reported that many of the Donax specimens
186<br />
a<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 43.—Pinnotheres mccainae Schmitt, ovigerous female, cl 5.3 mm, Pointe-Noire: a,b, third<br />
maxilliped; c, chela; d, second pereiopod; e, third pereipod; /, fourth pereiopod; g, fifth<br />
pereiopod.<br />
collected by him were "habitee par un couple de<br />
cette espece"; however, he gave no details of the<br />
male specimens.<br />
MEASUREMENTS.—Our ovigerous females have<br />
carapace lengths varying from 4 to 6.5 mm, and<br />
carapace widths from 5.5 to 7 mm. The holotype,<br />
also an ovigerous female had the carapace 6 mm<br />
long and 7 mm wide. The diameter of the eggs is<br />
about 0.2 mm.<br />
REMARKS.—The species was first described under<br />
the name Pinnotheres rouxi by Rossignol (1957)<br />
based on a single specimen. Silas and Alagarswami<br />
(1967) correctly pointed out that the name<br />
Pinnotheres rouxi had already been used by H.<br />
Milne Edwards (1837) for a different (Indo-<strong>West</strong><br />
Pacific) species and that therefore the <strong>West</strong> <strong>African</strong><br />
species needed a new name. It was Schmitt<br />
(in Schmitt, McCain, and Davidson, 1973), who<br />
provided this new name, Pinnotheres mccainae.<br />
BIOLOGY.—All the specimens so far mentioned<br />
in the literature (Rossignol, 1957, 1962) were<br />
found to be commensals of Donax sp. The present<br />
specimens were found in the mantle cavity of<br />
Donax rugosus Linnaeus. Rossignol (1962) indicated<br />
that a large proportion of the Donax found<br />
at Pointe-Noire was infested with this commensal;
NUMBER 306 187<br />
the bivalves were taken sublittorally in the sand<br />
at the lower limit of the lowest low tides.<br />
Rossignol's (1957) ovigerous female was collected<br />
in November, ours in April.<br />
DISTRIBUTION.—So far the species is only known<br />
from the area of Pointe-Noire, Congo. The holotype<br />
was reported from the beach at Pointe-Noire<br />
(Rossignol, 1957); additional material was mentioned<br />
by Rossignol (1962) from the Baie de<br />
Pointe-Noire. Also the present material is from<br />
that locality.<br />
Pinnotheres pinnotheres (Linnaeus, 1758)<br />
Pinnoteres pinnoteres.—Balss, 1922:79 [Cameroon, Gabon].<br />
Pinnoteres pinnotheres.—Monod, 1956:376 [references].<br />
Pinnotheres pinotheres.—Zariquiey Alvarez, 1968:409, fig.<br />
136b,f [Spain; references].<br />
Pinnotheres pinnotheres.—Schmitt, McCain, and Davidson,<br />
1973:68 [synonymy].<br />
SYNONYMS.—Pinnotheres veterum Bosc, 1802; Pinnotheres<br />
pinnae Leach, 1814; Pinnotheres montagui<br />
Leach, 1815; Pinnotheres pinnophylax H. Milne Edwards,<br />
1853.<br />
DISTRIBUTION.—North Sea to Mediterranean;<br />
possibly Gulf of Guinea; commensal in bivalves<br />
and ascidians.<br />
Pinnotheres pisum (Linnaeus, 1767)<br />
Cancer Pisum Linnaeus, 1767:1039.<br />
Pinnotheres pisum.—Capart, 1951:175 [Mauritania].—Forest<br />
and Games, 1960:353 [Morocco].—Zariquiey Alvarez,<br />
1968:408, figs. 7b, 14f, 135c,d, 136a,c-e [Spain; references].—Christiansen,<br />
1969:88, figs. 36, 37, map 30 [Scandinavia].—Schmitt,<br />
McCain, and Davidson, 1973:72 [synonymy].<br />
Pinnoteres pisum.—Monod, 1956:375 [references].<br />
SYNONYMS.—Cancer nutrix Scopoli, 1763; Cancer<br />
mytilorum albus Herbst, 1783; Cancer mytilorum juscus<br />
Herbst, 1783; Cancer varians Olivier, 1791; Pinnotheres<br />
mytili Leach, 1814; Pinnotheres modioli Leach,<br />
1814; Pinnotheres cranchii Leach, 1814; Pinnotheres<br />
latreillii Leach, 1817; Cancer eubolinus Nardo, 1847.<br />
DISTRIBUTION.—Eastern Atlantic, from Norway<br />
southward to Port Etienne, Mauritania; Mediter-<br />
ranean; commensal in bivalves and possibly ascidians;<br />
intertidal to about 45 m.<br />
Pinnotheres tellinae, new species<br />
FIGURES 44, 45<br />
Pinnoteres sp. A.—Monod, 1956:376 [part; not figs. 502-<br />
507].—Longhurst, 1958:88.<br />
Pinnotheres sp. A.—Silas and Alagarswami, 1967:1213<br />
[part].—Schmitt, McCain, and Davidson, 1973:91 [synonymy;<br />
part].<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Sierra Leone: Kissy, near Freetown, from<br />
mantle cavity of Tellina nymphalis Lamarck, 29 Nov 1954, A.<br />
G. Humes, 16, 109 (holotype; 2 ov) (L, W). Bullom Shore,<br />
opposite Freetown, from mantle cavity of Tellina nymphalis<br />
Lamarck, 12 Nov 1954, A. G. Humes, 11
188<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 44.—Pinnotheres tellinae, new species, paratypes, Kissy: a, adult female, cl 4.7 mm;<br />
b, adult male, cl 3.0 mm.<br />
tooth of fixed finger placed before that of dactylus.<br />
Remainder of cutting edges of chela with<br />
widely spaced tubercles; subdistally, cutting<br />
edges with short ridge, upper margin minutely<br />
tuberculate. Upper margin of chela with fringe<br />
of long hairs, latter also present on lower part of<br />
inner surface of palm and on fingers; shorter hairs<br />
present on remainder of chela. No tubercles present<br />
on either surface of palm. Carpus short, conical,<br />
slightly shorter than merus. Carpus and<br />
merus both with long hairs on dorsal and lower<br />
inner surface.<br />
Pereiopods of right side of equal size and shape<br />
as those on left side. First 3 walking legs (pereiopods<br />
2-4) (Figures 45
NUMBER 306 189<br />
a<br />
FIGURE 45.—Pinnotheres tellinae, new species, paratypes, Kissy. Female, cl 5.9 mm: a, front; b,<br />
third maxilliped; c, cheliped; d, second pereiopod; e, third pereiopod; /, fourth pereiopod; g,<br />
fifth pereiopod. Male, cl 2.5 mm: h, chela; i, second pereiopod; j, third pereiopod; k, fifth<br />
pereiopod; /, abdomen; m, gonopod.
190<br />
his single ovigerous female as 5.5 mm. The diameter<br />
of the eggs is 0.2 to 0.3 mm.<br />
REMARKS.—The species is characterized by the<br />
almost equal walking legs that have the dactylus<br />
of equal length, by the third maxilliped that has<br />
the dactylus implanted on the lower margin of<br />
the propodus, reaching slightly beyond the top of<br />
the propodus, by the hairiness of the legs, and by<br />
the naked carapace. It resembles Pinnotheres pholadis<br />
de Haan from Japan, which, however, has<br />
more slender and hairless legs. The hairy legs of<br />
the female P. tellinae distinguish it from the <strong>West</strong><br />
<strong>African</strong> species P. mccainae and P. pinnotheres; in P.<br />
leloeuffi the carapace is much wider and the shape<br />
of the third maxilliped and that of the male<br />
abdomen are quite different.<br />
Monod (1956:376) referred an ovigerous female<br />
of a Pinnotheres found in Tellina sp. in Sierra Leone,<br />
to his Pinnotheres sp. A. The other specimens referred<br />
by Monod to his species A were commensals<br />
of Ostrea tulipa Lamarck (= Ostrea gasar) from<br />
Senegal. Monod's (1956, figs. 502-507) illustrations<br />
are made after the Ostrea commensals, and<br />
show a species that is clearly different from P.<br />
tellinae. In the Ostrea commensals the carapace is<br />
wider in the females (length 9 mm, width 10 mm)<br />
and narrower in the males (length and width<br />
2 mm), the pereiopods are more slender and<br />
without long hairs, the shapes of the dactylus and<br />
propodus of the third maxilliped are different,<br />
and the size of the females is much larger. Although<br />
it is highly likely that Monod's specimen<br />
from Sierra Leone belongs to P. tellinae, as it was<br />
taken at the type-locality and from the type-host<br />
of that species, it is certain that the specimens<br />
from the host Ostrea tulipa belong to a different<br />
species. Monod (1956:376) himself remarked that<br />
"la conspecificite de cet exemplaire [i.e., the female<br />
from Sierra Leone] avec les precedents [i.e.,<br />
those from Senegal] n'est pas certaine."<br />
TYPE-LOCALITY.—Kissy, near Freetown, Sierra<br />
Leone.<br />
DISPOSITION OF TYPES.—The holotype (USNM<br />
169498) is one of the females from Kissy; all of<br />
the other specimens are paratypes. The holotype<br />
is deposited in the National Museum of Natural<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
History, <strong>Smithsonian</strong> <strong>Institution</strong>, Washington,<br />
D.C. The paratypes are in the same museum and<br />
duplicates are placed in the Rijksmuseum van<br />
Natuurlijke Histoire, Leiden. The specimen mentioned<br />
by Monod (1956:376) and Longhurst<br />
(1958:88), also a paratype, most likely is in the<br />
British Museum (Natural History).<br />
ETYMOLOGY.—The specific epithet is from the<br />
generic name Tellina, referring to the association<br />
of this species with a species of that genus.<br />
BIOLOGY.—Monod (1956) reported the species<br />
from Tellina sp. His specimen was collected by A.<br />
Longhurst in Sierra Leone. Longhurst (1958:88)<br />
himself gave more detailed information of what<br />
evidently is the same specimen, indicating that<br />
his material was found in the mantle cavity of<br />
Tellina nymphalis Lamarck, taken in an estuarine<br />
habitat. The specimens dealt with in the present<br />
paper were obtained by Dr. A. G. Humes from<br />
the mantle cavity of the same host.<br />
The date of collecting of Monod's and Longhurst's<br />
ovigerous female is not known. Our ovigerous<br />
females were collected in November.<br />
DISTRIBUTION.—So far the species is only known<br />
from Sierra Leone (Monod, 1956; Longhurst,<br />
1958).<br />
Pinnotheres sp. A<br />
Pinnoteres sp. A.—Monod, 1956:376, figs. 502-507 [part;<br />
Senegal].<br />
Pinnotheres sp. A.—Silas and Alagarswami, 1967:1213<br />
[listed].—Schmitt, McCain, and Davidson, 1973:91<br />
[synonymy].<br />
DISTRIBUTION.—Senegal. Commensal of Ostraea<br />
gasar (Adanson) (= Ostrea tulipa Lamarck)<br />
(Monod, 1956).<br />
Pinnotheres sp. B<br />
Pinnoteres sp. B.—Monod, 1956:378, figs. 510-525 [Senegal].<br />
Pinnotheres sp. B.—Silas and Alagarswami, 1967:1213<br />
[listed].—Schmitt, McCain and Davidson, 1973:91 [synonymy].<br />
DISTRIBUTION.—Senegal, in Panopea aldrovandi<br />
Menard; sublittoral, to at least 5 m.
NUMBER 306 191<br />
Pinnotheres sp. D<br />
Pinnoteres sp. D — Monod, 1956:382, figs. 539, 540<br />
[Congo].—Rossignol, 1962:118 [Congo].<br />
Pinnotheres sp. D.—Silas and Alagarswami, 1967:1214<br />
[listed].—Schmitt, McCain and Davidson, 1973:92 [synonymy].<br />
DISTRIBUTION.—Pointe-Noire, Congo, host and<br />
depth range unknown.<br />
Pinnotheres sp.<br />
Pinnotheres sp.—Guinot and Ribeiro, 1962:64, figs. 30-33<br />
[Angola].—Schmitt, McCain and Davidson, 1973:94 [synonymy].<br />
DISTRIBUTION.—Baia de Santa Marta, Angola,<br />
among mussels on rocks.<br />
Family RETROPLUMIDAE Gill, 1894<br />
RETROPLUMIDAE Gill, 1894:1043.<br />
PTENOPLACIDAE Alcock, 1899c: 78.<br />
REMARKS.—This family, which contains a single<br />
genus, Retropluma Gill, 1894, is not represented<br />
in the eastern Atlantic.<br />
Family MICTYRIDAE Dana, 1851<br />
MYCTIRIDAE Dana, 1851e:247 [corrected to Mictyridae by<br />
Alcock, 1900:383].<br />
REMARKS.—This family, which contains a single<br />
genus, Mictyris Latreille, 1806, is not represented<br />
in the eastern Atlantic.<br />
Family PALICIDAE Rathbun, 1898<br />
CYMOPOLIDAE Dana, 1854:9 [corrected to Cymopoliidae by<br />
Faxon, 1895:38; name 406 on Official Index, under Cymopoliidae<br />
Faxon, 1895].<br />
PALICIDAE Rathbun, 1898:280 [name 376 on Official List].<br />
PALICAE Bouvier, 1898:56, 58.<br />
EASTERN ATLANTIC GENERA.—One, Palicus, occurring<br />
off <strong>West</strong> Africa and in the Mediterranean<br />
Sea.<br />
EASTERN ATLANTIC SPECIES.—One, Palicus caronii<br />
(Roux, 1830), represented by one lot in the<br />
Pillsbury collections. It was recorded by Monod<br />
(1956) as Cymopolia caroni.<br />
Genus Palicus Philippi, 1838<br />
Cymopolia P. Roux, 1830, pi. 21 [invalid junior homonym of<br />
Cymopolia Lamouroux, 1816 (alga); type-species: Cymopolia<br />
caronii Roux, 1830, by monotypy; gender: feminine; name<br />
1718 on Official Index].<br />
Palicus Philippi, 1838:11 [type-species: Palicus granulatus Philippi,<br />
1838, a subjective junior synonym ofCymopolia caronii<br />
Roux, 1830, by monotypy; gender: masculine; name 1640<br />
on Official List].<br />
REMARKS.—Holthuis and Gottlieb (1958:104)<br />
explained why Palicus must be used in place of<br />
the older Cymopolia.<br />
* Palicus caronii (P. Roux, 1830)<br />
Cymopolia caroni.—Monod, 1956:387, figs. 546-551.<br />
Palicus caroni.—Forest and Guinot, 1966:88.<br />
Palicus caronii.—Zariquiey Alvarez, 1968:411, fig. 135e<br />
[Spain; references].—Tiirkay, 1976b:61 [listed], 71.<br />
SYNONYM.—Palicus granulatus Philippi, 1838.<br />
MATERIAL EXAMINED.—Pillsbury material: Ghana:<br />
Sta. 24, 35-37 m, dark red bryozoans, 26, 19 ov (L, W).<br />
Other Material: Canary Islands: No specific locality, 30 m<br />
(?), Talisman, 16 (W).<br />
Cape Verde Islands: Ilheu Branco, Talisman, 26, 1? ov<br />
(W).<br />
Senegal: Near Bane du Seminole, Baie de Goree, 38 m, I.<br />
Marche-Marchad, 18 Feb 1954, 26 (W).<br />
DESCRIPTION.—Bouvier, 1940:5; A. Milne Edwards<br />
and Bouvier, 1902:46-47 (comparison off.<br />
affinis A. Milne Edwards and Bouvier and P.<br />
caronii).<br />
Figures: Monod, 1956, figs. 546-551.<br />
Male Pleopod: Monod, 1956, figs. 549-551 (Senegal).<br />
MEASUREMENTS.—Our males have carapace<br />
widths of 7.6 to 10 mm; the ovigerous females<br />
have carapace widths of 9.3 and 11 mm.<br />
BIOLOGY.—Palicus caronii is a sublittoral species,<br />
occurring off <strong>West</strong> Africa in depths between 23<br />
(8-30) and 100 m. Of 16 recent depth records, 12<br />
(75%) are from depths between 23 and 50 m and<br />
4 (25%) are from depths greater than 50 m but<br />
less than 100 m, namely 73 m, 75 m, 80 m, and
192<br />
97-98 m. The Pillsbury specimens were taken on<br />
bottom with dark red bryozoans. The specimens<br />
collected by the Calypso were taken on bottoms<br />
with calcareous algae (three stations), algae and<br />
calcareous algae, and sand, algae, and calcareous<br />
algae (Forest and Guinot, 1966).<br />
Off <strong>West</strong> Africa ovigerous females have been<br />
collected in March, May, June, November, and<br />
December (Monod, 1956; Forest and Guinot,<br />
1966; Pillsbury).<br />
DISTRIBUTION.—Eastern Atlantic, from the<br />
Mediterranean and adjacent Atlantic, including<br />
the Azores, Ilhas Desertas, Madeira, the Canary<br />
Islands, the Cape Verde Islands, off the mainland<br />
from Senegal and Ghana, and from the offshore<br />
islands of the Gulf of Guinea, Principe, Sao Tome,<br />
and Annobon, in depths between 23 m and<br />
100 m. Monod (1956) reported material from<br />
Senegal and Principe; records since 1956 include<br />
the following:<br />
Madeira: No specific locality; Ponta de Sao Lourengo, 80<br />
m (Tiirkay, 1976b).<br />
Principe: 01°43'10"N, 07°28'2O"E, 73 m; and 01°43'N,<br />
07°28'55"E, 37 m (Forest and Guinot, 1966).<br />
Sao Tome: 00°25'40"N, 06°40'10"E, 50 m; and OO°25'-<br />
15"N, 06°43'05"E, 8-30 m (Forest and Guinot, 1966).<br />
Annobon: N of San Antonio, 23 m (Forest and Guinot,<br />
1966).<br />
Family OCYPODIDAE Rafinesque, 1815<br />
OCYPODIA Rafinesque, 1815:% [corrected to Ocypodidae by<br />
MacLeay, 1838:63, 64; name 375 on Official List].<br />
DOTINAE Dana, 1851c:286 [unavailable, based on a homonym].<br />
MACROPHTHALMINAE Dana, 1851c:286.<br />
UCAINAE Dana, 1851c:289.<br />
HELOECIACAEA H. Milne Edwards, 1852:153.<br />
DOTILLIDAE Stimpson, 1858b:98 [p. 44 on separate].<br />
CAMPTANDRIIDAE Stimpson, 1858b: 106 [p. 52 on separate].<br />
Gelasimiden Nauck, 1880:8, 17, 23, 64, 66.<br />
GELASIMIDAE Miers, 1886:viii.<br />
SCOPIMERIOAE Alcock, 1900:290, 295, 363.<br />
CAMPTANDRIINAE Serene and Kumar, 1971:75.<br />
CAMPTANDRINI Pretzmann, 1974:443.<br />
CLEISTOTOMATINI Pretzmann, 1977:66.<br />
EASTERN ATLANTIC GENERA.—Five, Calabarium,<br />
new genus, Ecphantor, new genus, Lillyanella, new<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
genus (see addendum), Ocypode, Telmatothrix, new<br />
genus, and Uca, each represented by species off<br />
tropical <strong>West</strong> Africa.<br />
EASTERN ATLANTIC SPECIES.—Six, all occurring<br />
in tropical <strong>West</strong> Africa. Two species were taken<br />
by the Pillsbury: Ocypode cursor and Uca tangeri.<br />
Three species were recorded by Monod (1956)<br />
under the same names used here, although he<br />
used Ocypoda for Ocypode.<br />
Three species of this family reported from <strong>West</strong><br />
Africa appear to be either erroneously labeled or<br />
misidentified:<br />
Ocypode ceratophthalmus (Pallas, 1772). An Indo-<br />
<strong>West</strong> Pacific species reported from Fernando Poo<br />
by Pesta (1911:54) (see Monod, 1956:530).<br />
Ocypode quadrata (Fabricius, 1787). A western<br />
Atlantic species reported under the name 0. albicans<br />
Bosc from "<strong>West</strong>afrika" by Balss (1922:79)<br />
(see Monod, 1956:399).<br />
Uca burgersi Holthuis, 1967. A western Atlantic<br />
species reported from Liberia under the name<br />
Uca mordax (Smith, 1870) by Rathbun (1900a:<br />
276; 1918:393; see also Monod, 1956:404) and<br />
Crane (1975:172, 326, 327, 604) and from Cameroon<br />
by Crane (1975:327, 604).<br />
It seems highly unlikely that these records are<br />
correct. We believe that these species should not<br />
be included in the list of <strong>West</strong> <strong>African</strong> Ocypodidae.<br />
Most modern authors (Balss, 1957:1663-1665;<br />
Guinot, 1967a:280-283; Serene, 1968:97-101)<br />
recognized three subfamilies within the family<br />
Ocypodidae: Ocypodinae, Scopimerinae, and<br />
Macrophthalminae. As the family group name<br />
Dotillidae Stimpson, 1858, is older than Scopimerinae<br />
Alcock, 1900, it has priority and should<br />
be used for the subfamily which contains both<br />
the genera Dotilla Stimpson, 1858, and Scopimera<br />
de Haan, 1833. We agree with Serene (1974:59-<br />
68) that a fourth subfamily, Camptandriinae<br />
Stimpson, 1858, should be recognized; it contains<br />
those genera formerly placed in the Macrophthalminae<br />
in which the male gonopods are strongly<br />
recurved. The four subfamilies of Ocypodidae<br />
that we now recognize are, therefore: Ocypodinae<br />
Rafinesque, 1815; Dotillinae Stimpson, 1958;<br />
Macrophthalminae Dana, 1851; and Camptan-
NUMBER 306<br />
driinae Stimpson, 1858. Only the first and the<br />
last are represented in the eastern Atlantic.<br />
Subfamily CAMPTANDRIINAE Stimpson, 1858<br />
REMARKS.—Stimpson (1858b: 106) erected a<br />
new family, Camptandriidae, to include a single<br />
genus, Camptandrium, which he was also newly<br />
describing. Later authors usually have synonymized<br />
the Camptandriidae with the subfamily<br />
Macrophthalminae Dana, 1851. However, the<br />
differences between the genera Macrophthalmus<br />
Desmarest, 1823, and Australoplax Barnes, 1966,<br />
on the one hand and the genera Camptandrium,<br />
Cleistostoma, Paracleistostoma, Tylodiplax, and Leipocten<br />
on the other are significant. It seems best,<br />
therefore, to separate the latter five genera as a<br />
distinct subfamily, for which the name Camptandriinae<br />
Stimpson, 1858, is available, confirming<br />
the views of Serene and Kumar (1971:75) and<br />
Serene (1974:59-68). Our reasons for recognizing<br />
Camptandriinae are practically the same as those<br />
noted by Serene and Kumar (1971) and Serene<br />
(1974).<br />
The characters that distinguish the Camptandriinae<br />
from the Macrophthalminae are the following:<br />
The most important is the shape of the<br />
Key to Genera of Camptandriinae<br />
193<br />
male gonopods, which are strongly recurved in<br />
the Camptandriinae, straight or slightly bent in<br />
the Macrophthalminae. The external maxillipeds<br />
in the Macrophthalminae have the merus shorter<br />
than the ischium and the last two segments thick;<br />
in the Camptandriinae the merus of the maxillipeds<br />
is as long as or longer than the ischium and<br />
the distal two segments are slender.<br />
The Camptandriinae are represented in <strong>West</strong><br />
Africa by four genera, all of which are new. In<br />
order to ascertain their affiliations with the other<br />
genera of Camptandriinae, we had to study each<br />
genus within the subfamily Camptandriinae.<br />
These genera are very poorly defined and their<br />
limits are rather vague, so that a revision on the<br />
generic level was badly needed. Fortunately a fair<br />
number of species of Camptandriinae was available<br />
to us and made it possible to obtain a better<br />
idea of the various genera and their limits. At the<br />
same time the material showed the necessity to<br />
erect several new genera, as a number of species<br />
did not fit the older genera as redefined by us.<br />
For this reason we give here a diagnosis of the<br />
genera of Camptandriinae recognized by us with<br />
a discussion of the species that we assign to them,<br />
those of which we consider of dubious status and<br />
those that in our opinion should be eliminated<br />
from the Camptandriinae.<br />
1. Male abdomen constricted near fifth somite so that gonopods are partly<br />
visible when abdomen is pressed against thorax 2<br />
Male abdomen gradually tapering towards terminal somite, completely<br />
covering gonopods, which are not visible when abdomen is pressed<br />
against thorax 6<br />
2. Gonopod of male with 2 distinct appendages at the distal end 3<br />
Gonopod of male tapering to narrow pointed apex, without<br />
appendages 4<br />
3. Appendages at tip of male gonopod longer than distal recurved part of<br />
shaft. Carapace subhexagonal Camptandrium<br />
Appendages at tip of male gonopod short, at most half length of recurved<br />
part of shaft. Carapace transverse quadrangular Paratylodiplax<br />
4. Anterolateral margin of carapace with teeth. Meri of ambulatory legs<br />
with distinct subdistal dorsal tooth. Male gonopod drawn out into<br />
slender, narrow tip Calabarium, new genus
194 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Anterolateral margin of carapace entire. Meri of ambulatory legs without<br />
subdistal dorsal tooth. Male gonopod with tip either short and triangular<br />
or with subdistal lobe 5<br />
5. Carapace only slightly wider than long, subhexagonal, without dorsal<br />
carina. Legs slender, merus of third (= longest) leg more than 3 times<br />
as long as wide. Male gonopod with broad subapical lobe. Fifth<br />
abdominal somite of male only slightly constricted<br />
Ecphantor, new genus<br />
Carapace distinctly wider than long, transversely quadrangular, with<br />
distinct transverse ridge. Legs broad, merus of third leg about twice as<br />
long as wide. Male gonopod regularly narrowing to triangular tip,<br />
without subapical lobe. Fifth male abdominal somite strongly constricted<br />
in basal part Deiratonotus, new genus<br />
6. First somite of male abdomen much wider than other somites and reaching<br />
coxae of fifth pereiopods 7<br />
First somite of male abdomen, though usually slightly wider than second,<br />
separated from coxae of fifth pereiopods by considerable distance .. 8<br />
7. Male gonopod ending in a pointed apex, which is not swollen. Chelipeds<br />
with strong sexual dimorphism Cleistostoma<br />
Male gonopod ending in blunt swollen apex. Males with chelipeds small,<br />
not different from those of females (dactylus of chelipeds without molarlike<br />
tooth) Ilyogynnis, new genus<br />
8. Merus of pereiopods with blunt spines on lower (flexor)<br />
surface Leipocten<br />
Merus of pereiopods without ventral spines 9<br />
9. Carapace without tubercles or transverse ridges dorsally. Anterolateral<br />
margins of carapace without teeth 10<br />
Carapace with transverse dorsal ridges and tubercles. Anterolateral margins<br />
of carapace with distinct or indistinct teeth. (Third maxilliped<br />
with small lobe at anterolateral angle of merus near base of carpus.<br />
Male gonopod without distinct appendages) 11<br />
10. Male gonopods swollen distally, provided with 1 or more distal appendages,<br />
which are at least half as long as recurved part of shaft. Apex of<br />
gonopod usually lying well free from basal part of shaft. Carapace with<br />
greatest width before middle Paracleistostoma<br />
Male gonopods tapering to narrow, pointed or blunt apex, not widened<br />
distally and without distal appendages. Gonopods so strongly recurved<br />
that tip lies over basal part of shaft. Carapace with greatest width<br />
behind middle Serenclla, new genus<br />
11. Anterolateral margin of carapace with 2 teeth, of which posterior very<br />
distinct, larger than anterior. Lateral margins of carapace converging<br />
posteriorly. Chelipeds showing strong sexual dimorphism. Propodi of<br />
second to fifth pereiopods longer than wide. Male gonopod not widened<br />
distally, distal margin deeply incised; morphological outer half of apex<br />
of gonopod slender and digitiform Telmatothrix, new genus
NUMBER 306 195<br />
Anterolateral margin of carapace with 2 low teeth, posteriormost less<br />
distinct than anterior. Lateral margins of carapace diverging posteriorly.<br />
Chelae of male small, of same shape and size as those of female.<br />
Propodi of second to fifth pereiopods hardly longer than wide. Male<br />
gonopod slightly widened distally, without an appendage, apex of<br />
gonopod not incised; morphological outer half of apex of gonopod a<br />
wide and blunt lobe, wider than long Tylodiplax<br />
Genus Calabarium, new genus<br />
TYPE-SPECIES.—Calabarium crinodytes, new species.<br />
ETYMOLOGY.—Derived from the type-locality,<br />
the New Calabar River; the gender of the generic<br />
name is neuter.<br />
DIAGNOSIS.—A genus of Camptandriinae. Carapace<br />
flat and uneven, hardly wider than long,<br />
subhexagonal, with prominent front. Regions<br />
faintly indicated, with several elevations. Dorsal<br />
surface of carapace with sparse pubescence of<br />
short, curved, dark colored hairs, densest at elevations<br />
and at margins. Prominent front with<br />
wide V-shaped incision on anterior margin. Epigastric<br />
tubercles obscure, placed on base of front.<br />
Anterolateral margins of carapace with distinct<br />
teeth. Posterior margin of orbits lying in single<br />
transverse line, showing no indentations.<br />
Cornea well developed, but narrower than<br />
eyestalk. Antennules obliquely folded. Antennae<br />
entering orbit. Lower margin of orbit distinct in<br />
frontal view, suborbital ridge lying some distance<br />
below it; both showing only a few denticles.<br />
Epistome short and concave, with anterior and<br />
posterior margins elevated. Third maxillipeds not<br />
filling oral cavity, distinct space present between<br />
anterior margin of merus of maxilliped and posterior<br />
margin of epistome. Merus about as long<br />
as ischium and partly covering exopod. Anterior<br />
margin of merus sloping backwards towards median<br />
line of body. Inner anterior angle of ischium<br />
produced obliquely forwards.<br />
Chelipeds equal, those of males usually not<br />
enlarged, of about same size, or smaller, than<br />
those of females. Third and fourth pereiopods<br />
longer than second and fifth. Merus of second to<br />
fifth pereiopods with strong subdistal anterodorsal<br />
tooth, least pronounced in fifth leg. Dark<br />
colored, curved short hairs and longer plumose<br />
soft setae present on legs.<br />
Female abdomen with 7 somites, all somites<br />
free, abdomen broadly oval with apex slightly<br />
produced. Male abdomen elongate triangular,<br />
with the second, third, and fourth somites fused.<br />
First somite not wider than second, not reaching<br />
coxae of fifth pereiopods. Abdomen narrowing<br />
suddenly at end of fourth and beginning of fifth<br />
somite, with definite constriction, so much so that<br />
gonopods partly exposed, even when abdomen<br />
fully pressed against thorax. Male gonopods are<br />
strongly recurved, ending in simple, narrow,<br />
drawn out apex.<br />
REMARKS.—The shape of the carapace of Calabarium<br />
shows a remarkable resemblance to that<br />
of Camptandrium, being hexagonal with distinct<br />
anterolateral teeth and being almost as long as<br />
wide. Also the male abdomen, which is so strongly<br />
constricted that the gonopods become partly exposed,<br />
resembles that of Camptandrium. However,<br />
the fact that the male gonopods in Calabarium end<br />
in a single slender point and do not show distal<br />
appendages immediately distinguishes the present<br />
genus from the Indo-<strong>West</strong> Pacific Camptandrium.<br />
Furthermore, the walking legs in Camptandrium<br />
do not possess dorsal teeth, and in most<br />
species the chelipeds show a distinct sexual dimorphism.<br />
The male gonopod of Calabarium somewhat<br />
resembles that of Deiratonotus, but has the<br />
apex narrowly drawn out. Deiratonotus furthermore<br />
differs from Calabarium by the wide and<br />
ridged carapace, which is more quadrangular<br />
than hexagonal, by the unarmed anterolateral<br />
margins of the carapace, and by the walking legs,<br />
which do not show any dorsal teeth on the merus;<br />
also in Deiratonotus there is a strong sexual dimorphism<br />
in the chelipeds.
196<br />
Calabarium crinodytesy new species<br />
FIGURE 46<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Nigeria: New Calabar River at Okpo<br />
waterside, Niger delta, 04°52'N, 06°54'E, 25 Jul 1978, C. B.<br />
Powell, 26, 3$ ov (one male is holotype) (L, W). Same<br />
locality, 7 Aug 1978, C. B. Powell, 26, 1 9 ov (W). Same<br />
locality, 15 Nov 1978, C. B. Powell, llcj, 5? (3 ov) (L, W).<br />
DESCRIPTION.—Carapace (Figure 46a) subhexagonal,<br />
slightly broader than long if lateral teeth<br />
included in width, slightly longer than broad<br />
exclusive of lateral teeth. Surface flat but uneven.<br />
Median area of carapace with following elevations<br />
from front to back: 2 indistinct epigastric, 2<br />
submedian mesogastric, 2 submedian cardiac, 1<br />
intestinal, latter five much stronger than epigastric,<br />
almost developed into tubercles. Each branchial<br />
region with 2 elevations, outermost placed<br />
slightly more posteriorly than innermost. Regions<br />
indistinctly indicated, cervical groove discernible.<br />
Lateral margins of carapace with strong tooth in<br />
anterior half, about twice as close to outer orbital<br />
angle as to posterolateral angle of carapace. This<br />
tooth about as strong as outer orbital tooth;<br />
between them are 2 smaller teeth on anterolateral<br />
margin. Posterior margin of orbit slightly sinuous,<br />
merging with lower orbital angle under broadly<br />
rounded curve. Neither orbital margin with incision,<br />
but lower orbital margin with distinct<br />
blunt tooth slightly before middle. Front horizontal,<br />
wide, anterior margin formed into 2 broad<br />
triangular teeth, each terminating in blunt point,<br />
separated by wide, V-shaped incision. Dorsal surface<br />
of carapace bearing short, stiff, dark brown,<br />
curved hairs, placed closest together on elevations<br />
and on margins. Few long, soft, plumose, uncolored<br />
hairs also present, especially on front.<br />
Eyes (Figure 46a,6) almost filling orbits, tapering<br />
distally, with distinct cornea. Some short,<br />
dark brown hairs placed on anterior surface of<br />
eyestalk. Lower margin of orbit with narrow denticulated<br />
process near base of antenna. Lateral to<br />
this process lower orbital margin with 1 or 2 small<br />
teeth on inner part, remainder of margin un-<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
armed. Suborbital ridge lying distinctly behind<br />
and below orbit, with 2 denticles.<br />
Antennules covered by front, folding obliquely.<br />
Antennae short, entering orbit, latter widely open<br />
at its base.<br />
Epistome wide, somewhat concave, with anterior<br />
and posterior margins elevated, ridge-like;<br />
surface with some irregular pits.<br />
Third maxillipeds (Figure 466) not completely<br />
filling oral cavity, space present between anterior<br />
margin of merus and posterior margin of epistome.<br />
Outer margin of merus much longer than<br />
inner, anterior margin oblique. Outer anterior<br />
angle of merus produced, but not conspicuously<br />
so. Ischium about as long as merus, with inner<br />
anterior angle produced forward. Merus partly<br />
covering exopod.<br />
First pereiopods of male (Figure 46^) small,<br />
equal, much shorter than second pereiopods and<br />
not always larger than chelipeds of female. Fingers<br />
of male cheliped about as long as palm. Tips<br />
of chelipeds hoof-shaped and closing, proximal<br />
half of cutting edges gaping. Cutting edge of<br />
dactylus with blunt tooth in proximal third, proximal<br />
half of cutting edge of fixed finger with<br />
several small denticles. Palm about twice as long<br />
as broad when not swollen, slightly longer than<br />
broad when swollen. Carpus somewhat shorter<br />
than palm, unarmed. Merus slightly shorter than<br />
chela, with inner and outer lower anterior angles<br />
broadly rounded. Ischium short, with tooth on<br />
inner margin and several tubercles near distal<br />
margin. All segments of first pereiopod, up to<br />
palm, bearing stiff, short and curved, dark hairs<br />
similar to those on dorsal surface of carapace; a<br />
few long, soft, plumose hairs also present. Cheliped<br />
of female (Figure 46
NUMBER 306 197<br />
a<br />
FIGURE 46.—Calabarium crinodytes, new genus, new species. Ovigerous female paratype:<br />
a, carapace; b, frontorbital region, ventral view; c, cheliped; d, abdomen. Male paratype:<br />
e, cheliped; /, second pereiopod; g, third pereiopod; h, fourth pereiopod; i, fifth pereiopod;<br />
j, abdomen in situ; k, basal part of abdomen. Holotype, male: /, gonopod.<br />
implanted on both upper and lower margins.<br />
Propodus slightly less than 1.5 times as long as<br />
dactylus, slightly widening proximally, bearing<br />
usual dark, short, and curved setae and some<br />
longer plumose hairs. Carpus about as long as<br />
dactylus. Merus, slightly longer than propodus,<br />
conspicuous by having large, sharp, forwardly<br />
curved, subdistal dorsal tooth. Pubescence of carpus<br />
and merus similar to that of propodus. Third<br />
leg (Figure 46g) longer and stronger than second,<br />
of about same structure and with same pubescence,<br />
except that lower margin of propodus and<br />
usually to lesser extent also that of merus, densely<br />
hairy, with numerous soft and long plumose hairs.
198<br />
Fourth leg (Figure 46/*) very similar in size, shape,<br />
and pubescence to third. Fifth leg (Figure 46?)<br />
distinctly shortest of all walking legs. Dactylus<br />
almost as long as propodus and longer than carpus.<br />
Subdistal short tooth of merus reduced to<br />
blunt lobe. In females legs longer and more slender<br />
than those of males, with short dark hairs<br />
more distinct and fewer long soft hairs.<br />
Female abdomen (Figure 46V) wide, reaching<br />
to coxae of pereiopods, broadly oval to almost<br />
circular, with all somites free. Seventh somite<br />
with apex somewhat broadly triangularly produced.<br />
Male abdomen (Figure 46/,A:) elongate triangular.<br />
First somite very short, about as wide as<br />
second, failing to reach base of fifth pereiopod,<br />
separated from coxa of that leg by considerable<br />
distance. Second, third, and fourth abdominal<br />
somites fused, but lines between them rather distinct.<br />
Second somite shortest of the three, but<br />
longer than first. Deep incision in lateral margin<br />
of abdomen indicating separation of second and<br />
third somites. Third and fourth somites of about<br />
equal length, fourth narrowing distinctly in distal<br />
part. Fifth and sixth somites free, fifth much<br />
narrower than third or basal part of fourth, narrowest<br />
at base, widening distally. Sixth somite<br />
about as wide as fifth but somewhat shorter,<br />
narrowing slightly distally. Seventh somite about<br />
as long as fifth but somewhat shorter, distinctly<br />
narrower, ending in almost semicircular distal<br />
margin. Outer margin of depression in thoracic<br />
sternum, which receives abdomen when latter<br />
completely folded against body, regularly triangular<br />
in shape; apex of depression rounded like<br />
seventh abdominal somite, lateral margins not<br />
showing any constriction; space present between<br />
margin of depression and outer margin of abdomen<br />
at level of constriction between fourth and<br />
fifth somites filled by part of ventral surface of<br />
gonopods, latter partly exposed even when abdomen<br />
completely pressed against thorax.<br />
Male gonopod (Figure 46/) with characteristic<br />
recurved shape of Camptandriinae. Anterior limit<br />
of curve just failing to reach separation between<br />
sternites of first and second pereiopods. Gonopod<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
tapering regularly from base to top, widening<br />
somewhat in distal fourth, then narrowing rapidly<br />
to slender, sharp, drawn out point; latter<br />
simple, lacking appendages. Distal part of gonopod<br />
ornamented with few, scattered, minute tubercles,<br />
with some tubercles arranged in short<br />
row on inner margin proximal to tip.<br />
MEASUREMENTS.—Carapace lengths of males<br />
3.8 to 7.8 mm, of nonovigerous females 3.0 to<br />
5.0 mm, of ovigerous females 5.5 to 9 mm. Eggs<br />
numerous and small, diameter 0.3 to 0.35 mm.<br />
REMARKS.—The three new genera and new<br />
species, Calabrium crinodytes, Ecphantor modestus, and<br />
Telmatothrix powelli, are so far the only known<br />
<strong>West</strong> <strong>African</strong> Camptandriinae. Ecphantor can immediately<br />
be distinguished from the other two by<br />
the absence of anterolateral teeth on the carapace.<br />
The present new species is characterized by the<br />
shape of the front, which is more deeply incised<br />
anteriorly; furthermore, the chelipeds in Telmatothrix<br />
show a strong sexual dimorphism. Telmatothrix<br />
lacks the strong subdistal dorsal tooth on<br />
the merus of the walking legs. Also, the male<br />
abdomens and gonopods of the three species show<br />
significant differences, which necessitate recognition<br />
of a separate genus for each of the three <strong>West</strong><br />
<strong>African</strong> species.<br />
TYPE-LOCALITY.—New Calabar River at Okpo<br />
waterside, Niger Delta, Nigeria, 04°52'N, 06°-<br />
54'E.<br />
DISPOSITION OF TYPES.—The holotype (Crust.<br />
D. 31950), a male, cl 5.7 mm, cb 6.3 mm, is in<br />
the Rijksmuseum van Natuurlijke Historie, Leiden.<br />
Paratypes have been deposited in that institution<br />
as well as in the National Museum of<br />
Natural History, <strong>Smithsonian</strong> <strong>Institution</strong>, Washington,<br />
D.C.<br />
ETYMOLOGY.—The specific epithet is derived<br />
from the generic name of the aquatic lily, crinum,<br />
on which the species was found.<br />
BIOLOGY.—Mr. C. B. Powell (in litt., 26 July<br />
1978) informed us as follows:<br />
Specimens were collected from submerged leaves of the<br />
aquatic lily Crinum natans Baker in the zone of mixed Rhizophora<br />
and Pandanus in the New Calabar River. The water at<br />
the site is nearly fresh, but the tidal range is about 2 meters
NUMBER 306 199<br />
and there is considerable disturbance to the littoral zone<br />
caused by the wash from motorboats and barges moving at<br />
top speed—when they pass by, one has to scramble to shore<br />
to avoid the large waves. The other Malacostraca present<br />
are Parorchestia n. sp., Cirolana, n. sp., Metastenasellus n. sp.,<br />
Caridina africana Kingsley, Palaemonetes africanus Balss, Macrobrachium<br />
vollenhovenii (Herklots), M. felicinum Holthuis, Potamalpheops<br />
monodi (Sollaud), P. pylorus Powell (a single specimen),<br />
a new Potamalpheops, Telmatothrix powelli, Metagrapsus<br />
curvatus (H. Milne Edwards), Sesarma alberti Rathbun, S.<br />
angolense De Brito Capello, S. elegans Herklots, and S. buettikoferi<br />
De Man.<br />
Ovigerous females were collected in July, August,<br />
and November.<br />
DISTRIBUTION.—Known only from the typelocality.<br />
Genus Camptandrium Stimpson, 1858<br />
FIGURE 47<br />
Camptandrium Stimpson, 1858b: 106 [type-species: Camptandrium<br />
sexdentatum Stimpson, 1858, by monotypy; gender:<br />
neuter; name 289 on Official List].<br />
DIAGNOSIS.—Carapace hexagonal rather than<br />
transversely quadrangular, with 2 or 3 anterolateral<br />
teeth, posteriormost reaching sideways. Surface<br />
of carapace flat, uneven, with elevations that<br />
may form longitudinal or transverse ridges. Epigastric<br />
lobes rather conspicuous, placed in basal<br />
part of front. Chelipeds equal, with conspicuous<br />
sexual dimorphism (except in the aberrant C.<br />
elongatum); dactylus in male with single tooth on<br />
cutting edge, tooth being absent from fixed finger;<br />
in female neither finger with tooth. Meri of<br />
pereiopods without spines, but lower distal angle<br />
may be produced. First somite of male abdomen<br />
only slightly wider than second and not extending<br />
to coxae of fifth pereiopods. Second to fifth somites<br />
of male abdomen fused, but lines between<br />
somites more or less distinctly visible. Abdomen<br />
gradually narrowing distally, but constricted at<br />
level of fifth somite so that gonopods partly visible,<br />
even when abdomen folded fully back against<br />
thorax. Male gonopods strongly recurved, each<br />
with apex somewhat swollen, ending in 2 long<br />
slender appendices, one practically straight, the<br />
FIGURE 47.—Camptandrium sexdentatum Stimpson: a, female,<br />
dorsal view; b, male abdomen; c, male chela; d, gonopod.<br />
(All from Shen, 1932, figs. 138, 140a,c,e.)<br />
other sinuous and longer of the two; appendages<br />
longer than recurved part of gonopod.<br />
REMARKS.—Shen (1935:30) assigned the following<br />
four species to the genus Camptandrium: C.<br />
sexdentatum Stimpson, 1858 (Figure 47); C. elongatum<br />
Rathbun, 1929; C. anomalum Shen, 1935; and<br />
C. paludicola Rathbun, 1909. Since 1935 the following<br />
three new species have been described in<br />
Camptandrium: C. starmuehlneri Pretzmann, 1968;<br />
C. ambonensis Serene and Moosa, 1971; and C.<br />
rathbunae Takeda, 1972. In our opinion four of<br />
these species are true Camptandrium, viz., C. sexdentatum,<br />
C. elongatum, C. ambonensis, and C. rathbunae.<br />
Camptandrium starmuehlneri Pretzmann (1968:16;<br />
see also Pretzmann, 1969:5, figs 1-4), which is<br />
known only from the female holotype, probably<br />
is a good Camptandrium near C. elongatum; but since<br />
the shape of the abdomen and the gonopods of<br />
the male are not known, we are not quite certain<br />
of its generic status. Serene and Moosa (1971:6)
200<br />
suggested that the species might belong either to<br />
the genera Shenius Serene or Ilyograpsus Barnard,<br />
which, however, seems unlikely. Camptandrium paludicola<br />
Rathbun belongs to the grapsid genus<br />
Ilyograpsus Barnard, 1955, being synonymous with<br />
the type and only known species of the genus,<br />
Ilyograpsus rhizophorae Barnard, 1955, the correct<br />
name of which thus is Ilyograpsus paludicola (Rathbun,<br />
1909) (see Crosnier, 1965:31). Camptandrium<br />
anomalum Shen was removed from Camptandrium<br />
by Serene (1971:916, 917) and made the type of<br />
the new genus Shenius Serene. Shenius should be<br />
removed from the Camptandriinae because the<br />
male gonopods of Shenius anomalus (Shen) are not<br />
recurved. Serene (1974:60) placed the genus Shenius<br />
in the subfamily Scopimerinae. Shenius also<br />
was left out of his (1974:62-66) lists of the genera<br />
and species of Camptandriinae. However, it was<br />
included in Serene's (1974:66) key to the genera<br />
of Camptandriinae.<br />
The range of the genus extends from India to<br />
Japan, Korea, the Malay Archipelago, and perhaps<br />
New Caledonia.<br />
Genus Cleistostoma de Haan, 1833<br />
FIGURE 48<br />
Cleistostoma de Haan, 1833:5, pi. B, pi. 7: fig. 3 [misspelled<br />
Cleistotoma on pi. 7] [type-species: Ocypode (Cleistostoma)<br />
dilatata de Haan, 1833, by monotypy; gender: neuter].<br />
DIAGNOSIS.—Carapace convex both in longitudinal<br />
and transverse directions, not showing<br />
any appreciable ridges dorsally, but evenly hairy<br />
with short, dark pubescence. Carapace quadrangular<br />
in outline, being wider than long. Epigastric<br />
ridges not very distinct, extending forward far<br />
beyond base of front. Latter shallowly concave in<br />
middle, with anterior margin rounded, lacking<br />
anterolateral lobes. Suborbital ridge reaching far<br />
forward, protruding beyond distinct lower orbital<br />
margin. Upper orbital margin showing no incisions.<br />
Anterolateral margin of carapace lacking<br />
teeth apart from outer orbital angle, continuing<br />
posteriorly into posterolateral margin. No additional<br />
carina present on posterolateral part of<br />
a<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 48.—Cleistostoma dilatatum (de Haan): a, female, dorsal<br />
view; b, male, frontal view; c, male abdomen; d, male<br />
chela; e, gonopod; /, tip of gonopod, enlarged. (From Shen,<br />
1932, figs. 145, 146, 147b,c, 148.)<br />
carapace. Chelipeds equal, left and right, but<br />
showing strong sexual dimorphism: much larger<br />
and stronger in male than in female and with<br />
different shape. Dactylus of male cheliped with<br />
distinct molariform tooth on cutting edge; such<br />
tooth absent on fixed finger of male cheliped or<br />
on either finger of that of female. Pereiopods<br />
lacking spines on merus. First somite of male<br />
abdomen very wide, reaching to coxae of fifth<br />
pereiopods; second somite much narrower. Second<br />
to fourth somites of male abdomen fused;<br />
fifth and sixth free. Lateral margins of abdomen<br />
somewhat sinuous, fitting snugly against thoracic<br />
sternum, covering gonopods completely. Strongly<br />
recurved male gonopods gradually narrowing toward<br />
apex, lacking distal appendages. Detailed<br />
figures of the type-species, C. dilatatum, are pro-
NUMBER 306 201<br />
vided by Shen (1932:236, figs. 145-148, pi. 10;<br />
fig. 4).<br />
REMARKS.—Guinot and Crosnier (1963) listed<br />
the following species as belonging to the genus<br />
Cleistostoma: C. dilatatum (de Haan, 1833) (Figure<br />
48), C. wardi Rathbun, 1926, C. dotilliforme Alcock,<br />
1900, C. macneilli Ward, 1933, C. edwardsii Mac-<br />
Leay, 1838, and C. algoense Barnard, 1954. We<br />
have examined material of all these species except<br />
C. macneilli. In our opinion, only C. dilatatum, the<br />
type-species of the genus, which is known from<br />
Japan, North China, and Korea, is a true Cleistostoma;<br />
all the other species should be transferred<br />
to other genera. Cleistostoma wardi and C. dotilliforme,<br />
perhaps also C. macneilli, are placed in the<br />
genus Paracleistostoma (p. 208), and C. edwardsii<br />
and C. algoense are referred below to the genus<br />
Paratylodiplax (p. 209), as they differ in so many<br />
respects from Cleistostoma that it is impossible to<br />
assign them to that genus.<br />
Ocypode {Cleistostoma) dilatata de Haan, 1833, is<br />
a senior homonym of Ocypode (Macrophthalmus)<br />
dilatata de Haan (1835:55, pi. 15: fig. 2). We<br />
propose the replacement name, Macrophthalmus<br />
abbreviatus, new name, for the invalid Ocypode<br />
{Macrophthalmus) dilatata de Haan, 1835, for<br />
which, so far as we can ascertain, no other name<br />
has ever been proposed. The epithet abbreviatus is<br />
chosen because de Haan (1835:26) himself used<br />
it for the species, be it as a nomen nudum.<br />
In the original publication of the genus, de<br />
Haan (1833) used two different spellings: Cleistostoma<br />
in the text (p. 5) and on plate B, and<br />
Cleistotoma on plate 7: figure 3. In later parts of<br />
the Fauna Japonica de Haan was not consistent<br />
either; he mostly used Cleistostoma (part 2(1835):<br />
26, 27, 55, 56; part 3(1837), pi. 16: fig. 1), but at<br />
the end used again Cleistotoma (part 7(1849):233,<br />
234), namely in the index to the work. This 1849<br />
use of Cleistotoma was perhaps because H. Milne<br />
Edwards (1837:67, 68) had adopted that spelling;<br />
de Haan in his index (1849:233) referred to u Cleistotoma,<br />
n. Edw^ From de Haan's original publication<br />
it is not clear which spelling is the correct<br />
one; neither de Haan's (1835, 1837, 1849) later<br />
uses of the two spellings, nor H. Milne Edwards'<br />
(1837) use of Cleistotoma, indicates why a certain<br />
spelling is used. Thus none of these actions can<br />
be construed as first reviser actions. The actual<br />
first reviser is Dana (1852b:312, 313) who used<br />
the spelling Cleistostoma, and said in a footnote on<br />
p. 312 "not Cleistotoma." By Dana's action Cleistostoma<br />
is the correct spelling of the generic name;<br />
Cleistotoma, being an incorrect original spelling,<br />
has no nomenclatural standing.<br />
Agassiz (1846:89, 90) emended Cleistotoma to<br />
Clistotoma. Clistotoma Agassiz, 1846, thus is an<br />
unjustified emendation of the incorrect original<br />
spelling, but this notwithstanding is an available<br />
name, even though it is invalid as a junior objective<br />
synonym of Cleistostoma de Haan, 1833.<br />
Genus Deiratonotus, new genus<br />
FIGURE 49<br />
TYPE-SPECIES.—Paracleistostoma cristatum De<br />
Man, 1895.<br />
ETYMOLOGY.—From the Greek, deirado (ridge)<br />
and noton (back); gender of name is masculine.<br />
DIAGNOSIS.—Carapace about quadrangular in<br />
outline, distinctly broader than long with lateral<br />
FIGURE 49.—Deiratonotus cnstatus (De Man): a, male, dorsal<br />
view; b, male abdomen; c, abdomen in situ, male, cl. 7.5<br />
mm, Japan; d, male chela; e, gonopod; /, tip of gonopod,<br />
enlarged (a,M-/from Shen, 1932, figs. 141, 143b,c, 144).
202<br />
margins convex. Surface very flat, except for 3<br />
elevations forming a transverse ridge slightly behind<br />
middle, and short but distinct epigastric<br />
ridges. Ridges accentuated by covering of short<br />
dark brown hairs absent from rest of carapace<br />
except for marginal rim and additional ridge in<br />
posterolateral area. Inner dorsal orbital angle<br />
rounded. Front slightly concave in middle and<br />
with anterolateral angles angularly rounded but<br />
not produced. Suborbital ridge lying somewhat<br />
below distinct lower orbital margin, but reaching<br />
farther forward. Anterolateral margin of carapace<br />
unarmed. Sharp carina placed on posterolateral<br />
part of carapace, joining lateral margin anteriorly.<br />
Chelipeds with distinct sexual dimorphism.<br />
In male, chelipeds large, chelae with large molariform<br />
tooth on cutting edge of dactylus, but not<br />
on fixed finger. Meri of walking legs unarmed.<br />
Male abdomen with first somite somewhat wider<br />
than any of other somites, but failing to reach<br />
coxae of fifth legs by considerable distance. First<br />
somite of male abdomen free, second, third,<br />
fourth, and fifth fused, sutures only faintly indicated;<br />
incision in lateral margin marking separation<br />
between second and third somites. Outline<br />
of male abdomen narrowing gradually towards<br />
telson, but strongly constricted at level of the fifth<br />
somite, constriction revealing part of gonopods,<br />
even when abdomen is fully pressed against thoracic<br />
sternum. Exposed parts of gonopods, however,<br />
not flush with surface of abdomen. Sixth<br />
somite of male abdomen and telson are free. Male<br />
gonopod strongly recurved, ending in sharp point,<br />
neither swollen distally nor bearing appendages<br />
there.<br />
REMARKS.—This genus, in the shape of the<br />
abdomen, which leaves part of the gonopods<br />
exposed, resembles the genera Camptandrium and<br />
Paratylodiplax, and differs in this respect from all<br />
other Camptandriine genera. In Camptandrium<br />
and Paratylodiplax, however, the gonopods have<br />
two distinct appendages distally, while in Deiratonotus<br />
such appendages are entirely lacking. In<br />
the general shape of the carapace, with elevated<br />
hairy ridges the present new genus resembles the<br />
<strong>West</strong> <strong>African</strong> genus Telmatothrix.<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
The type-species is the Indo-<strong>West</strong> Pacific<br />
Paracleistostoma cristatum De Man, 1895 (Figure<br />
49). A second species that we assign to this genus<br />
is Paracleistostoma japonicum Sakai. Although we<br />
saw no material, Sakai's (1934:321, fig. 26, pi. 18:<br />
fig. 1) good description indicates the main characters<br />
of Deiratonotus: ridged carapace, male abdomen<br />
revealing the gonopods, and the shape of<br />
the male gonopod. Deiratonotus cristatus is known<br />
from Japan, China and Korea, D. japonicus only<br />
from Japan.<br />
Genus Ecphantor, new genus<br />
TYPE-SPECIES.—Ecphantor modestus, new species.<br />
ETYMOLOGY.—From the Greek word ekphantor<br />
(revealer), referring to the partly exposed gonopods<br />
of the male; gender of name is masculine.<br />
DIAGNOSIS.—A genus of Camptandriinae. Carapace<br />
subhexagonal, being only slightly (1.1 to<br />
1.2 times) wider than long. Surface of carapace<br />
flat but slightly uneven, with dense, uniform,<br />
short pubescence. Front directed obliquely down.<br />
Epigastric lobes strong, placed on base of front;<br />
no other distinct ridges present. Anterolateral<br />
margins without teeth. Orbits transverse, situated<br />
in common straight line. Eyes with distinct pigmented<br />
cornea. Antennules obliquely folded. Antennae<br />
entering orbit. Lower margin of orbit<br />
straight, without tubercles or teeth. Epistome<br />
short, concave. Third maxilliped filling entire<br />
oral field; merus and ischium covering palp and<br />
part of exopod. Merus shorter than ischium. Anterolateral<br />
angle of merus widely rounded, not<br />
produced. Inner anterior angle of ischium produced<br />
forward. Chelipeds equal, left and right;<br />
those of adult male much stronger and more<br />
robust than in female. Dactylus of cheliped in<br />
male with strong tooth in basal half of cutting<br />
edge, no other teeth present. No teeth on either<br />
finger in female. Tips of fingers of chelipeds<br />
spoon-shaped. Ambulatory legs longer than chelipeds;<br />
third and fourth pereiopods longer than<br />
second and fifth. No spines or teeth on any segments<br />
of ambulatory legs, latter covered by uniform<br />
short pubescence and scattered longer hairs.
NUMBER 306 203<br />
Male abdomen narrow; all somites distinct, regularly<br />
tapering from first to sixth. First somite<br />
failing to reach coxa of fifth pereiopods. Fifth<br />
somite slightly constricted in basal part, first gonopods<br />
thereby slightly exposed. Female abdomen<br />
very wide, almost semicircular. Male gonopods<br />
strongly recurved; pointed apex with broad subdistal<br />
lobe, but without appendages.<br />
REMARKS.—The shape of the carapace of this<br />
new genus is somewhat similar in its hexagonal<br />
form to that of Calabarium, but it can immediately<br />
be distinguished by the total lack of anterolateral<br />
teeth. Like Camptandrium, Calabarium, Paratylodiplax,<br />
and Deiratonotus, Ecphantor has the abdomen<br />
constricted in such a way that the gonopods are<br />
partly visible even when the abdomen is firmly<br />
pressed against the sternum. The distal part of<br />
the gonopod, which carries a broad subdistal lobe<br />
but no appendages, distinguishes Ecphantor immediately<br />
from the other genera in the subfamily.<br />
Ecphantor modestus, new species<br />
FIGURE 50<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Nigeria: Elechi Creek at College of Science<br />
and Technology, Port Harcourt, 04°47.3'N, 06°58.6'E, 19<br />
Oct 1979, C. B. Powell, Ic5 holotype (L). Same locality, 21<br />
Feb 1979, C. B. Powell, 1$ paratype (W). Same locality, 25<br />
Feb 1979, C. B. Powell, 1? paratype (L).<br />
DESCRIPTION.—Carapace (Figure 50a) subhexagonal,<br />
slightly (1.1 to 1.2 times) wider than long.<br />
Surface flat, margins slightly elevated; with pubescence<br />
removed, surface appearing slightly uneven,<br />
smooth, shining and pitted. Hair cover of<br />
carapace uniform, consisting of short, dark,<br />
curved, stiff hairs; cover dense enough to obscure<br />
surface, especially so since mud particles usually<br />
are caught between hairs. Central part of cervical<br />
groove rather distinct, with another median transverse<br />
depression some distance behind it; latter,<br />
however, very vague. Front directed obliquely<br />
down, separated from rest of carapace by distinct<br />
epigastric lobes, falling off precipitously anteriorly,<br />
separated from each other by short, distinct<br />
median groove; short longitudinal lateral groove,<br />
less distinct than median groove, present on either<br />
side of frontal lobes. Width of front at level of<br />
epigastric lobes slightly less than half frontorbital<br />
width; distal width of front 2/5 frontorbital<br />
width. Upper surface of front rather flat, slightly<br />
concave in middle. Anterior margin produced<br />
somewhat forward in middle, top slightly emarginate.<br />
Anterolateral angles of front broadly<br />
rounded. Lateral margin of front merging with<br />
orbital margin under wide curve. Orbital margin<br />
with indistinct notch in inner, posterior part,<br />
latter otherwise slightly convex. Outer orbital<br />
angle rectangularly rounded. Anterolateral margin<br />
of carapace unarmed. Angle between anterolateral<br />
and posterolateral margins blunt and<br />
wide. Posterolateral margin less sharply defined<br />
than anterolateral. Angle between anterolateral<br />
and posterolateral margins lying in anterior half<br />
of body, forming widest part of carapace. Faint<br />
but broad rim present on posterior margin; latter<br />
about 2/3 as wide as frontorbital width.<br />
Eyes with cornea distinct and globular but<br />
narrower than eyestalk. Lower margin of orbit<br />
straight, smooth, bearing several long hairs, lacking<br />
granules or teeth. Suborbital ridge similarly<br />
unarmed.<br />
Antennules folding obliquely, almost transversely.<br />
Antennae entering orbit. Flagellum short,<br />
consisting of 4 articles and terminating in long<br />
seta.<br />
Epistome wide, short, concave, anterior margin<br />
almost straight or slightly sinuous, posterior margin<br />
ending posteriorly in median carina separating<br />
the two halves of the oral field. No granules<br />
present.<br />
Third maxillipeds filling entire oral field. In<br />
ventral view merus and ischium covering larger<br />
part of both exopod and palp, of both only basal<br />
part being visible. Merus shorter than ischium,<br />
with anterolateral angle broadly rounded, not<br />
produced or auricular. Palp articulating in middle<br />
of anterior margin of merus. Exopod rather<br />
broad, with well-developed flagellum.<br />
First legs (Figure 50c,A) showing conspicuous<br />
sexual dimorphism. Chelipeds of male relatively<br />
small, shorter than any of other legs, but distinctly
204<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 50.—Ecphantor modestus, new genus, new species. Male, holotype: a, dorsal view;<br />
b, anterior part of body, ventral view; c, cheliped; d, second pereiopod; e, abdomen;/, gonopod;<br />
g, apex of gonopod, enlarged. Smaller female, paratype: A, cheliped; i-l, second to fifth<br />
pereiopods.
NUMBER 306 205<br />
more robust than those of female. Left and right<br />
chelipeds equal in both sexes. In adult male<br />
fingers of chela somewhat shorter than palm,<br />
gaping basally, tips spoon-shaped, with small<br />
horn-colored hoofs. Cutting edge of dactylus<br />
bearing strong tooth in gap, no other teeth present<br />
on it or on fixed finger. Palm slightly swollen,<br />
about 2/3 as high as long, highest in distal half.<br />
Carpus short, cup-shaped, slightly more than half<br />
as long as palm and distinctly less high. Merus<br />
elongate, slightly more than half as high as palm<br />
and about 2.5 times as long as wide. Chelipeds<br />
slenderer in female. Fingers narrow and elongate,<br />
somewhat shorter than palm, with spoon-shaped<br />
tips and fully unarmed cutting edges. Palm elongate,<br />
being about 3 times as long as high, increasing<br />
slightly in height distally. Carpus about as<br />
long as and as high as palm. Merus about 4 times<br />
as long as high, being slightly higher than carpus<br />
or palm.<br />
Of following legs (Figure 50d, i-l) second and<br />
third (= third and fourth pereiopods) longest.<br />
Second and fifth pereiopods (= first and fourth<br />
ambulatory legs) shorter, but still longer than<br />
cheliped. Second pereiopod slightly more slender<br />
than fifth, especially in shape of propodus. Dactylus<br />
of each ambulatory leg slender, about as<br />
long as propodus, unarmed. Carpus about as long<br />
or slightly shorter than propodus and about 2/3<br />
as long as merus. Merus totally lacking subdistal<br />
dorsal tooth, at most barely noticeable slight<br />
elevation there. All legs covered with same short,<br />
stiff, curved, dark hairs as are on carapace. Long,<br />
soft, somewhat plumose hairs present between<br />
short hairs, more distinct on anterior and posterior<br />
margins of segments. Longer hairs denser on<br />
posterior margins of dactylus and propodus than<br />
on anterior margins, and denser on posterior legs<br />
than on anterior ones.<br />
Male abdomen (Figure 50
206<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
type as follows:<br />
into anterolateral angles, latter triangularly pro-<br />
The specimen was taken from a bed of leaf fragments etc.,<br />
with no sand and not much mud. At low tide seepage water<br />
drains through the bed, and the crab was in a handful of the<br />
duced. Posterior margin of orbit sinuous, with<br />
distinct incision at inner end. Suborbital ridge<br />
placed below distinct lower orbital margin, but<br />
litter I had scooped from the edge of the main trickle of reaching farther forward. Anterolateral margin of<br />
water. The most common macro-invertebrate present in the<br />
litter was the amphipod Quadrivisio,<br />
carapace showing 2 blunt lobes behind outer<br />
orbital angle. Sharp carina placed in posterolat-<br />
According to Powell (1979:126), Elechi Creek eral part of carapace above and mesial to postero-<br />
is a mangrove creek of<br />
lateral margin, forming pseudo-posterolateral<br />
margin, about parallel with true posterolateral<br />
about 6-12 meters wide and 1-2 meters deep, draining a<br />
large area of mangrove peat flats. The vegetation consists margin but not touching it. Chelipeds of male<br />
mainly of Avicemia africana and Lagunculana racemosa; also small, equal, similar to those of female; fingers<br />
present are Rhizophora spp. incl. R. mangle, the fern Acrostichumwith<br />
spoon-shaped tips, neither showing molari-<br />
aureum and the grass Paspalum vaginatum. The tidal range isform<br />
tooth. No sexual dimorphism shown by<br />
close to 2 meters so that at low tide the creek is dry except<br />
chelipeds. Meri of walking legs lacking spines.<br />
for a small shallow stream of drainage water running along<br />
the central sandy part of the creek bottom. . .. The maxi- Male abdomen about triangular in shape. First<br />
mum salinity of the creek water is probably about 20% . . . somite widest and shortest, reaching laterally as<br />
the normal dry-season salinity range is in the order of 5- far as coxae of fifth pereiopods. Second to fifth<br />
20%. In the rainy-season the maximum salinity is probably<br />
nearly as high . . . but the average and minimum salinities<br />
must be somewhat less.<br />
somites fused, separated by indistinct sutures<br />
only. Second somite much narrower than first<br />
and about as wide as third. Sixth somite and<br />
The type-locality of Ecphantor modestus is also telson free. Lateral margins of male abdomen<br />
the type-locality of Potamalpheops pylorus Powell, from third somite to telson about straight, being<br />
1979, the description of which is cited here. Pow- only slightly concave in sixth somite. Abdomen<br />
ell (1979:127) also provided a list of other inver- fitting closely against thoracic sternum, entirely<br />
tebrates present at the type locality.<br />
covering gonopods. Male gonopods recurved,<br />
Ovigerous females were collected in February. ending in widened bulbous apex, lacking distal<br />
DISTRIBUTION.—Known only from the type- appendage.<br />
locality.<br />
REMARKS.—The only species that we assign to<br />
this genus is Paracleistostoma microcheirum Tweedie<br />
(Figure 51), of which we examined material from<br />
Genus Ilyogynnis, new genus<br />
Singapore (USNM 137238). The genus Ilyogynnis<br />
FIGURE 51<br />
is primarily distinguished from Paracleistostoma by<br />
the very wide first segment of the male abdomen,<br />
TYPE-SPECIES.—Paracleistostoma microcheirum which reaches sideways as far as the coxae of the<br />
Tweedie, 1937.<br />
fifth pereiopods, the absence of sexual dimorph-<br />
ETYMOLOGY.—From the Greek ilyos (mud) and ism in the chelipeds, and by the shape of the male<br />
gynnis (a womanish man); gender of generic name gonopod, which lacks a distal appendage. In other<br />
is masculine.<br />
respects the male gonopod strongly resembles that<br />
DIAGNOSIS.—Carapace transversely quadran- of Paracleistostoma. Also the shape of the orbit with<br />
gular, only slightly wider than long, with lateral the distinct incision at the inner end of the pos-<br />
margins convex. Surface of carapace flat, but terodorsal margin, and the rather distinct regions<br />
regions rather well indicated by grooves. Dense of the carapace serve to distinguish the new genus.<br />
pubescence covering entire carapace. Epigastric The correct spelling of the name of the present<br />
ridges distinct. Front deeply grooved longitudi- species is Ilyogynnis microcheirum (Tweedie). Tweenally<br />
in middle, with oblique grooves extending die (1937) did not give the derivation of the
NUMBER 306 207<br />
FIGURE 51.—Ilyogynnis microcheirum (Tweedie): a, anterior part<br />
of carapace of male; b, abdomen in situ, male, cl 6.5 mm,<br />
South China Sea; c, gonopod. (a,c, from Tweedie, 1937, fig.<br />
6a,d).<br />
specific epithet microcheirum, and since cheirum is<br />
not a known adjective, the word "microcheirum"<br />
must be considered either a noun or an arbitrary<br />
combination of letters; in either case the removal<br />
of the epithet from a neuter genus (Paracleistostoma)<br />
to one of which the name is masculine<br />
{Ilyogynnis) does not necessitate a change in spelling<br />
of the epithet.<br />
Genus Leipocten Kemp, 1915<br />
FIGURE 52<br />
Leipocten Kemp, 1915:243 [type-species: Leipocten sordidulum<br />
Kemp, 1915, by monotypy; gender: neuter].<br />
DIAGNOSIS.—Carapace subquadrangular,<br />
slightly convex in both longitudinal and transverse<br />
directions. Upper surface with regions<br />
weakly indicated and bearing hairs and granules.<br />
Epigastric lobes faint, placed on base of front.<br />
Anterolateral teeth present. Third maxilliped<br />
with merus hardly auriculate. Part of exopod of<br />
third maxilliped visible. Chelipeds with distinct<br />
sexual dimorphism. Male chela with molariform<br />
tooth on cutting edge of dactylus, but not on<br />
fixed finger. Female with strong spines on chela,<br />
fingers about as long as palm. Walking legs short,<br />
robust, with spines on lower margin of merus.<br />
Male abdomen with first somite narrow, not<br />
reaching coxae of fifth pereiopods. Second and<br />
third abdominal somites of male fused. Male<br />
gonopod strongly recurved, ending in 2 processes,<br />
one narrow, pointed, the other lobiform, with<br />
several curved spines.<br />
REMARKS.—Only one species, the type, L. sordidulum<br />
(Figure 52), is known in this genus. It has<br />
an Indo-<strong>West</strong> Pacific distribution (India to Formosa<br />
and Australia).<br />
Serene (1974:62, 64, 66) justifiably intimated<br />
that Baruna Stebbing, 1904, probably is a senior<br />
synonym of Leipocten Kemp, 1915. Unfortunately,<br />
Stebbing's (1904:3, 4, pi. 1: fig. A) description<br />
and illustrations of his new genus Baruna and its<br />
single new species, Baruna socialis Stebbing, 1904,<br />
are insufficient by modern standards. For one<br />
thing the male pleopods have neither been described<br />
nor figured. On the other hand, the great<br />
similarity of the shapes of the male chelipeds, of<br />
the peculiar pereiopods, of the male abdomen,<br />
and of the anterolateral margin of the carapace<br />
of Leipocten and Baruna, as well as the sizes and<br />
the habitats of their species, strongly suggest that<br />
the two genera might be synonymous. The differences<br />
between the two, so far as can be judged by<br />
the available data, are minor: Baruna socialis has<br />
the carapace wider (1.4 times as wide as long)<br />
than Leipocten sordidulum (1.27-1.33 times); the<br />
exopod of the third maxilliped seems wider in<br />
Baruna than in Leipocten; the posterior anterolateral<br />
tooth of the carapace in Leipocten is granular,<br />
whereas in Baruna it is smooth, having been described<br />
by Stebbing as simple (i.e., not subdivided<br />
into small teeth); the fact that Stebbing's illustration<br />
of the front and the orbit of Baruna socialis<br />
show these without tubercles may be due to the<br />
inaccuracy of the drawing. Should the gonopods<br />
of Baruna and Leipocten prove to be very similar<br />
there would be no good reason not to synonymize<br />
the two names and the older of them, Baruna<br />
Stebbing, 1904, would have to be used.<br />
Baruna socialis Stebbing, 1904, so far the only<br />
species assigned to Baruna, is known only from the
208<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
a d<br />
FIGURE 52.—Leipocten sordidulum Kemp: a, female, dorsal view; b, female, frontal view; c, male<br />
abdomen; d, male chela; e, female chela;/, fourth pereiopod. (From Kemp, 1915, figs. 16, 18,<br />
19, 20a, and pi. 12: fig. 8.)<br />
type-material, which originated from brackish<br />
water of Lake Negombo, Ceylon.<br />
Genus Paracleistostoma De Man, 1895<br />
FIGURE 53<br />
Paracleistostoma De Man, 1895a:580 [type-species: Paracleistostoma<br />
depression De Man, 1895, by selection by Guinot and<br />
Crosnier, 1963:608; gender: neuter].<br />
DIAGNOSIS.—Carapace flat, being only slightly<br />
curved down at margins, lacking ridges, with only<br />
depressed H-shaped groove in center. Surface of<br />
carapace hairy or glabrous, outline quadrangular,<br />
being wider than long. Epigastric ridges indistinct,<br />
placed at base of front. Anterior margin of<br />
front with blunt or triangular projection at either<br />
lateral angle. Anterolateral margin of carapace<br />
lacking teeth. Suborbital ridge forming lower<br />
margin of orbit, true lower margin being barely<br />
visible as row of granules inside orbit. Shape and<br />
sexual dimorphism of chelipeds similar to Cleistostoma.<br />
Walking legs lacking spines on merus.<br />
First somite of male abdomen only slightly wider<br />
than second, failing by far to reach coxae of fifth<br />
pereiopods. Second to fifth somites of male abdomen<br />
fused, although lines separating these somites<br />
only faintly or more distinctly indicated by<br />
sutures. Male abdomen narrowing regularly distally,<br />
showing no constrictions; abdomen pressed<br />
against thorax, completely covering male gonopods.<br />
Latter recurved, strongly and bulbously<br />
widened apically, carrying distal appendage.<br />
REMARKS.—Guinot and Crosnier (1963) listed<br />
the following species as belonging to Paracleistostoma:<br />
P. depressum De Man, 1895 (Figure 53); P.<br />
leachii (Audouin, 1826); P. cristatum De Man,<br />
1895; P. eriophorum Nobili, 1903; P. dentatum Tesch,<br />
1918; P. longimanum Tweedie, 1937; P. fossulum<br />
Barnard, 1955; P. microcheirum Tweedie, 1937; and<br />
P. japonicum Sakai, 1934. In our opinion only the<br />
following species actually belong to Paracleistostoma:<br />
P. depressum, P. longimanum, P. wardi, and P.<br />
dotilliforme (we have examined material of all<br />
except P. longimanum). The last two species (P.<br />
wardi and P. dotilliforme) formerly had been assigned<br />
to Cleistostoma (p. 200). We doubtfully refer<br />
the following two species to Paracleistostoma:<br />
Cleistostoma mcneilli Ward, 1933: This species<br />
was placed by Barnes (1967:246) in Paracleistos-
NUMBER 306 209<br />
FIGURE 53.—Paracleistostoma depression De Man: a, male, dorsal<br />
view; b, male abdomen; c, male chela; d, female chela; e,<br />
gonopods in situ; /, gonopod. (a-d from De Man, 1897, pi.<br />
14: figs. 13, I3d-f; ejfrom Gordon, 1931, fig. 26).<br />
toma. However, the anterolateral margin of the<br />
carapace has distinct teeth, and the male gonopod<br />
figured by Barnes (1967, fig. 16d) differs somewhat<br />
from that of the typical Paracleistostoma species.<br />
It is possible that C. mcneilli is a true Paracleistostoma,<br />
and that the definition of the genus as<br />
given above should be somewhat modified.<br />
Paracleistostoma eriophorum Nobili, 1903: Judging<br />
by Nobili's (1903:23) description of the carapace,<br />
the chelipeds, and the male abdomen, this species<br />
could well be a true Paracleistostoma. However,<br />
nothing is known about the shape of the male<br />
gonopods, while also some important characters<br />
of the abdomen are not mentioned by Nobili. Of<br />
this species, which so far has not been illustrated,<br />
we have not seen any material.<br />
The following five species are excluded from<br />
the genus Paracleistostoma:<br />
Paracleistostoma leachii (Audouin, 1826): The<br />
shape of the male gonopod, which ends in a<br />
narrow apex, shows that the species is not a<br />
d<br />
Paracleistostoma and we make it the type of a new<br />
genus Serenella (p. 211).<br />
Paracleistostoma cristatum De Man, 1895, and P.<br />
japonicum Sakai, 1934, are both placed here in the<br />
new genus Deiratonotus (p. 201).<br />
Paracleistostoma microcheirum Tweedie, 1937, is<br />
made the type of a new genus, Ilyogynnis (p. 206).<br />
Paracleistostoma dentatum Tesch, 1918: The general<br />
shape of the carapace is wholly unlike that of<br />
Paracleistostoma and resembles more that of Camptandrium.<br />
Since the only known specimen is a small<br />
female, very little can be said about the generic<br />
status of this species.<br />
Paracleistostoma fossulum Barnard, 1955: In this<br />
species the dorsal surface of the carapace shows<br />
transverse ridges. The anterolateral margins of<br />
the carapace have some feeble teeth. The third<br />
maxillipeds are separated by a wide gap and the<br />
peduncle of the exopod is fully exposed. The<br />
female chelipeds are larger than those in Paracleistostoma,<br />
have the fingers shorter than the palm<br />
and the cutting edges of both fingers show several<br />
teeth. The species certainly is no Paracleistostoma<br />
and possibly not even an ocypodid. Unfortunately<br />
it is only known from a female specimen.<br />
Genus Paratylodiplax Serene, 1974<br />
FIGURE 54<br />
Paratylodiplax Serene, 1974:62 [type-species: Cleistostoma blephariskios<br />
Stebbing, 1924, by original designation; gender:<br />
feminine].<br />
DIAGNOSIS.—Carapace quadrangular or oval,<br />
wider than long, rather flat, somewhat more convex<br />
in longitudinal than in transverse direction.<br />
Dorsal surface smooth, showing only deep, Hshaped<br />
depression formed by central part of cervical<br />
groove and anterior part of cardiac grooves.<br />
Carapace pubescent or naked. Epigastric ridges<br />
present as rather indistinct rounded elevations at<br />
base of front. Latter somewhat concave in middle.<br />
Anterior margin of front with angularly rounded,<br />
but not produced, anterolateral angles. Inner orbital<br />
angle broadly rounded. Suborbital ridge<br />
lying distinctly below lower margin of orbit,
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 54.—Paratylodiplax derijardi (Guinot and Crosnier): a, carapace of male, dorsal view; b,<br />
male abdomen; c, male chela; d, female chela; e, gonopod. Paratylodiplax edwardsii (MacLeay):<br />
/, abdomen, male, cl 7.8 mm, South Africa; g, gonopod; h, tip of gonopod, enlarged, (a-e from<br />
Guinot and Crosnier, 1963, figs. 1-3, 8, 9; gji from Barnard, 1954, fig. 2e,f).<br />
reaching only slightly farther forward. Anterolateral<br />
teeth of carapace distinct or obscure. Anterolateral<br />
margin continuing into posterolateral; no<br />
additional ridge present posterolaterally on carapace.<br />
Chelipeds equal in female, subequal in<br />
male, showing strong sexual dimorphism, chelipeds<br />
of male being much stronger than those of<br />
female and having different shape. Dactylus of<br />
male cheliped with distinct molariform tooth in<br />
basal part of cutting edge; this tooth absent in<br />
female. Pereiopods lacking spines on merus. First<br />
somite of male abdomen widest of all abdominal<br />
somites, failing by considerable distance to reach<br />
coxae of fifth pereiopods. Second somite of male,<br />
abdomen slightly narrower than first, either free<br />
(P. derijardi, P. blephariskios) or fused with abdom-<br />
h<br />
inal somites 3 to 5. Latter fused, showing only<br />
traces of sutures between somites; sixth somite<br />
and telson free. Somites narrowing gradually<br />
from first to sixth, outline of male abdomen being<br />
bluntly triangular, except for constriction caused<br />
by concavity in lateral margin at level of fifth<br />
somite. Depressed area of thoracic sternum, which<br />
receives abdomen when latter is fully pressed<br />
against thorax, regularly triangular, leaving an<br />
opening between abdomen and thoracic sternite;<br />
through opening part of gonopod visible. Exposed<br />
part of gonopod completely filling gap between<br />
abdomen and sternum, exposed surface lying<br />
flush with lower surface of abdomen. Male gonopod<br />
strongly recurved, ending in 2 rather short<br />
appendages, inner slender, curved or sinuous,
NUMBER 306 211<br />
outer lobiform; both shorter than recurved part<br />
of shaft.<br />
REMARKS.—The following four species are here<br />
assigned to this genus; material of the first three<br />
has been examined by us:<br />
Paratylodiplax edwardsii (MacLeay, 1838:64)<br />
(Figure 54/-A), originally described as Cleistotoma<br />
edwardsii, was included in this genus by Serene<br />
(1974:62). In it the second somite of the male<br />
abdomen is fused with the following three; in our<br />
specimen the suture between the second and third<br />
somites is rather distinct in the right half, but<br />
altogether absent in the left. Barnard's (1950:150)<br />
statement (in his definition of the genus Cleistostoma<br />
in which he placed the present species) that<br />
all somites of the abdomen are free certainly does<br />
not apply here.<br />
Paratylodiplax algoensis (Barnard, 1954:122, figs.<br />
1, 2a-d) originally described as Cleistostoma algoense,<br />
was correctly considered by Barnard to be<br />
close to P. edwardsii. Barnard (1954) described the<br />
chelipeds and the male gonopods, but gave no<br />
description or figure of the entire animal nor of<br />
the male abdomen. Examination of specimens (1<br />
male and 1 female) from Knysna estuary, South<br />
Africa (USNM collection, uncataloged), confirmed<br />
our idea that the species belongs in Paratylodiplax.<br />
It fully agrees with the above-cited<br />
definition of the genus. The male abdomen has<br />
the second somite either free or, if fused with the<br />
third, separated by a deep suture. The fifth somite<br />
is constricted, so that in ventral view the gonopods<br />
show next to the closed abdomen. Serene<br />
(1974:62) believed that this species probably was<br />
conspecific with P. edwardsii.<br />
Paratylodiplax blephariskios (Stebbing, 1924:3, pi.<br />
116), the type-species, was originally described as<br />
Cleistostoma blephariskios. Stebbing's (1924) description<br />
was rather poor, and Barnard (1950:<br />
816, 817) gave some additional details, e.g., of the<br />
chelipeds and the male gonopods, which agree<br />
with those of Paratylodiplax. A rather poorly preserved<br />
specimen at our disposal shows that the<br />
shape of the male abdomen was incorrectly described<br />
and figured by Stebbing as consisting of<br />
seven free somites. The male abdomen shows the<br />
shape typical for the present genus. There is a<br />
distinct suture between the second and third abdominal<br />
somites, but not between the third,<br />
fourth, and fifth somites. The male gonopods are<br />
visible lateral to the closed abdomen. The examined<br />
specimen came from Cape Town, South<br />
Africa, collected by J. Day, 1973, in USNM<br />
collection, uncataloged.<br />
Paratylodiplax derijardi (Guinot and Crosnier,<br />
1963:612, figs. 1-3, 5-11) (Figure bAa-e), was<br />
originally placed in the genus Tylodiplax. The<br />
species is well described and figured in the original<br />
publication, and shows all the characters<br />
listed in the above generic diagnosis of Paratylodiplax,<br />
in which genus we therefore do not hesitate<br />
to place it, following Serene (1974:62).<br />
The present genus differs from Cleistostoma (1)<br />
in the less strong development of the suborbital<br />
ridges, (2) by the fact that the first abdominal<br />
segment of the male does not reach the coxae of<br />
the fifth legs, (3) in showing part of the male<br />
gonopods when the abdomen is pressed against<br />
the thorax, and (4) by the presence of distal<br />
appendages at the end of the male gonopods.<br />
Of the four known species of Paratylodiplax,<br />
three (P. algoensis, P. blephariskios, and P. edwardsii)<br />
are known only from South Africa. The fourth,<br />
P. derijardi, has been reported from Madagascar.<br />
Genus Serenella, new genus<br />
FIGURE 55<br />
TYPE-SPECIES.—Macrophthalmus leachii Audouin,<br />
1826.<br />
ETYMOLOGY.—This genus is named for Dr.<br />
Raoul Serene in recognition of his many valuable<br />
contributions to the study of Brachyura; the gender<br />
of the generic name is feminine.<br />
DIAGNOSIS.—Carapace broadly quadrangular,<br />
being distinctly wider than long. Dorsal surface<br />
of carapace smooth, without ridges or tubercles,<br />
with only shallow grooves indicating regions.<br />
Front curved down, anterior margin at either end<br />
with 2 angular lobes, inner larger than outer and<br />
reaching farther forward. Epigastric ridges dis-
212<br />
FIGURE 55.—Serenella leachu (Audouin), male: a, dorsal view;<br />
b, gonopods in situ; c, tip of gonopod (L, Crust. D. 26887),<br />
Melita Bay, Red Sea; d, abdomen (L, Crust. D. 26888),<br />
Museri Island, Red Sea. (a, from Paulson, 1875, pi. 8: fig. 6;<br />
b, from Gordon, 1931, fig. 27 left.)<br />
tinct, placed before base of front. Lateral margins<br />
of carapace without teeth, converging posteriorly.<br />
Orbits transverse, situated in single straight line.<br />
Upper orbital border convex, lacking incisions.<br />
Eyes with cornea well developed although small.<br />
Antennules transversely folded. Antennae not entering<br />
orbit. Suborbital ridge fused with outer<br />
third of lower orbital margin, provided with granules.<br />
Epistome short, with transverse, smooth,<br />
sharp ridge; posterior margin produced, tonguelike,<br />
in middle between third maxillipeds, latter<br />
filling entire oral field. Merus and ischium covering<br />
part of exopod and entire flagellum; merus<br />
wider than ischium, somewhat auriculate, lacking<br />
lobe near base of carpus. Chelipeds equal, those<br />
of adult males larger than those of females and<br />
with quadrangular tooth on cutting edge of dactylus.<br />
Third pereiopod longer than other legs,<br />
fifth pereiopod shortest. No spines present on any<br />
pereiopod. Male abdomen narrow, not constricted<br />
at fifth abdominal somite. First somite<br />
only slightly wider than second and failing considerably<br />
to reach coxa of fifth pereiopod. Second,<br />
third, and fourth abdominal somites fused, only<br />
some faint grooves indicating borders between<br />
them. Fifth and sixth somites free. Female abdo-<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
men almost circular, with all somites free. Male<br />
gonopods strongly recurved, so much so that apex<br />
overlaps base of shaft. Distal part of gonopod not<br />
widened, ending in narrow, bluntly rounded<br />
apex. Morphological inner side of apex bearing<br />
strong spines, curved at tip and on one side<br />
bearing some granules. Tip of gonopod bearing<br />
short, wide triangular lobe.<br />
REMARKS.—So far S. leachii is the only species<br />
known of this genus. It is possible that Tylodiplax<br />
indica Alcock, 1900, also should be placed here (p.<br />
217), but this species differs in several important<br />
respects from the type-species. In the first place<br />
Alcock's species evidently shows no sexual dimorphism<br />
in the chelipeds, while also the shape<br />
of the third maxilliped and the male gonopod are<br />
different. Stephensen's (1945, fig. 58D) illustration<br />
shows the male abdomen of T. indica slightly<br />
constricted at the fifth somite, but no information<br />
is available on whether or not this constriction is<br />
so strong that the gonopods become visible. A<br />
closer study of Alcock's species is necessary to<br />
solve its generic position.<br />
Genus Telmatothrix, new genus<br />
TYPE-SPECIES.— Telmatothrix powelli, new species.<br />
ETYMOLOGY.—The name is derived from the<br />
Greek words telma (mud of a pool) and thrix (hair)<br />
in allusion to the hairs of the animal, which are<br />
made quite conspicuous by the mud caught between<br />
them; the gender of the generic name is<br />
feminine.<br />
DIAGNOSIS.—Carapace transversely quadrangular,<br />
being wider than long. Surface of carapace<br />
flat, but regions rather distinct, some with elevations<br />
forming blunt transverse ridges. Short, dark<br />
pubescence visible on dorsal surface, especially<br />
dense and conspicuous on ridges. Front curved<br />
down. Epigastric ridges strong, placed on base of<br />
front. Anterolateral margins of carapace with<br />
distinct teeth. Orbits transverse, situated in common<br />
straight line; upper orbital border lacking<br />
incisions. Eyes with cornea well developed. Antennules<br />
obliquely folded. Antennae entering or-
NUMBER 306 213<br />
bit. Lower orbital margin and suborbital ridge<br />
visible in front view, granulated. Epistome short,<br />
with smooth, sharp, transverse ridge. Third maxillipeds<br />
filling entire oral field; merus and ischium<br />
covering palp and part of exopod. Merus longer<br />
than ischium, and with anterolateral angle<br />
rounded or somewhat flattened, not produced,<br />
but with small lobe near base of carpus. Inner<br />
anterior angle of ischium bluntly triangularly<br />
produced. Chelipeds equal; those of adult males<br />
much larger than those of females and with different<br />
shape; dactylus in males with strong molariform<br />
tooth on cutting edge, no such tooth<br />
present on fixed finger of either sex or on dactylus<br />
of female. Third and fourth pereiopods longer<br />
than second and fifth. No spine on any of segment<br />
of pereiopods, but pubescence present on several.<br />
Male abdomen narrow, with second, third, and<br />
fourth somites fused. First somite only slightly<br />
wider than second, largely failing to reach coxae<br />
of fifth pereiopods. Female abdomen almost circular,<br />
consisting of 7 free somites. Male gonopods<br />
strongly recurved, ending in cleft apex, latter<br />
narrowing gradually, lacking distal appendages.<br />
The type and only species so far known of this<br />
genus is described below.<br />
Telmatothrix powelli, new species<br />
FIGURES 56, 57<br />
"a new ocypodid".—Powell, 1976:315.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Nigeria: Mayuku Creek at Ugbekoko (=<br />
Gbekoko), approximately 10 mi [16 km] W of Sapele, Midwest<br />
State, at about 05°54'N, O5°37'E, among mangroves<br />
(Rhizophora), oligohaline, 5 Oct 1975, C. B. Powell, 36, 3 juv<br />
9, 1 juv (L). Same locality, Oct-Dec 1975, C. B. Powell, 3
a<br />
FIGURE 56.—Telmatothrix powelli, new genus, new species: a, male, dorsal view; b, female,<br />
dorsal view. (Drawn by J. Wessendorp.)
NUMBER 306 215<br />
distinctly granular, slightly sinuous, showing no<br />
indentations. Inner half of posterior orbital margin<br />
widened ventrally, with granules in lower<br />
part of widened area. Between widened area and<br />
base of eye short sharp carina present, branching<br />
off from orbital margin. Outer orbital tooth<br />
bluntly angular. Behind this tooth anterolateral<br />
margin of carapace bearing 2 distinct teeth, anteriormost<br />
lower, more rounded than second; latter<br />
longer, more triangular, directed more sideways;<br />
lateral 3 teeth separated from each other<br />
by deep triangular incisions, margins granular.<br />
Posterolateral margin longer than anterolateral,<br />
unarmed but granular; just before reaching posterior<br />
margin row of granules curving inward, not<br />
touching posterior margin. Second granular ridge<br />
present branching off from posterolateral ridge<br />
slightly behind posterior lateral tooth, lying more<br />
medially from posterolateral ridge. This second<br />
ridge, stronger than posterolateral, extending posteriorly,<br />
ending in sharp, elongate posterolateral<br />
tubercle, lying near margin of carapace above<br />
base of fifth pereiopod. Posterolateral margin of<br />
carapace with blunt tooth protruding between<br />
bases of fourth and fifth pereiopods. Posterior<br />
margin of carapace, like posterolateral, somewhat<br />
granular and elevated; another parallel granular<br />
ridge present, placed more anteriorly.<br />
Eyes (Figure 57a) elongate, cornea terminal,<br />
not surrounding stalk completely; lower margin<br />
of orbit finely granular, bearing long hairs that<br />
partly cover eye when latter retracted. Below<br />
lower orbital margin ridge present with about 11<br />
to 16 low but distinct tubercles, outer smaller,<br />
slightly more closely placed than inner.<br />
Antennules folding obliquely, almost transversely.<br />
Antenna entering orbit; flagellum short,<br />
consisting of about 6 segments, bearing long bristle,<br />
slightly longer than flagellum itself.<br />
Front (Figure 57a) touching epistome in middle.<br />
Epistome showing smooth transverse ridge in<br />
middle, posterior margin, strongly produced posteriorly<br />
in middle, distinctly granular.<br />
Third maxillipeds (Figure 57 d) fill entire oral<br />
cavity. In ventral view only part of exopod and<br />
part of palp visible, slender dactylus, somewhat<br />
FIGURE 57.—Telmatothrix powclli, new genus, new species: a,<br />
front; b, male abdomen; c, base of male abdomen in situ; d,<br />
third maxilliped; e, male chela; /, female chela; g, chela of<br />
juvenile male; h, gonopod.<br />
less slender propodus, part of carpus, and distal<br />
part of exopod hidden behind merus and ischium.<br />
Merus with anterolateral angle rounded or somewhat<br />
flattened, but not auricular, forming shallow,<br />
anteriorly directed lobe just outside base of<br />
palp. Latter articulating in middle of anterior<br />
margin of merus. Inner anterior angle of merus<br />
rectangularly rounded. Ischium almost as long as<br />
and as wide as merus. Inner margin of both<br />
segments elevated, rim-like, otherwise no longitudinal<br />
carinae or grooves present.<br />
First legs (Figure 51e-g) showing conspicuous<br />
sexual dimorphism. In adult males chelipeds<br />
large, robust, being far larger than following legs<br />
and more than twice as long as carapace. Fingers<br />
slightly more than half as long as palm, gaping<br />
distinctly. Finger tips slightly spoon-shaped, each
216<br />
bearing row of long hairs on inner margin. Cutting<br />
edges of fingers bearing small granules,<br />
distalmost larger than proximals. Cutting edge of<br />
dactylus with wide molariform tooth, occupying<br />
slightly more than 1/4 of length of edge; no such<br />
tooth present on cutting edge of fixed finger.<br />
Palm highest distally, narrowing proximally,<br />
bearing scattered small granules, densest dorsally.<br />
Carpus about half as long as palm, merus slightly<br />
shorter than palm. Both carpus and merus showing<br />
granules, but bearing no spines. Left and<br />
right chelipeds in male equal in shape and size.<br />
In female chelipeds much shorter than second<br />
pereiopods, of about same length as carapace,<br />
slender and equal. Fingers somewhat longer than<br />
palm, gaping and having spoon-shaped tips bordered<br />
with long hairs. Cutting edges entire. Carpus<br />
about as long as palm and less than 2/3 as<br />
long as merus.<br />
Dactyli of following legs simple, slightly shorter<br />
than propodi, provided on both dorsal and ventral<br />
surfaces with 3 longitudinal pubescent<br />
grooves separated by 2 ridges. Propodi about<br />
twice as long as wide and longer than carpi,<br />
neither bearing any granules. Merus about as<br />
long as propodus and carpus combined but wider,<br />
bearing distinct granules on lower surface, especially<br />
along margins; some granules visible also<br />
in upper part of outer (= posterior) surface. Third<br />
and fourth pereiopods distinctly longer and wider<br />
than second and fifth; third leg largest, fifth<br />
smallest of walking legs. Outer (= posterior) surface<br />
of these legs bearing numerous short black<br />
hairs, among which some longer hairs visible,<br />
especially on upper and lower margins. Third,<br />
fourth, and fifth pereiopods showing numerous<br />
soft, long, woolly hairs on lower surface of merus;<br />
such woolly hairs also visible in outer distal part<br />
of carpus and dorsal part of propodus of third<br />
and fifth legs, and on inner, upper, and lower<br />
surfaces of propodus and distal part of carpus of<br />
fourth legs. Woolly hairs more distinct in males<br />
than in females. No teeth or spines on any of<br />
segments of legs; anterodorsal and anteroventral<br />
angles of merus bluntly rounded.<br />
Male abdomen (Figure 57b,c) slender. First<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
somite only slightly wider than second, largely<br />
failing to reach coxae of fifth pereiopods. Second<br />
somite slightly wider than third, of about same<br />
length as first and about half as long as third.<br />
Third somite about as long as but somewhat<br />
wider than fourth. Second, third, and fourth somites<br />
fused, but grooves indicating lines separating<br />
somites, while deep incisions in lateral margins of<br />
fused part also indicate limits of somites. Distal 3<br />
somites of about equal width. Fifth somite longest<br />
of three, also longer than fourth; differences<br />
slight. Sixth and seventh somites of about equal<br />
length. Sixth quadrangular, seventh with tip<br />
semicircularly rounded. Weak, low transverse carina,<br />
interrupted in middle, present in distal half<br />
of exposed surface of seventh somite. Other somites<br />
not showing any carinae, except first, traversed<br />
by distinct smooth carina. In adult female<br />
abdomen with all somites free, almost semicircular,<br />
reaching to, or overlapping, bases of the<br />
pereiopods.<br />
Male gonopods (Figure 57h) reaching to line<br />
between sternites of second and third pereiopods,<br />
strongly recurved, tapering gradually towards tip.<br />
Tip ending in 2 points, of which 1 minutely<br />
dentate on outer margin. Tip neither widened<br />
nor bearing any appendages. Male sexual openings<br />
placed on sternum.<br />
MEASUREMENTS.—The largest male examined<br />
(the holotype) has the carapace 10 mm long and<br />
14 mm wide, the largest females were of the same<br />
length and 14 and 15 mm wide. Females, with<br />
the abdomen wide and reaching to the bases of<br />
the pereiopods, ranged in length from 7 to 10 mm<br />
and in width from 9 to 15 mm. Ovigerous females<br />
had a carapace length of 9 and 9.5 mm and a<br />
carapace width of 12 and 13 mm. Juveniles (including<br />
females with a narrow abdomen and<br />
males in which the chelae still showed the female<br />
type) ranged from carapace width 2.5 to 8 mm.<br />
The eggs are numerous and have a diameter of<br />
about 0.35 mm, they are spherical.<br />
A male with a carapace length of 6 mm and a<br />
carapace width of 8 mm showed the chelipeds<br />
very similar to those of the females, the fingers<br />
(Figure big) were longer than the palm, with only
NUMBER 306 217<br />
faint traces of the molariform tooth of the dactylus<br />
and of the granulation of the cutting edge<br />
of the fixed finger are visible. In still smaller<br />
males the chelipeds are indistinguishable from<br />
those of the female, e.g., in a male with cl 3.5 mm<br />
and cb 5 mm, in which the abdomen and the<br />
gonopods showed the typical male shape.<br />
TYPE-LOCALITY.—Mayuku Creek at Ugbekoko<br />
(= Gbekoko), approximately 10 miles [16km] west<br />
of Sapele, Midwest State, Nigeria, at 05°54'N,<br />
05°37'E. This locality was described in detail by<br />
Powell (1976:315), who also dealt with the various<br />
animal species found in it.<br />
DISPOSITION OF TYPES.—The holotype (Crust.<br />
D. 31508), a male collected 1-3 November 1975,<br />
forms part of the collection of the Rijksmuseum<br />
van Natuurlijke Historie, Leiden; the greater part<br />
of the paratypes are in the same museum. Series<br />
of paratypes are also deposited in the National<br />
Museum of Natural History, <strong>Smithsonian</strong> <strong>Institution</strong>,<br />
Washington, D.C., and in the Koninklijk<br />
Museum voor Centraal Afrika, Tervuren, Belgium.<br />
ETYMOLOGY—The specific epithet is given in<br />
honor of the collector of the material, C. B.<br />
Powell, University of Port Harcourt, Nigeria, who<br />
by careful collecting activities, and even more by<br />
his investigations, has greatly furthered the<br />
knowledge of <strong>West</strong> <strong>African</strong> fresh and brackish<br />
water Decapoda.<br />
Genus Tylodiplax De Man, 1895<br />
FIGURE 58<br />
Tylodiplax De Man, 1895a:598 [type-species: Tylodiplax tetralylophora<br />
De Man, 1895, by monotypy; gender: feminine].<br />
DIAGNOSIS.—Carapace almost semicircular,<br />
with anterolateral margins widely rounded and<br />
long, provided with 2 indistinct teeth of which<br />
anterior more pronounced than posterior. Posterolateral<br />
margins of carapace short. Dorsal surface<br />
of carapace somewhat convex, with 2 transverse<br />
ridges in anterior half and 4 tubercles in<br />
posterior half. Epigastric ridges not high, placed<br />
on base of front. Anterolateral angles of front<br />
distinct, little produced. Dorsal margin of orbit<br />
showing no incisions. Ischium and merus of third<br />
maxilliped covering part of exopod and part of<br />
palp; merus slightly longer than ischium and not<br />
auriculate, but with small lobe on anterior margin<br />
near base of carpus. Chelipeds of adult male<br />
small, not larger than those of female, and without<br />
teeth on cutting edges. Third and fourth<br />
pereiopods larger than second or fifth. First somite<br />
of male abdomen narrow, not reaching coxae<br />
of fifth pereiopods; second to fourth somites fused,<br />
fifth and sixth free. Abdomen of male not constricted<br />
at fifth somite and in reflexed position<br />
covering gonopods. Male gonopods strongly recurved,<br />
ending in somewhat swollen distal parts,<br />
the one on morphological inner half bearing<br />
broad lobe and that on morphological outer half<br />
several strong and some small spines.<br />
REMARKS.—So far three species have been described<br />
as belonging in the genus Tylodiplax: T.<br />
tetratylophora De Man, 1895; T. indica Alcock,<br />
1900; and 71 derijardi Guinot and Crosnier, 1963.<br />
Some authors also placed Cleistostoma blephariskios<br />
Stebbing, 1924, in this genus. As has been explained<br />
(p. 211), both T. derijardi and C. blephariskios<br />
are assigned to the genus Paratylodiplax, differing<br />
from Tylodiplax in several important characters.<br />
Also Tylodiplax indica has to be removed<br />
from Tylodiplax; it is here provisionally placed in<br />
Serenella (p. 211), but its generic position is still<br />
far from clear, and can only be decided upon<br />
after examination of additional material.<br />
The only species thus remaining in the genus<br />
Tylodiplax is its type-species, T. tetratylophora De<br />
Man, which is known only from Malaya.<br />
Subfamily OCYPODINAE Rafinesque, 1815<br />
Genus Ocypode Weber, 1795<br />
Ocypode Weber, 1795:92 [type-species: Cancer ceratophthalmus<br />
Pallas, 1772, by selection by Holthuis, 1962:244, 245;<br />
gender: feminine; name 1637 on Official List].<br />
Ocypode Fabricius, 1798:312, 347 [invalid junior objective<br />
synonym and homonym of Ocypode Weber, 1795; typespecies:<br />
Cancer ceratophthalmus Pallas, 1772, by selection by<br />
Latreille, 1810:95, 422; gender: feminine; name 1738 on<br />
Official Index].
218<br />
a<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 58—Tylodiplax tetratylophora De Man: a, male, dorsal view; b, male abdomen; c, carapace,<br />
dorsal view; d, third maxilliped; e, gonopod;/, tip of gonopod. (a,b, from De Man, 1897, pi. 14:<br />
fig. 15, 15d; c-f, from Serene and Kumar, 1971, figs. 1, 3-5.)<br />
Ocypoda Lamarck, 1801:149 [incorrect spelling of Ocypode<br />
Weber, 1795; name 1737 on Official Index].<br />
Monolepis Say, 1817:155 [type-species: Monolepis inermis Say,<br />
1817, a subjective junior synonym of Cancer quadratic Fa-<br />
SYNONYM.—Ocypoda hexagonura Hilgendorf,<br />
1882.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
bricius, 1787, by selection by Fowler, 1912:457; gender: Other Material: Liberia: Grand Cape Mount, near Rob-<br />
feminine].<br />
ertsport, 1880-1882, J. Biittikofer and J. H. Sala, 12 speci-<br />
Ceratophthalma MacLeay, 1838:64 [type-species: Cancer cursor mens (L). Monrovia, Apr 1894, O. F. Cook and G. N.<br />
Linnaeus, 1758, by monotypy; gender: feminine]. Collins, 1 juv (W). Ocean beach in front of Camp Johnson,<br />
Parocypoda Neumann, 1878:26 [type-species: Cancer ceratophMonrovia,<br />
G. C. Miller, 21 Jul 1952, \6, 19 (W). Mouth of<br />
thalmus Pallas, 1772, by monotypy; gender: feminine]. Mesurado River, Monrovia, O. F. Cook, 1
NUMBER 306 219<br />
Angola: Musserra, P. Kamerman, lectotype, Id "from<br />
Congo coast" (L, reg. no. D. 235). Musserra, P. Kamerman,<br />
8 specimens (L).<br />
DESCRIPTION.—Capart, 1951:176.<br />
Figures: Monod, 1956, figs. 555-558.<br />
Male Pleopod: Monod, 1956, figs. 557, 558<br />
(Ghana).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths of 8 to 34 mm.<br />
REMARKS.—De Man (1881:253-255), when describing<br />
the present species, based it on a male<br />
(Crust. D. 235) from "Congo" [almost certainly<br />
Musserra, Angola], collected by P. Kamerman.<br />
However, De Man also assigned to his new species<br />
a specimen from Liberia, not seen by him, but<br />
reported upon by Hilgendorf (1869:81), and<br />
noted as ZMB 3118. Both specimens are syntypes<br />
of De Man's species and the one from "Congo"<br />
is now selected to be the lectotype. Monod (1956:<br />
395) already indicated that specimen as the<br />
holotype of Ocypode africana.<br />
BIOLOGY.—Like all species of Ocypode, 0. africana<br />
is an inhabitant of sandy beaches, making its<br />
burrows above the tide line. Gauld and Buchanan<br />
(1956) and Gauld (1960) found the species "very<br />
common in sandy ground above high water<br />
marks, such as coconut groves; juveniles are<br />
found, with O. cursor, below high water mark but<br />
adults rarely so" (Gauld, 1960:71). According to<br />
Gauld and Buchanan (1956:295, 299), 0. africana<br />
occupies a habitat more distant from the sea than<br />
0. cursor with, in certain places, "no evidence of<br />
overlap of the two species . . . . O. africana is quite<br />
terrestrial in habit, emerging from its burrows at<br />
night." Longhurst (1958:53), on the other hand,<br />
found that "in Sierra Leone—at least on the<br />
beaches examined—both species occur together<br />
and excavate burrows above H.W. of springs;<br />
from these, isolated individuals of both species<br />
emerge during the daytime to feed along the lines<br />
of foam left by the surf. At night, the beaches are<br />
crowded with feeding individuals of both species,<br />
which appear to gather—as in the daytime—at<br />
the edge of the surf."<br />
DISTRIBUTION.—<strong>West</strong> Africa, from southern<br />
Mauritania to South-<strong>West</strong> Africa as far as 19°<br />
23'S. Monod (1956) enumerated the records of<br />
the species then known. To these the following<br />
can now be added:<br />
Senegal: Plage de Camberene, N of Dakar and Plage de<br />
Bargny Gouddou (as Bargny), E of Dakar (Sourie, 1957).<br />
Guinea: lies de Los (Uschakov, 1970).<br />
Sierra Leone: No specific locality (Longhurst, 1958).<br />
Liberia: No specific locality (Biittikofer, 1890).<br />
Ghana: No specific locality (Gauld, 1960). Denu and<br />
Labadi (Gauld and Buchanan, 1956).<br />
Nigeria: Lagos (Bruce-Chwatt and Fitz-John, 1951).<br />
Principe: No specific locality (Forest and Guinot, 1966).<br />
Congo: Baie de Pointe-Noire (Rossignol, 1957, 1962).<br />
Zaire: Banana to Vista (Dubois, 1957).<br />
Angola: Lobito (Bott, 1964). Moc.amedes (Guinot and<br />
Ribeiro, 1962).<br />
South-<strong>West</strong> Africa: Rocky Point, 18°59'S, 12°29'E (Penrith<br />
and Kensley, 1970; Kensley, 1970). Near Kunene River<br />
mouth, 17°15'S, 11° 45'E; Dunedin Star wreck site, 18°13'S,<br />
11°56'E; False Cape Frio, 18°29'S, 12°01'E; <strong>West</strong>ies Mine<br />
Camp, 19°12'S, 12°37'E; and Mowe Point, 19° 23'S,<br />
12°42'E (all Kensley, 1970).<br />
* Ocypode cursor (Linnaeus, 1758)<br />
lOcypode rhomba.—Pechiiel-Loesche, 1882:287 [not Ocypode<br />
rhombea Fabricius, 1798].<br />
Ocypode cursor.—Hilgendorf, 1879:802.—Biittikofer, 1890:<br />
465, 487.—Johnston, 1906:862.—Rossignol, 1957:86, pi.<br />
2: fig. 1.—Bott, 1964:31.—Forest and Guinot, 1966:89.—<br />
Voss, 1966:30.—Kensley, 1970:180.—Penrith and Kensley,<br />
1970b:252, 261— Hartmann-Schroder and Hartmann,<br />
1974:13, 23.—Sakai and Tiirkay, 1977:178.<br />
Ocypode ippeus.—Monod, 1933b: 548.<br />
Ocypoda cursor.—Capart, 1951:178, fig. 68.—Monod, 1956:<br />
391, 632, figs. 552-554.—Dubois, 1957:7, fig. 22.—Rossignol,<br />
1957:119 [key].—Sourie, 1957:14, 31, 43, 45.—<br />
Longhurst, 1958:53, 88.—Gauld, 1960:71.—Nicou, 1960:<br />
140.—Guinot and Ribeiro, 1962:65.—Rossignol, 1962:<br />
119—Ribeiro, 1964:14.<br />
Ocypoda hippeus.—Gauld and Buchanan, 1956:295, 296, 298,<br />
301; 1959:127.<br />
Ocypode.—Voss, 1966:52.—Bayer, 1966:98, 102.<br />
SYNONYM.—Ocypode ippeus Olivier, 1804.<br />
MATERIAL EXAMINED.—Pillsbury Material: Nigeria:<br />
Sta 224, Lagos, sand beach, 1
220<br />
north of Dakar, 3 Dec 1975, W. Bohme, 1
NUMBER 306 221<br />
Mesuca Bott, 1973b:316 [type-species: Cancer letragonon Herbst,<br />
1790, by original designation; gender: feminine].<br />
Latuca Bott, 1973b:317 [type-species: Mesuca (Latuca) neocultnmana<br />
Bott, 1973, by original designation; gender: feminine].<br />
Tubuca Bott, 1973b:322 [type-species: Gelasimus urvillei H.<br />
Milne Edwards, 1852, by original designation; gender:<br />
feminine].<br />
Austruca Bott, 1973b:322 [type-species: Gelasimus annulipes H.<br />
Milne Edwards, 1837, by original designation; gender:<br />
feminine].<br />
Paraleptuca Bott, 1973b:322 [type-species: Gelasimus chlorophthalmus<br />
H. Milne Edwards, 1837, by original designation;<br />
gender: feminine].<br />
Heteruca Bott, 1973b:323 [type-species: Gelasimus heteropleurus<br />
Smith, 1870, by original designation and monotypy; gender:<br />
feminine].<br />
Planuca Bott, 1973b:324 [type-species: Uca thayeri Rathbun,<br />
1900, by original designation; gender: feminine].<br />
Leptuca Bott, 1973b:324 [type-species: Gelasimus stenodactylus<br />
H. Milne Edwards and Lucas, 1843, by original designation;<br />
gender: feminine].<br />
Deltuca Crane, 1975:21 [type-species: Gelasimus forcipatus<br />
Adams and White, 1848, by original designation; gender:<br />
feminine].<br />
Australuca Crane, 1975:62 [type-species: Gelasimus bellator<br />
Adams and White, 1848, by original designation; gender:<br />
feminine].<br />
Thalassuca Crane, 1975:75 [type-species: Cancer tetragonon<br />
Herbst, 1790, by original designation; gender: feminine].<br />
Amphiuca Crane, 1975:96 [type-species: Gelasimus chlorophthalmus<br />
H. Milne Edwards, 1837, by original designation;<br />
gender: feminine].<br />
Borboruca Crane, 1975:109 [type-species: Uca thayeri'Rathbun,<br />
1900, by original designation and monotypy; gender:<br />
feminine].<br />
Afruca Crane, 1975:116 [type-species: Gelasimus tangeri<br />
Eydoux, 1835, by original designation and monotypy;<br />
gender: feminine].<br />
Celuca Crane, 1975:211 [type-species: Uca deichmanni Rathbun,<br />
1935, by original designation; gender: feminine].<br />
REMARKS.—Latreille (1817b:5l7), when introducing<br />
the generic name Gelasimus, cited it as<br />
"Gelasimus (Buffon)." This caused Stebbing<br />
(1905:40) to remark: "Latreille at the first institution<br />
of Gelasimus attributed the genus to Buffon,<br />
though in 1820 he claims it as his own. He gave<br />
no reference for the name to any part of Buffon's<br />
works, and no such reference has since been discovered."<br />
Also Crane (1975:20) remarked: "I<br />
have been unable to trace the use of the word by<br />
Buffon." The solution of this problem may be<br />
that Latreille did not intend to refer to George<br />
Louis Leclerc, comte de Buffon, but that the word<br />
"Buffon" is merely cited as a translation of the<br />
latin word "Gelasimus," which means "buffoon,"<br />
"jester," or "mocker."<br />
* Uca tangeri (Eydoux, 1835)<br />
Cancer Uka una, Brasiliensis Seba, 1759:44, pi. 18: fig. 8.<br />
Cancer vocans major Herbst, 1782:83, pi. 1: fig. 11.<br />
Ocypoda heterochelos Lamarck, 1801:150.—Bosc, 1802:197.—<br />
Desmarest, 1830:250.<br />
Cancer Uka Shaw and Nodder, 1803, pi. 588.<br />
Uca una Leach, 1814:430.<br />
Gelasimus Tangeri Eydoux, 1835, pi. 17.<br />
Gelasimus />«7a/uj.—Hilgendorf, 1879:806.—De Man, 1879:<br />
66.—Pechiiel-Loesche, 1882:288.—Biittikofer, 1890:464,<br />
487.—Aurivillius, 1893:31.—Johnston, 1906:862.<br />
Uca tangeri.—Maccagno, 1928:33, fig. 19.—Frade, 1950:11,<br />
26.—Capart, 1951:180, fig. 69.—Monod, 1956:399, figs.<br />
559, 560.—Dubois, 1957:7, fig. 23.—Rossignol, 1957:86,<br />
119 [key].—Sourie, 1957:14, 50.—Longhurst, 1958:53,<br />
88.—Gauld, 1960:71.—Guinot-Dumortier and Dumortier,<br />
1961:136.—Nicou, 1960:135-156, figs. 1-4.—Guinot<br />
and Ribeiro, 1962:67.—Rossignol, 1962:119.—Forest and<br />
Guinot, 1966:90.—Monod, 1967:180, pi. 17: fig. 4.—Uschakov,<br />
1970:448, 455, fig. 4.—Bright and Hogue, 1972:<br />
13.—Bott, 1973a:311-314, figs. 1, 3; 1973b:316, fig. 1.—<br />
Hartmann-Schroder and Hartmann, 1974:19.—Pauly,<br />
1975:57.—Powell, 1979:127.<br />
Gonoplax speciosus Monod, 1933b:548 [footnote; nomen nudum].<br />
Gelasimus (Uca) tangeri.—Bruce-Chwatt and Fitz-John, 1951:<br />
117 [also Gelasimus tangeri, pp. 117 and 119; and Gelasimus<br />
Tangery, p. 119].<br />
Uca tangeri typique Monod and Nicou, 1959:988, figs. 2, 4, 5.<br />
Uca tangeri matandensis Monod and Nicou, 1959:988, figs. 1,<br />
3,6.<br />
Uca (Minuca) tangeri.—Bott, 1968:168.<br />
Uca (Afruca) tangeri.—Crane, 1975:118, figs. 27D-F, 37E,<br />
45E-I, 45EE-II, 46F, 63D, 81E, 82F, 99, pi. 18A-D.<br />
SYNONYMS.—Gelasimus perlatus Herklots, 1851;<br />
Gelasimus cimatodus De Rochebrune, 1883.<br />
MATERIAL EXAMINED—Pillsbury Material: Nigeria: Sta<br />
1, Lagos harbor, shore, 26, 19 (W).<br />
Other Material: Morocco: Tangier, H. Milne Edwards,<br />
?type-material, lc?, 19 (dry, L).<br />
Senegal: Dakar, 1887, J. Biittikofer. 1$ (L). Dakar, 3 May<br />
1882, O. F. Cook, 26, 1? (W).<br />
Liberia: Grand Cape Mount near Robertsport, 1882, J.
222 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Biittikofer, 4(5, 29 (L). Monrovia, Apr 1894, O. F. Cook and<br />
G. N. Collins, New York State Colonization Society, 16* (W).<br />
Rock Spring, Monrovia, Apr 1894, O. F. Cook and G. N.<br />
Collins, 1(5 (W). Mouth of Mesurado River, Monrovia; O.<br />
F. Cook: 3(5 (W). Tive's farm, Bushrod Island, Monrovia, 16<br />
Nov 1953, G. C. Miller, 2(5 (W).<br />
Ghana: Locality not specified, 1840-1855, H. S. Pel,<br />
paralectotypes of Gelasimus perlatus Herklots, 46* (L). Butre<br />
(04°50'N, 01°56'W), 1841-1850, H. S. Pel, types oiGelasimus<br />
perlatus Herklots, 16* lectotype, 3$ paralectotypes (L; lectotype<br />
Crust. D. 262). Elmina, 27 Nov 1889, U. S. Eclipse<br />
Expedition, 36*, 4$ (W). Accra, 1868-1869, M. Sintenis, 6(5,<br />
4$ (L).<br />
Nigeria: Lagos, Apr 1964, F. Kliige, 26*, 1? (L). Tarkwa<br />
Bay, Lagos, Oct 1957, J. Crane, 29(5, 179 (W). S bank of<br />
Escravos River near Ajudaibo, Niger delta, 05°34.5'N,<br />
05° 11.75'E, 20 Jul 1975, C. B. Powell, 26* (L). Wof Forcados<br />
near confluence of Odimodi Creek and Forcados River,<br />
05°22'N, 05°26'E, 28 Feb 1976, C. B. Powell, 15 specimens<br />
(L). Between Brass and Port Harcourt, Niger delta, May-<br />
Aug 1960, H. J. G. Beets, 12 specimens (L).<br />
Cabinda: Chinchoxo (= Quinchoxo, = Tschintschotscho,<br />
05°09.24'S, 12°03.75'E) (see Holthuis and Manning, 1970:<br />
250, 251), 1873-1876, J. Falkenstein, don. Mus. Berlin, 1(5<br />
(W). "Quilla Miindung" (= mouth of the Quila River,<br />
05°58'S, 14°47'E), 1873-1876, J. Falkenstein, don. Mus.<br />
Berlin, 1(5 (W).<br />
Zaire: Congo, Jan 1895, H. C. Kooiman, 19 (L). Banana,<br />
mouth of the Congo River, Aug 1915, H. Lang, American<br />
Museum Congo Expedition, 10(5, 99 (W). Banana, R. I.<br />
Meyer, 26*, 19 (L).<br />
Angola: Musserra, 1882, P. Kamerman, 20 specimens (L).<br />
Santo Antonio do Zaire, mouth of the Congo River, Aug<br />
1915, H. Lang, American Museum Congo Expedition, 1(5<br />
(W). Luanda (= St. Paul do Loanda), 11 Dec 1889, U. S.<br />
Eclipse Expedition, 16* (W). Samba, Luanda, Sep 1957, J.<br />
Crane, 686*, 349, 33 juv (W). Ilha de Cabo, Luanda, Sep<br />
1957, J. Crane, 946*, 989 (W). Luanda, 18 June 1967, G.<br />
Hartmann, 26*, 19 (L). Morro da Cruz, between Luanda and<br />
Cuanza, 23 km S of Luanda, 20 Jun 1967, G. Hartmann,<br />
26", 19 (L). Baia Farta, near Benguela, 3 Jul 1967, G.<br />
Hartmann, 16*, 19 (L).<br />
DESCRIPTION.—Crane, 1975:118-124.<br />
Figures: Capart, 1951, fig. 69; Monod, 1956,<br />
figs. 559, 560; Bott, 1973a, figs. 1, 3; Crane, 1975,<br />
figs. 27D-F, 37E, 45E-1,45EE-II, 46F, 63D, 8IE,<br />
82F, 99, pi. 18A-D.<br />
Male Pleopod: Bott, 1973b, fig. 1 (Zaire); Crane,<br />
1975, fig. 63D (no locality).<br />
MEASUREMENTS.—The examined males have<br />
the carapace width between 13 and 42 mm, the<br />
females between 12 and 28 mm.<br />
REMARKS.—Bott (1973a) very convincingly<br />
showed that the specimen which Seba (1759)<br />
showed on his figure 8 of plate 18 as "Cancer Uka<br />
una, Brasiliensis" belongs to the species that is best<br />
known as Uca tangeri (Eydoux, 1835), and not to<br />
the American species which Rathbun (1918:381)<br />
in her monograph of the American grapsoid crabs<br />
had indicated with the name Uca heterochelos (Lamarck,<br />
1801), Lamarck's species being based on<br />
Seba's figure. Holthuis (1959b:277; 1962:239,<br />
240) rejected the specific epithet heterochelos Lamarck,<br />
1801, and replaced it by its senior objective<br />
synonym major Herbst, 1782; he used the<br />
name Uca major (Herbst) for the American species,<br />
accepting Rathbun's interpretation of the identity<br />
of Ocypoda heterochelos Lamarck, and thus of<br />
Seba's specimen. Holthuis (1962:240) selected the<br />
specimen figured by Seba (1759, pi. 18: fig. 8) as<br />
the lectotype of Cancer vocans major Herbst, 1782,<br />
Ocypoda heterochelos Lamarck, 1801, and Uca una<br />
Leach, 1814, and thereby made these species<br />
objective synonyms of each other. As Uca una<br />
Leach, 1814, is the type-species (by monotypy) of<br />
the genus Uca Leach, 1814, the correct name of<br />
that type-species becomes Uca major (Herbst,<br />
1782).<br />
Bott's (1973a) discovery that the epithets major<br />
Herbst, 1782, heterochelos Lamarck, 1801, and una<br />
Leach, 1814, do not pertain to the American<br />
species (as Rathbun and most later authors<br />
thought), but to the European and <strong>West</strong> <strong>African</strong><br />
Uca tangeri (Eydoux), would cause considerable<br />
confusion in the nomenclature of the group, unless<br />
action by the International Commission on<br />
Zoological Nomenclature prevents this. Bott<br />
(1973a) showed that the oldest available name<br />
for the American species is Gelasimus platydactylus<br />
H. Milne Edwards, 1837, and he suggested that<br />
henceforth this species be known as Uca platydactylus<br />
(H. Milne Edwards, 1837). This change in<br />
itself would not have been disturbing, as the<br />
species is not a very common one and had been<br />
known under several names before 1918 when<br />
Miss Rathbun's monograph stabilized the use of<br />
the name Uca heterochelos for it; not until 1959 was<br />
the epithet major reintroduced for it. However,
NUMBER 306 223<br />
the name Uca major is now generally used for it<br />
and is also adopted in the recent monumental<br />
monograph of the genus Uca published by Crane<br />
(1975:136).<br />
If the change of the name Uca major sensu<br />
Crane to Uca platydactylus (H. Milne Edwards) is<br />
somewhat disturbing, this is very much so for the<br />
other nomenclatural change that results from<br />
Bott's discovery. The species which at present is<br />
best known as Uca tangeri (Eydoux, 1835) under<br />
the strict application of the International Code of<br />
Zoological Nomenclature should have to be given the<br />
name Uca major (Herbst, 1782). This species occurs<br />
on the extreme southwest coast of the Iberian<br />
peninsula and in <strong>West</strong> Africa as far south as<br />
Angola. Since 1835 the epithet tangeri (also spelled<br />
tangieri) has been mostly used for it and since 1900<br />
this usage has been unanimous. Uca tangeri is a<br />
far better known species than U. platydactylus, and,<br />
being the only species of Uca occurring in Europe,<br />
its behavior has been studied by many European<br />
zoologists and the ethological literature concerning<br />
it is extensive (see Crane, 1975:124, for a<br />
listing of this literature). The change of the epithet<br />
tangeri to major would be most undesirable,<br />
the more so as the epithet major at present is<br />
regularly in use for a different species. Also the<br />
fact that Crane adopted the epithet tangeri in her<br />
fundamental monograph, which will be the basis<br />
for all taxonomic and ethological studies of Uca<br />
by generations of biologists to come, is a most<br />
important argument for not changing this name.<br />
Dr. Bott (1973a:313, 314) was of the opinion that<br />
the names Cancer vocans major Herbst, Ocypoda heterochelos<br />
Lamarck, and Uca una Leach could be<br />
considered "nomina oblita" and should be conveniently<br />
ignored. This is not true, however. In<br />
the first place, only Uca una Leach under the Code<br />
before it was revised in 1972 (at the International<br />
Congress of Zoology held in Monaco) could qualify<br />
as a nomen oblitum, but certainly not the<br />
other two names, as those have been repeatedly<br />
used in the last 50 years (be it for a species<br />
different from U. tangeri, which, however, is irrelevant<br />
here). Secondly the present revised Code<br />
does not allow rejection of so-called "nomina<br />
oblita" without a definite action by the International<br />
Commission on Zoological Nomenclature.<br />
As far as we can see, there are two ways open<br />
to save the use of the name Uca tangeri for the<br />
present species:<br />
1. To request the International Commission<br />
on Zoological Nomenclature to use its plenary<br />
powers to suppress the names Cancer vocans major<br />
Herbst, 1782, Ocypoda heterochelos Lamarck, 1801,<br />
Cancer uka Shaw and Nodder, 1803, and Uca una<br />
Leach, 1814. This is a complicated procedure and<br />
is especially inadvisable as the names are those of<br />
the type-species of the genus Uca Leach. The<br />
results of such an action by the Commission<br />
would be that the valid name for the <strong>African</strong><br />
species would remain Uca tangeri, while that of the<br />
American species would become U. platydactylus.<br />
The Commission then also has to decide which of<br />
these two species (or any other) should be made<br />
the type of the genus Uca.<br />
2. To request the International Commission to<br />
indicate under their plenary powers a specimen<br />
of Uca platydactylus (H. Milne Edwards) to be the<br />
neotype specimen of Cancer vocans major Herbst,<br />
1782. In this way the name Uca major (Herbst)<br />
would become the valid name for Uca platydactylus<br />
and the name Uca tangeri would become that of<br />
the <strong>African</strong> species. This action would preserve<br />
the current usage of both names and legalize the<br />
names used by Crane (1975) for the two species.<br />
Furthermore, no change in the name of the<br />
type-species of the genus Uca is then necessary.<br />
The second of these two actions seems to us to<br />
be preferable, and an application has been submitted<br />
on these lines to the International Commission<br />
on Zoological Nomenclature. In it the<br />
Commission is requested to use its plenary powers<br />
to validate the selection of the type-specimen of<br />
Uca platydactylus (H. Milne Edwards) to be the<br />
neotype of Cancer vocans major Herbst, 1782. This<br />
specimen, a full grown male from Cayenne, now<br />
forms part of the collection of the Museum national<br />
d'Histoire naturelle in Paris (see Crane,<br />
1975:601). In case H. Milne Edwards' (1837)<br />
original material of Gelasimus platydactylus consisted<br />
of more than one specimen, the type-spec-
224 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
imen examined by Crane (1975) is made the<br />
lectotype of that species. Of all the specimens of<br />
Uca platydactylus known at present, the neotype of<br />
Cancer vocans major Herbst originates from a locality<br />
(Cayenne) that is closest to the originally<br />
indicated type-locality of Herbst's species, viz.<br />
Brazil. By this neotype selection Cancer vocans major<br />
Herbst, 1782, Ocypoda heterochelos Lamarck, 1801,<br />
Cancer uka Shaw and Nodder, 1803, Uca una Leach,<br />
1814, and Gelasimus platydactylus H. Milne Edwards,<br />
1837, become objective synonyms.<br />
Bott's (1973a) discovery that Seba's specimen<br />
is identical with Uca tangeri also has consequences<br />
for the nomenclature of the subgenera of Uca. In<br />
1968 Bott placed Uca tangeri in the subgenus<br />
Minuca Bott, 1954, but in 1973 he (Bott, 1973b:<br />
316) split the old genus Uca into 10 genera (7<br />
new), one of which consisted of 2 subgenera (both<br />
new). The genus Uca was restricted to two species:<br />
Uca tangeri (Eydoux, 1835) and Uca marionis (Desmarest,<br />
1825) [= Uca vocans (Linnaeus, 1758)]. In<br />
Crane's (1975) monograph of Uca, all the fiddler<br />
crabs were still placed in a single genus, Uca,<br />
within which Crane recognized 9 subgenera (7 of<br />
which were given new names by her). Uca tangeri<br />
was placed by itself in a monotypic subgenus<br />
Afruca Crane, 1975. The limits of Crane's subgenera<br />
in many instances do not coincide at all<br />
with those of Bott's genera and subgenera. It<br />
furthermore is most unfortunate that Bott's<br />
(1973b) preliminary paper was published before<br />
Crane's (1975) monograph, the manuscript of<br />
which was in press too early to take either Bott's<br />
system into consideration or to adopt his new<br />
names. Bott's final revision of the genus Uca sensu<br />
lato was never published, as he died in 1974<br />
before finishing it. We have now the most unpleasant<br />
situation that Crane's subgenera are well<br />
defined and exhaustively treated in an ideal way,<br />
while Bott's names, published in a short, not too<br />
well-documented paper, have priority. Von<br />
Hagen (1976:223), in his review of Crane's monograph,<br />
has already pointed to this unfortunate<br />
nomenclatural situation and stated:<br />
Because of their being objective synonyms Crane's names<br />
Thalassuca, Amphiuca and Borboruca must be replaced by Bott's<br />
Mesuca, Paraleptuca and Planuca, respectively. When accepting<br />
Crane's limits of species groups, one has also to use Tubuca<br />
Bott instead of Deltuca Crane and should use Leptuca (not<br />
Austruca) Bott instead oiCeluca Crane. Only Australuca Crane<br />
and (provided the shift of the type species name in consequence<br />
of Bott, 1973a, is avoidable) Afruca Crane remain<br />
valid.<br />
Unfortunately, in practice (though not in theory) a chaos<br />
of names seems inevitable. Many non-taxonomists, taking<br />
Crane's monograph as the sound reference work that it<br />
actually is, will unsuspiciously use her new but invalid<br />
subgeneric names, while taxonomists would, of course, have<br />
to use Bott's names (regardless, whether on a subgeneric or<br />
generic level).<br />
If the International Code of Zoological Nomenclature<br />
is strictly applied, the name of the present species<br />
would be Uca (Uca) major (Herbst, 1782); if Bott's<br />
system is followed it should be named Uca tangeri<br />
(Eydoux, 1835); and under Crane's system it is<br />
Uca (Afruca) tangeri (Eydoux, 1835). Pending the<br />
decision of the International Commission on Zoological<br />
Nomenclature on this case, the well<br />
known name Uca tangeri (Eydoux) is used here.<br />
We also follow Von Hagen's (1976) suggestion<br />
not to use subgeneric names for the time being.<br />
Another change, in both Crane's (1975) and<br />
Bott's (1973b) nomenclature is needed. If Crane's<br />
system is adopted, then the name Thalassuca<br />
Crane, 1975, should not be replaced by Mesuca<br />
Bott, 1973, as Von Hagen (1976) thought, but by<br />
Gelasimus Latreille, 1817, because the type-species<br />
of Gelasimus, Cancer vocans Linnaeus, 1758, belongs<br />
in Crane's Thalassuca (see Crane, 1975:20). Under<br />
Bott's classification Gelasimus Latreille, 1817, must<br />
fall as a junior synonym of Uca Leach, 1814, and<br />
for the genus that Bott (1973b:323) indicated<br />
with the name Gelasimus, the generic name Acanthoplax<br />
H. Milne Edwards, 1852, would become<br />
available.<br />
DISTRIBUTION.—The species is found in the<br />
eastern Atlantic, its range extending from the<br />
south coast of Portugal to southern Angola; it<br />
does not occur in the Mediterranean. The records<br />
from America (<strong>West</strong> Indies, Brazil; cf. Rathbun,<br />
1918:388 and Bott, 1973a) in all likelihood are<br />
erroneous, due to incorrect labeling. Monod<br />
(1956) enumerated the localities from which the
NUMBER 306 225<br />
species was then known. Since that time it has<br />
been reported from the following <strong>West</strong> <strong>African</strong><br />
localities (including those overlooked by Monod;<br />
Crane's (1975) records are not duplicated here):<br />
Senegal: Saloum (Sourie, 1957). Ngor, near Dakar (Nicou,<br />
1960).<br />
Guinea-Bissau: Bissau (Maccagno, 1928; Frade, 1950).<br />
Guinea: lie Marara, mouth of Rio Pongo (Uschakov,<br />
1970). Boulbinet, Tanene, and Ralompa, all near Conakry<br />
(Monod and Nicou, 1959).<br />
Liberia: No specific locality (Hilgendorf, 1879; Biittikofer,<br />
1890; Johnston, 1906).<br />
Sierra Leone: Robene Point and Cline Bay (as Kline)<br />
(Longhurst, 1958).<br />
Ivory Coast: Lagoon of Abidjan, 05°16'12"N, 04°00'-<br />
20"W (Forest and Guinot, 1966).<br />
Ghana: "Coast of Guinea" (De Man, 1879). Estuaries of<br />
Volta and Ankobra rivers (Gauld, 1960). Sakumo Lagoon<br />
(Pauly, 1975).<br />
Nigeria: Lagos (Bruce-Chwatt and Fitz-John, 1951). Elechi<br />
Creek, Port Harcourt, 04°47'15"N, 06°58'45"E (Powell,<br />
1979).<br />
Cameroon: No specific locality (Aurivillius, 1893). Douala<br />
(Monod and Nicou, 1959).<br />
Principe: No specific locality (Maccagno, 1928; Forest<br />
and Guinot, 1966). Praia Ponta da Mina and Santo Antonio<br />
(Forest and Guinot, 1966).<br />
Sao Tome: No specific locality (Maccagno, 1928; Forest<br />
and Guinot, 1966).<br />
Congo: No specific locality (Maccagno, 1928). Loango<br />
(Pechiiel-Loesche, 1882; Rossignol, 1957). Mouth of Songololo<br />
River (Rossignol, 1957, 1962). Djeno (Rossignol, 1957).<br />
Cabinda: Chinchoxo (Hilgendorf, 1879).<br />
Zaire: Banana (Dubois, 1957; Bott, 1973a,b).<br />
Angola: Luanda (Hilgendorf, 1879; Bott, 1968). Baia de<br />
Luanda, Lobito; Baia Farta, Benguela; and Baia dos Tigres<br />
(Guinot and Ribeiro, 1962). Between Cacuaco and Lobito-<br />
Benguela (Hartmann-Schroder and Hartmann, 1974).<br />
Family GRAPSIDAE MacLeay, 1838<br />
GRAPSIDAE MacLeay, 1838:63, 65.<br />
SESARMINAE Dana, 1851c:288.<br />
PLAGUSINAE Dana, 1851c:288 [corrected to Plagusiinae by<br />
Miers, 1878:147; name 377 on Official List].<br />
VARUNACEA H. Milne Edwards, 1853:175 [corrected to Varuninae<br />
by Alcock, 1900:288, 400].<br />
CYCLOGRAPSACEA H. Milne Edwards, 1853:191.<br />
EASTERN ATLANTIC GENERA.—Thirteen, 12 of<br />
which are represented by species occurring from<br />
localities between Mauritania and Angola. The<br />
other genus is: Eriocheir de Haan (1835:32), typespecies:<br />
Grapsus (Eriocheir) japonicus de Haan,<br />
1835, by selection by H. Milne Edwards (1854:<br />
146); gender: masculine.<br />
EASTERN ATLANTIC SPECIES.—Twenty-five, 19<br />
of which occur in tropical waters (the included<br />
Atlantic species of Brachynotus is not known to<br />
occur south of Mauritania so is not strictly tropical).<br />
The other species are as follows:<br />
Brachynotus foresti Zariquiey Alvarez, 1968.<br />
Mediterranean; littoral (Zariquiey Alvarez, 1968;<br />
Froglia and Manning, 1978).<br />
Brachynotus gemmellari (Rizza, 1839). Mediterranean;<br />
sublittoral (Froglia and Manning, 1978).<br />
Brachynotus sexdentatus (Risso, 1827). Mediterranean;<br />
introduced into Great Britain in artificially<br />
heated water at Swansea; northern part of<br />
Suez Canal and Black Sea (Zariquiey Alvarez,<br />
1968; Froglia and Manning, 1978).<br />
Eriocheir sinensis H. Milne Edwards, 1853. A<br />
native of China, introduced into NW Europe and<br />
now reported from Finland, Sweden, and Norway<br />
to England and the Atlantic coast of France, in<br />
rivers (Christiansen, 1969); and from Lake Erie,<br />
North America (Nepszy and Leach, 1973).<br />
Pachygrapsus marmoratus (Fabricius, 1787). Mediterranean<br />
and Atlantic coast of Europe from<br />
Brittany (France) southward to Morocco, including<br />
Madeira, the Azores, and the Canary Islands;<br />
intertidal (Zariquiey Alvarez, 1968).<br />
Pachygrapsus maurus (Lucas, 1846). <strong>West</strong>ern<br />
Mediterranean, Madeira, the Azores and the Canary<br />
Islands; intertidal (Zariquiey Alvarez, 1968).<br />
Four species of Grapsidae at one time or another<br />
reported from <strong>West</strong> Africa in all probability<br />
do not belong to the <strong>West</strong> <strong>African</strong> fauna, and the<br />
material on which these records are based either<br />
is incorrectly identified or is labeled with a wrong<br />
locality. These species are as follows:<br />
Metopograpsus messor (Forskal, 1775). A species<br />
known with certainty from the Red Sea, the east<br />
coast of Africa, Madagascar and the Persian Gulf<br />
(Holthuis, 1977a). Monod (1956:422, 423) listed<br />
the records of this species from <strong>West</strong> Africa and<br />
indicated that most probably they were based on<br />
incorrect identifications. There was, however, a
226<br />
correctly identified specimen of Metopograpsus messor<br />
in the collection of the Museum national<br />
d'Histoire naturelle, Paris, labeled "Gabon," the<br />
correctness of which was doubted by Monod as<br />
this same lot included a specimen of another<br />
species, Pseudograpsus elongatus (see below), which<br />
also occurs only in the Indo-<strong>West</strong> Pacific region.<br />
Platychirograpsus spectabilis De Man, 1896. This<br />
species was originally described by De Man<br />
(1896:292, fig. 1) from Gabon. Apart from the<br />
types of this very characteristic species, no material<br />
has ever been reported from <strong>West</strong> Africa,<br />
whereas it has been found repeatedly in fresh<br />
water on the E coast of Mexico and also has been<br />
introduced into Florida (Marchand, 1946:93-<br />
100). As pointed out by Monod (1956:426-428)<br />
there is a definite possibility that De Man's material<br />
did not originate from <strong>West</strong> Africa but from<br />
Mexico.<br />
Pseudograpsus elongatus (A. Milne Edwards,<br />
1873). A well known species from the Indo-<strong>West</strong><br />
Pacific (Red Sea, E Africa, Madagascar, the Seychelles,<br />
and New Caledonia) (Holthuis, 1977a),<br />
but not known from <strong>West</strong> Africa other than from<br />
the specimen labeled as originating from Gabon<br />
and found together with Metopograpsus messor (see<br />
above).<br />
Sesarma roberti H. Milne Edwards, 1853. Originally<br />
described from Goree, Senegal, but otherwise<br />
only reported from the <strong>West</strong> Indies and the<br />
Atlantic coasts of Central and South America. As<br />
stated by Monod (1956:443) and Chace and<br />
Hobbs (1969:184), the type-locality indication is<br />
very likely erroneous, so that this species, as well<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Subfamily GRAPSINAE MacLeay, 1838<br />
Genus Geograpsus Stimpson, 1858<br />
Geogmpsus Stimpson, 1858b: 101 [p. 47 on separate] [typespecies:<br />
Grapsus lividus H. Milne Edwards, 1837, by subsequent<br />
designation by Rathbun, 1918:231; gender: masculine].<br />
Orthograpsus Kingsley, 1880a: 188, 194 [type-species: Orthograpsus<br />
hillii Kingsley, 1880, a subjective junior synonym<br />
of Grapsus lividus H. Milne Edwards, 1837, by present<br />
selection; gender: masculine].<br />
* Geograpsus lividus (H. Milne Edwards, 1837)<br />
Geograpsus livtdus.—Monod, 1956:410, figs. 562, 563.—Dubois,<br />
1957:7.—Guinot and Ribeiro, 1962:69.—Ribeiro,<br />
1964:15.—Forest and Guinot, 1966:91.<br />
SYNONYMS.—Grapsus brevipes H. Milne Edwards,<br />
1853; Orthograpsus hillii Kingsley, 1880.<br />
MATERIAL EXAMINED.—Pillsbury Material: Nigeria:<br />
Sta 1, Lagos harbor, shore, 2$ (L).<br />
Other Material: Zaire: Banana, mouth of the Congo River,<br />
Aug 1915, H. Lang, American Museum Congo Expedition,<br />
2c5, 2$ (W).<br />
Angola: Lobito, P. Kamerman, 1$ (L).<br />
DESCRIPTION.—Rathbun, 1921:442; Chace and<br />
Hobbs, 1969:157.<br />
Figures: Rathbun, 1921, pi. 15: fig. 1, pi. 22:<br />
figs. 2, 3; Monod, 1956, figs. 562, 563; Chace and<br />
Hobbs, 1969, fig. 48.<br />
Male Pleopod: Monod, 1956, fig. 563 (Senegal);<br />
Chace and Hobbs, 1969, fig. 52a-c (<strong>West</strong> Indies).<br />
MEASUREMENTS.—The females collected by the<br />
Pillsbury have the carapace width 12 and 13 mm.<br />
DISTRIBUTION.—The species inhabits the tropi-<br />
as Metopograpsus messor, Pseudograpsus elongatus, and cal Atlantic, and has also been reported from the<br />
Platychirograpsus spectabilis, had best be removed eastern Pacific. In the Indo-<strong>West</strong> Pacific area it<br />
from the list of the <strong>West</strong> <strong>African</strong> Brachyura. is represented by the closely related Geograpsus<br />
Other than the subgeneric names of the <strong>West</strong><br />
stormi De Man. In <strong>West</strong> Africa the species is<br />
<strong>African</strong> Sesarma , our resurrection of Goniopsis pelii<br />
known from the Cape Verde Islands and Senegal<br />
to Angola and Sao Tome. Monod (1956) listed<br />
from the synonymy of Goniopsis cruentata, and our<br />
all <strong>West</strong> <strong>African</strong> records known to him, to which<br />
use of Metagrapsus instead of Sarmatium for S.<br />
now the following can be added:<br />
curvatum, the grapsid names used by Monod<br />
(1956) have not changed so we have not listed Cape Verde Islands: Bafa das Gatas, Sao Vicente (Guinot<br />
them separately for this family.<br />
and Ribeiro, 1962; Ribeiro, 1964). Sao Tiago (Forest and<br />
Guinot, 1966).
NUMBER 306 227<br />
Fernando Poo: No specific locality (Forest and Guinot,<br />
1966).<br />
Annobon: No specific locality (Forest and Guinot, 1966).<br />
Zaire: No specific locality (as Belgian Congo) (Dubois,<br />
1957).<br />
Angola: Baia Farta, Benguela; Baia da Caota, Benguela;<br />
Baia de Santa Maria; and Praia Amelia, Mocamedes (all<br />
Guinot and Ribeiro, 1962).<br />
Genus Goniopsis de Haan, 1833<br />
Goniopsis de Haan, 1833:5 [genus established without included<br />
nominal species; the first nominal species to be<br />
assigned to the genus, by de Haan (1835:33), are Cancer<br />
strigosus Herbst, 1799, Grapsus pictus Latreille, 1803, and<br />
Grapsus cruentatus Latreille, 1803; type-species: Grapsus<br />
cruentatus Latreille, 1803, by selection by Rathbun, 1918:<br />
236; gender: feminine].<br />
* Goniopsis pelii (Herklots, 1851)<br />
FIGURE 59<br />
Grapsus (Grapsus) Pelii Herklots, 1851:8, 23, pi. 1: figs. 6, 7.<br />
Grapsus (Grapsus) simplex Herklots, 1851:9, 23, pi. 1: fig. 8.<br />
Grapsus (Goniopsis) cruentatus.—Won Martens, 1872:105, 106<br />
[not Grapsus cruentatus Latreille, 1803].<br />
Grapsus Pelii.—De Man, 1879:68; 1900, pi. 2: fig. 6 [Peli on<br />
pp. 43-46, 64].<br />
Grapsus simplex.—Be Man, 1879:68.—Miers, 1886:255.—De<br />
Man, 1900:43, pi. 2: fig. 7.<br />
Goniopsis cruentatus.—Kingsley, 1880a: 190.—Studer, 1882:<br />
333 [listed].—Biittikofer, 1890:487.—Thallwitz, 1891:<br />
52.—Benedict, 1893:538.—Doflein, 1900:142.—Johnston,<br />
1906:862.—Vilela, 1949:63.—Frade, 1950:11, 26 [not<br />
Grapsus cruentatus Latreille, 1803].<br />
Grapsus pelii.—Kingsley, 1880a: 190 [in synonymy].<br />
Pachygrapsus simplex.—Kingsley, 1880a:201.<br />
Goniograpsus cruentatus.—Osorio, 1887:227; 1888:191; 1889:<br />
130, 134, 139; 1890:46; 1892:199; 1895a:54; 1895b:57, 58;<br />
1898:193 [not Grapsus cruentatus Latreille, 1803].<br />
Grapsus Peli.—Dc Man, 1900:43, 64, pi. 2: fig. 6 [Pelii on<br />
plate].<br />
Goniopsis cruentata.—Rathbun, 1900a:278.—Nobili, 1906b:<br />
311.—Rathbun, 1918:237; 1921:443, pi. 39.—Balss, 1922:<br />
80.—Monod, 1927:620; 1928:124.—Irvine, 1947:292, fig.<br />
197.—Bruce-Chwatt and Fitz-John, 1951:117.—Capart,<br />
1951:183, fig. 7.—Sourie, 1954a:84, 112, 293, 297, 306.—<br />
Monod, 1956:412, figs. 564-567.—Rossignol, 1957:89,<br />
121.—Longhurst, 1958:88.—Gauld, 1960:71.—Rossignol,<br />
1962:119.—Forest and Guinot, 1966:91—Uschakov,<br />
1970:443, 455 [listed].—Pauly, 1975:57. [Not Grapsus cruentatus<br />
Latreille, 1803.]<br />
Gonopsis cruentatus.—J. Roux, 1927:237 [not Grapsus cruentatus<br />
Latreille, 1803].<br />
Goniopsis cruenta.—Hartmann-Schroder and Hartmann,<br />
1974:19 [erroneous spelling].<br />
Goniopsis pelii.—Powell, 1979:127.<br />
Not Goniograpsus cruentatus.—Osorio, 1892:199 [=Grapsus grapsus<br />
(Linnaeus), see Forest and Guinot, 1966:90].<br />
Not Pachygrapsus simplex.—Doflein, 1904:129.—Lenz, 1910:<br />
125.—Lenz and Strunck, 1914:283 [= P. gracilis (Saussure),<br />
see Balss, 1922:81].<br />
MATERIAL EXAMINED.—Pillsbury Material: Nigeria:<br />
Sta 1, Lagos harbor, shore, 2
228 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
specimens varied from 16 to 49 mm. There is no<br />
obvious size difference in males and females.<br />
Ovigerous females have carapace widths between<br />
25 and 47 mm. The male lectotype of G. pelii has<br />
a carapace width of 48 mm, the male lectotype of<br />
G. simplex one of 20 mm.<br />
REMARKS.—A comparison of the present <strong>West</strong><br />
<strong>African</strong> material with specimens of Goniopsis cruentata<br />
from the north coast of South America and<br />
the Caribbean shows some differences, which,<br />
although small, seem to be constant and induce<br />
us to distinguish the two populations as distinct<br />
species. A closer study based on much more material<br />
and taking also the <strong>West</strong> American G.<br />
pulchra into account may establish that the three<br />
should be considered subspecies of a single species.<br />
Goniopsis pelii differs from G. cruentata in the<br />
following points.<br />
1. The front is relatively shorter and wider.<br />
The anterior margin of the front not only is<br />
emarginate in the middle, but in the middle of<br />
each half as well; this secondary emargination is<br />
less distinct in G. cruentata. The surface of the front<br />
bears coarse tubercles, some of which, near the<br />
anterior margin, form more or less distinct transverse<br />
ridge-like rows; in G. cruentata such transverse<br />
ridges, if present, are usually short and<br />
inconspicuous. This latter character, which was<br />
already described by Von Martens (1872), however,<br />
is not very constant.<br />
2. The pereiopods in G. pelii have the propodi<br />
more slender than in G. cruentata, and the dactyli<br />
are more robust. In G. pelii the length of the<br />
dactylus of the last pereiopod (Figure 59b) is<br />
distinctly less than that of the lower margin of<br />
the propodus, while in G. cruentata (Figure 59
NUMBER 306 229<br />
FIGURE 59.—Goniopsispelii (Herklots), ovigerous female, Cameroon: a, dorsal view; b, propodus<br />
and dactylus of fifth pereiopod. Goniopsis cruentata (Latreille): c, propodus and dactylus of fifth<br />
pereiopod.<br />
the exception of the third maxillipeds, which are<br />
pale with a dark purple spot on the carpus and in<br />
the distal part of the merus. The distal four<br />
somites of the abdomen are whitish; the basal<br />
three are pale purple.<br />
The specimens from Nigeria in which the color<br />
is well preserved show the same color pattern,<br />
although the color itself is more reddish. Also the<br />
Liberian material agrees well with this description.<br />
In most other specimens the color has faded,<br />
but in a few some traces of the original pattern<br />
are visible, agreeing, as far as can be ascertained,<br />
with the above description.<br />
The color pattern of the <strong>West</strong> <strong>African</strong> speci-<br />
mens differs strikingly from that of G. cruentata, of<br />
which an excellent description is provided by<br />
Chace and Hobbs (1969:161), who also provided<br />
a beautiful figure of the color pattern. In Goniopsis<br />
cruentata (1) the carapace is lighter, instead of<br />
having small light spots on a purple background,<br />
it shows small purple or red spots on a pale<br />
background; (2) instead of having a pale streak<br />
along the posterolateral margin, it shows in the<br />
posterolateral area a number of characteristic<br />
white spots that are surrounded by a dark purple<br />
or red ring; (3) the eyestalks are marbled with<br />
purple or red instead of being uniformly colored;<br />
(4) the carpus of the cheliped is "scarlet with
230<br />
purple lines" (Chace and Hobbs, 1969:161) and<br />
does not show the bright white tubercles of G.<br />
pelii; (6) the merus of the chelipeds is of a uniform<br />
red color inside or orange red with darker marblings<br />
and shows on the outer surface conspicuous<br />
large pale spots surrounded by a dark red ring;<br />
(7) the pereiopods are paler than in G. pelii, the<br />
upper margins of merus, carpus, and propodus<br />
are blocked with dark and light color; the dactylus<br />
is not conspicuously paler than the propodus.<br />
The merus has conspicuous white spots with<br />
dark rings around them.<br />
Only very few authors give some information<br />
on the color of the present species. Herklots (1851:<br />
8) described the carapace as "ruber, viridi flavo<br />
marmoratus." Von Martens (1872:106) when<br />
dealing with a specimen from Liberia observed<br />
that the coloration of the carapace was more<br />
"kleinfleckig" than in his American material. De<br />
Man (1900:43) stated that the types of Grapsus<br />
pelii at that time had still "une belle couleur<br />
rouge jaunatre, moins intense sur les pattes anterieures."<br />
Rathbun (1921:443) published the following<br />
color description of a living animal "A<br />
very showy crab, distinguished by its colors; carapace<br />
brownish yellow or brick-red; legs red, with<br />
spots of a darker red, extremities yellow. Chelipeds<br />
red, except the palms, which are almost<br />
white, and the fingers, which are yellow." Irvine<br />
(1947:292) described the colors as follows: "The<br />
carapace is dark reddish black. . . . The eyes are<br />
black. The claws are pale, with yellowish tips,<br />
their upper segments being of a beautiful purple<br />
colour. The legs are dark red in colour and are<br />
flat and covered with long stiff bristles."<br />
Capart (1951:183) gave the following account<br />
"Couleurs vives, carapace brun-rouge; pattes<br />
rouges, de meme que les chelipedes, sauf les<br />
pinces, qui sont jaune clair." Unfortunately most<br />
of these authors paid more attention to the colors<br />
than to the color pattern of their animals, while<br />
the latter is far more important. The color description<br />
provided by Rossignol (1957:89) is far<br />
superior to most others. It is as follows:<br />
Face dorsale: Carapace noire avec des points jaunes tres<br />
petits et tres serres, sauf sur les regions branchiales. Une<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
ligne mediane courte de couleur jaune sur la partie posterieure<br />
de la carapace (ligne continuee par une serie de points<br />
de meme couleur sur chaque segment abdominal). Une ligne<br />
blanche sur chaque bord lateral. Deux taches rouges entre la<br />
region gastrique et la region cardiaque.<br />
Pinces: face externe jaune clair a vert amande. Face<br />
interne: violacee. Sur le bord superieur de la main et du<br />
doigt, une serie de tubercules blancs ou jaunatres.<br />
Pattes: brun fonce, noir, avec une tache rouge a chaque<br />
articulation. Dactyle rouge.<br />
Face ventrale: jaunatre. Les deux premiers segments abdominaux<br />
violaces. Mxp. 3 blancs, tranchant sur le brun<br />
sale des regions pterygostomiales. [The species is also] tres<br />
facilement reconnaissable avec ses tubercules blancs sur le<br />
bord superieur des pinces.<br />
This description, obviously made after living<br />
specimens, agrees excellently with the one we<br />
made (quite independently from Rossignol's account,<br />
with which we were not aware at that<br />
time) after preserved specimens that had kept<br />
their coloration quite well. An illustration of our<br />
specimen is provided here to show the color pattern<br />
(Figure 59a).<br />
It is interesting to note that Dana (1855, atlas,<br />
pi. 21: fig. 7) published a colored figure ofGoniopsis<br />
cruentata from Brazil that shows a color pattern<br />
different from that of the <strong>West</strong> Indian form and<br />
in a few respects resembles that of G. pelii (presence<br />
of a white streak along the posterolateral<br />
margin of the carapace; pale dactyli of the pereiopods);<br />
however, it differs conspicuously from<br />
both types, and a more thorough study of Brazilian<br />
material of Goniopsis might yield interesting<br />
results.<br />
Herklots (1851:8, 9, pi. 1: figs. 6, 7, 8) described<br />
the present species under two different names,<br />
viz., Grapsus pelii and G. simplex. As pointed out<br />
by De Man (1900:43, pi. 2, figs. 6, 7), G. simplex<br />
is based on juvenile specimens of G. pelii. The<br />
types of both species are still extant in the Rijksmuseum<br />
van Natuurlijke Historie, Leiden, and<br />
after examining them we can fully confirm De<br />
Man's conclusions. We use here the specific name<br />
pelii in preference to simplex, as the former is based<br />
on fully adult, the latter on juvenile specimens.<br />
The two names are of the same data, and under<br />
the Law of Priority have equal rights. In an<br />
earlier paper De Man (1879:68) synonymized
NUMBER 306 231<br />
Grapsus pelii with Goniopsis cruentata and was folcommon<br />
among stones and debris on the ground<br />
lowed in this by all subsequent authors. The (Rathbun, 1921:444; Uschakov, 1970:443), and<br />
specimens brought by Doflein (1904:129), Lenz Sourie (1954a:84, 293) mentioned that near<br />
(1910:125), and Lenz and Strunck (1914:283) to Dakar the species lives "dans les anfractuosites<br />
Pachygrapsus simplex, as Balss (1922:81) has des maconneries; abondant au voisinage des man-<br />
pointed out, are not Herklots' species, but Pachygroves," and "commun sur les falaises" near Congrapsus<br />
gracilis (De Saussure).<br />
akry. Pauly (1975:57) noted that it was not com-<br />
Forest and Guinot (1966:91) reported on a mon near the mouth of the Sakumo Lagoon,<br />
specimen of Geograpsus lividus from Fernando Poo Ghana.<br />
in the collection of the Museu Bocage, which was This species has been reported to live in bur-<br />
identified as "Goniograpsus cruentatus"; it is possible rows that it makes in the mud of mangrove areas.<br />
that this is one of the specimens reported by It is commonly seen climbing on the aerial roots<br />
Osorio (1895b:57) from that island. The same<br />
of mangrove trees and also on slender vertical<br />
authors also showed that the material that Osorio<br />
stems, up to 5 feet [1.5 m] over the surface<br />
(1892:199) reported as Goniograpsus cruentata from<br />
(Rathbun, 1921). Irvine (1947:293) described this<br />
Binda, Sao Tome, actually belongs to Grapsus<br />
as follows:<br />
grapsus. On the other hand, they could confirm<br />
Osorio's (1889:139) identification of this species It is often seen climbing up an aerial root of the red<br />
from Praia Salgada, Principe.<br />
BIOLOGY.—The Pillsbury specimens were taken<br />
on a muddy bottom near the sea wall of Lagos<br />
harbor in brackish water. Most records in the<br />
mangrove with four legs on either side of it. The claws are<br />
kept underneath as it climbs, and are used as legs, working<br />
alternately in pushing the animal up the aerial roots. When<br />
pursued it can climb rapidly, and if closely pressed it often<br />
drops into the water and escapes along submerged roots.<br />
literature indicate the species from a similar<br />
brackish muddy habitat. The best characterization<br />
is that given by Rathbun (1921:444):<br />
They are very agile and when disturbed seek<br />
shelter between mangrove roots, or under stones<br />
and debris (Rathbun, 1921). They feed on the<br />
These crabs avoid the seashore, and on finding them in<br />
good numbers in mangrove swamps one might at first<br />
consider these their favorite habitat. However, they are not<br />
found in any of these swamps far inland for they remain<br />
near the mouth of rivers, where the salinity of the water is<br />
ground. Irvine (1947:293) also remarked that "it<br />
appears to feed on the bark of dead roots and<br />
other rotting vegetation, but may live on minute<br />
animals living in the bark. When feeding it brings<br />
less than in the sea but still greater than about the creeks its claws up to its mouth alternately in a most<br />
up-river where mangroves are still able to flourish. amusing manner."<br />
Irvine (1947:292, 293) described the species as<br />
living "mainly in the wet mud underneath the<br />
tangled mass of aerial prop roots of the red mangrove<br />
(Rhizophora). It occupies small holes hollowed<br />
out in the mud and is often to be seen<br />
crawling on the surface." Furthermore, he stated<br />
that they "are also seen to a smaller extent in the<br />
Humes (1957) reported that this species was<br />
the host of the harpacticoid copepod Cancrincola<br />
longiseta Humes at several localities on the <strong>West</strong><br />
<strong>African</strong> coast between Sierra Leone and the<br />
Congo.<br />
Ovigerous females have been reported during<br />
April, May, July, August, and November.<br />
slightly drier swamps round the banks of the The species seems to be of very little economic<br />
lagoons, where the land is covered with a growth value. Only Irvine (1947:293) remarked that it is<br />
of white mangrove (Avicennia nitida)." Several sold in the fish markets in Ghana.<br />
other authors also reported the species from man- VERNACULAR NAME.—Both Irvine (1947) and<br />
groves (e.g., Monod, 1927, 1956; Capart, 1951; Bruce-Chwatt and Fitz-John (1951) give this spe-<br />
Longhurst, 1958; Gauld, 1960; Rossignol, 1957, cies the vernacular name "Mangrove crab," a<br />
1962). However, Goniopsis pelii also is found to be name also found on the field label of the speci-
232<br />
mens from the mouth of the Junk River, Liberia,<br />
examined by us.<br />
DISTRIBUTION.—Goniopsis pelii is known from<br />
the <strong>West</strong> <strong>African</strong> coast from Dakar, Senegal, in<br />
the north to Mocamedes, Angola, in the south.<br />
Monod (1956:414, 415) listed many localities in<br />
Senegal, Guinea-Bissau, Guinea, Sierra Leone,<br />
Liberia, Ivory Coast, Ghana, Nigeria, Rio Muni,<br />
Gabon, and Congo, whence he had examined<br />
material of the present species. The other localities<br />
mentioned in the literature are as follows.<br />
Senegal: Dakar (Rathbun, 1900a, 1918; Sourie, 1954a).<br />
Casamance (Sourie, 1954a).<br />
Guinea-Bissau: No specific locality (Osorio, 1895a). Agongoon<br />
on Ilha Caravela, and Porto de Bissau on Ilha de<br />
Bissau (Vilela, 1949).<br />
Guinea: Conakry (Capart, 1951; Sourie, 1954a; Uschakov,<br />
1970).<br />
Sierra Leone: No specific locality (Longhurst, 1958).<br />
Liberia: No specific locality (Von Martens, 1872; Biittikofer,<br />
1890; Johnston, 1906). Monrovia (Rathbun, 1918).<br />
Rock Spring and mouth of Mesurado River, near Monrovia<br />
(Rathbun, 1900a, 1918). Cape Mesurado (Balss, 1922).<br />
Ivory Coast: Lagune Ebrie at Adiapo-doume (Humes,<br />
1957).<br />
Ghana: No specific locality (Gauld, 1960). Butre (typelocality;<br />
Herklots, 1851; De Man, 1879, 1900). Baya River<br />
near Elmina (Benedict, 1893, as Ashantee; Rathbun, 1918).<br />
Accra (Irvine, 1947). Sakumo lagoon, near mouth (Pauly,<br />
1975).<br />
Nigeria: Lagos (Balss, 1922; Bruce-Chwatt and Fitz-John,<br />
1951). Iru Fisheries Station near Lagos (Humes, 1957).<br />
Elechi Creek, Port Harcourt, 04°47'15"N, 06°58'45"E (Powell,<br />
1979).<br />
Cameroon: No specific locality (Doflein, 1900; Balss,<br />
1922; Monod, 1927, 1928). Douala (Monod, 1927, 1956).<br />
Rio Muni: Cabo San Juan (Nobili, 1906b). Rio Muni,<br />
20 km up from Elobey (Balss, 1922). Islas de Elobey (Osorio,<br />
1895a).<br />
Fernando Poo: San Carlos and Mongola (Osorio, 1895a;<br />
see "Remarks").<br />
Principe: No specific locality (Frade, 1950).—Praia Salgada<br />
(Osorio, 1889; Forest and Guinot, 1966).—Rio Papagaio<br />
(Forest and Guinot, 1966).<br />
Sao Tome: No specific locality (Osorio, 1888, 1889; Balss,<br />
1922; Frade, 1950; Forest and Guinot, 1966). Praia das<br />
Conchas (Osorio, 1889). Iogoiogo (Osorio, 1890). Binda<br />
(Osorio, 1892).<br />
Gabon: No specific locality (Kingsley, 1880a). Cap Lopez<br />
(Balss, 1922). Port-Gentil (Roux, 1927). Ogooue (Thallwitz,<br />
1891).<br />
Congo: Songololo River near Pointe-Noire; Loeme River,<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
about 18 km S of Pointe-Noire (Humes, 1957). Loango, and<br />
mouth of Songololo River (Rossignol, 1957, 1962). Djeno,<br />
near Pointe-Noire (Rossignol, 1962).<br />
Zaire: Banana (Rathbun, 1921; Capart, 1951). Moanda<br />
(Rathbun, 1921).<br />
Angola: No specific locality (Osorio, 1887). Santo Antonio<br />
do Zaire (as San Antonio) (Rathbun, 1921). Lobito (Osorio,<br />
1887). Mocamedes (Osorio, 1895a). Between Cacuaco and<br />
Lobito-Benguela (Hartmann-Schroder and Hartmann,<br />
1974).<br />
Genus Grapsus Lamarck, 1801<br />
Gmpsus Lamarck, 1801:150 [type-species: Cancer grapsus Linnaeus,<br />
1758, by tautonomy; gender: masculine].<br />
* Grapsus grapsus (Linnaeus, 1758)<br />
Grapsus maculatus.—Biittikofer, 1890:465, 487.—Johnston,<br />
1906:862.<br />
Grapsus grapsus.—Bruce-Chwatt and Fitz-John, 1951:117.—<br />
Capart, 1951:181, fig. 70.—Monod, 1956:407, fig. 561.—<br />
Dubois, 1957:7.—Rossignol, 1957:87, 121 [key].—Gauld<br />
and Buchanan, 1959:124.—Forest and Games, 1960:<br />
353.—Gauld, 1960:71.—Guinot and Ribeiro, 1962:68.—<br />
Rossignol, 1962:119.—Ribeiro, 1964:14.—Chace, 1966:<br />
640.—Forest and Guinot, 1966:90.—Zariquiey Alvarez,<br />
1968:422, fig. 139a [Portugal].—Kensley, 1970:181.—<br />
Penrith and Kensley, 1970b:246, 250, 252, 260.—Hartmann-Schroder<br />
and Hartmann, 1974:24.<br />
SYNONYMS.—Grapsus pictus Latreille, 1803;<br />
Grapsus maculatus H. Milne Edwards, 1853; Grapsus<br />
webbi H. Milne Edwards, 1853; Grapsus pictus<br />
ocellatus Studer, 1883; Cancer jumpibus Swire, 1938.<br />
MATERIAL EXAMINED.—Pillsbury Material: Fernando<br />
Poo: Sta 258, shore, 86, 3$, 3 juv (W).<br />
Annobon: Sta 271, shore, 36, 2$, 5 juv (L). Sta 273, shore,<br />
26, 3 juv (L). Sta 281, shore, 1 cast (L).<br />
Other Material: Azores: No specific locality, W. Trelease,<br />
3c5, 3$ ov (W). Near harbor of Ilha do Corvo, 4 Jun 1976,<br />
W. Backhuys, 6 spec (L). Ponta Delgada, Ilha de Sao Miguel,<br />
31 Aug 1949, G. J. Jacobs, lc? (W). Horta, Ilha do Faial, U.<br />
S. Exploring Expedition, 1$ (W). Pirn Bay, Horta, Ilha do<br />
Faial, L. Dexter, 1$ (W).<br />
Senegal: Dakar, in tide pool, 12 Feb 1969, D. E. Harvey,<br />
1
NUMBER 306 233<br />
Ghana: Virgins Pool, Takoradi, 27-29 Jul 1961, Bane<br />
and Richards, 46\ 3$ (1 ov) (W).<br />
Cameroon: Kribi, common on rocky boulders, below and<br />
above low tide line, very fast, 8 Mar and 6 Aug 1964, B. de<br />
Wilde-Duyfjes, 19, 1 juv (L).<br />
Zaire: Banana, mouth of the Congo River, Jul 1915, H.<br />
Lang, American Museum Congo Expedition, 26, 19 ov (W).<br />
DESCRIPTION.—Capart, 1951:181; Chace and<br />
Hobbs, 1969:163.<br />
Figures: Capart, 1951, fig. 70; Monod, 1956, fig.<br />
561; Chace and Hobbs, 1969, figs. 50, 52g-i.<br />
Male Pleopod: Chace and Hobbs, 1969, fig.<br />
52g-i (<strong>West</strong> Indies).<br />
MEASUREMENTS.—The carapace width of the<br />
examined males ranged between 16 and 70 mm,<br />
that of the non-ovigerous females between 23 and<br />
43 mm, that of the ovigerous females between 32<br />
and 71 mm, and that of the juveniles between 5<br />
and 14 mm.<br />
BIOLOGY.—The species can be found on rocks<br />
in the surf- or splash-zone. A good description of<br />
its habitat is given by Chace and Hobbs (1969:<br />
165, 166).<br />
The species is very agile and exceedingly difficult<br />
to catch in the daytime; at night it can be<br />
approached far easier. See also Chace and Hobbs<br />
(1969:166). Off <strong>West</strong> Africa ovigerous females<br />
have been collected in February, May, July, and<br />
August (Guinot and Ribeiro, 1962; Ribeiro, 1964;<br />
present paper).<br />
DISTRIBUTION.—The species has a wide range<br />
in the tropical Atlantic and East Pacific regions.<br />
In the East Atlantic it is found from the Azores<br />
and Morocco to South-<strong>West</strong> Africa; there is a<br />
single record from Setubal, Portugal (Osorio,<br />
1905), but this needs confirmation. Monod (1956)<br />
listed the <strong>West</strong> <strong>African</strong> records, to which the<br />
following can now be added.<br />
Morocco: Temara, S of Rabat (Forest and Gantes, 1960).<br />
Cape Verde Islands: Baia de Monte Trigo, Santo Antao;<br />
Matiota, Baia da Fateixa, and Baia da Calheta, Sao Vicente;<br />
Baia da Murdeira and Pedra Lurae, Sal; Baia do Tarrafal,<br />
Sao Tiago; and Ponta Rodrigo, Boavista (all Guinot and<br />
Ribeiro, 1962; Ribeiro, 1964).<br />
Liberia: No specific locality (Biittikofer, 1890; Johnston,<br />
1906).<br />
Ghana: No specific locality (Gauld, 1960). Teshi, 8 mi [13<br />
km] E of Accra (Gauld and Buchanan, 1959). Takoradi and<br />
Tema harbors (Gauld, 1960).<br />
Nigeria: Lagos (Bruce-Chwatt and Fitz-John, 1951).<br />
Principe: Praia Ponta da Mina (Forest and Guinot, 1966).<br />
Sao Tome: Praia Pantufo, Praia Melao, Ilheu das Cabras,<br />
Binda, and Ilheu Gago Coutinho (as Hot das Rollas) (Forest<br />
and Guinot, 1966).<br />
Annobon: San Antonio (Forest and Guinot, 1966).<br />
Congo: Djeno (Rossignol, 1957, 1962). Baie de Loango<br />
and Pointe-Noire (Rossignol, 1962).<br />
Zaire: No specific locality (as Belgian Congo) (Dubois,<br />
1957).<br />
Angola: Baia de Luanda; Lobito; Baia Farta, Baia da<br />
Caota, and Ponta da Carufta, Benguela; Bafa de Santa<br />
Marta; and Praia Amelia, Mocamedes (all Guinot and<br />
Ribeiro, 1962). Near Mocamedes (Hartmann-Schrbder and<br />
Hartmann, 1974).<br />
South-<strong>West</strong> Africa: Rocky Point, 18°59'S, 12°29'E (Kensley,<br />
1970; Penrith and Kensley, 1970b). Mbwe Point, 19°-<br />
23'S, 12°42'E (Kensley, 1970).<br />
Saint Helena: several localities (Chace, 1966).<br />
Genus Pachygrapsus Randall, 1840<br />
Pachygrapsus Randall, 1840:126 [type-species: Pachygrapsus<br />
crassipes Randall, 1840, by selection by Kingsley, 1880a:<br />
198; gender: masculine; name 1638 on Official List].<br />
Goniograpsus Dana, 1851c:247, 249 [type species: Goniograpsus<br />
innotatus Dana, 1851, a subjective junior synonym of<br />
Pachygrapsus transversus (Gibbes, 1850), by present selection;<br />
gender: masculine].<br />
* Pachygrapsus gracilis (De Saussure, 1858)<br />
Pachygrapsus gracilis.—Frade, 1950:11, 26.—Capart, 1951:<br />
187, fig. 74, pi. 3: fig. 19.—Monod, 1956:419, figs. 569,<br />
571, 574-577.—Rossignol, 1957:89, 122 [key].—Gauld,<br />
1960:71.—Guinot and Ribeiro, 1962:71.—Rossignol,<br />
1962:120.—Forest and Guinot, 1966:92.—Uschakov,<br />
1970:443, 444, 445.—Powell, 1979:127.<br />
Pachygrapsus "africanus".—Hartmann-Schrbder and Hartmann,<br />
1974:19 [error for P. gracilis].<br />
SYNONYM.—Grapsus guadulpensis Desbonne and<br />
Schramm, 1867.<br />
MATERIAL EXAMINED.—Pillsbury Material: Nigeria:<br />
Sta 1, Lagos harbor, shore, 156", 179 (8 ov) (L). Sta 227,<br />
Lagos harbor, shore, 36*, 2$ (W).<br />
Other Material: Liberia: No specific locality, 13 Mar 1953,<br />
G. C. Miller, 16*, 2$ (W). Freeport area, Monrovia, oyster<br />
cultch, 12 Mar 1953, G. C. Miller, 3c5, 8$ (4 ov) (W).<br />
Ghana: Virgins Pool, Takoradi, near sports club, 24 Jul<br />
1961, Bane and Richards, 19 (W).
234<br />
Dahomey: Cotonou, Point XI Lagune, 8 Apr 1964, H.<br />
Hoestlandt, 1$ ov (L).<br />
Nigeria: Ogudu River, 8 Aug 1926, A. S. Pearse, 2$ (W).<br />
Lagos, harbor, 18 Jul 1963, A. R. Longhurst, 1(5 (L). W of<br />
Forcados, near confluence of Odimodi Creek and Forcados<br />
River, 05°22'N, 05°26'E, 28 Feb 1976, C. B. Powell, 16,<br />
1 $ (L). Niger delta between Brass and Port Harcourt, May-<br />
Aug 1960, H. J. G. Beets, 3
NUMBER 306 235<br />
stones on river shore, 15 Jul 1915, H. Lang, American<br />
Museum Congo Expedition, 10
236<br />
terial of this species from the Pacific Ocean between<br />
roughly 41 °N and 35°S latitudes and from<br />
the Indian Ocean; no reliable records of the<br />
species from the Atlantic Ocean were known to<br />
him. Later records showed that the species is<br />
rather common in the South Atlantic Ocean,<br />
being found there far more frequently than P.<br />
minutus; Juanico (1976:149) is of the opinion that<br />
it is the only species of Planes there (but see p.<br />
237). Juanico reported an extensive material (28<br />
specimens) of the species from various localities<br />
on the Uruguay coast. Chace (1966:646) examined<br />
11 specimens from various parts of the coast<br />
of St. Helena. The only certain record of the<br />
species from the coast of tropical <strong>West</strong> Africa is<br />
the one by Crosnier (1967), who examined a<br />
female from a buoy off Gabon, 02°50'S, 08°50'E.<br />
The species has once been reported from the<br />
North Atlantic, viz., by Shirley (1974) from North<br />
Padre Island, Texas, U.S.A. The unconfirmed<br />
records of Planes minutus from the <strong>West</strong> coast of<br />
Africa (Gambia, Liberia, Cameroon and "Gulph<br />
of Guinea"), discussed below, might pertain to<br />
the present species.<br />
Planes minutus (Linnaeus, 1758)<br />
Cancer minutus Linnaeus, 1758:625.<br />
Grapsus minutus.—Leach, 1818, in 1817-1818:414.—Monod,<br />
1970:66.<br />
Planes minutus.—Monod, 1956:425, fig. 583.—Christiansen,<br />
1969:94, fig. 39, map 32 [Netherlands].<br />
SYNONYMS.—Cancer pusillus Fabricius, 1775;<br />
Grapsus pelagicus Say, 1818; Planes clypeatus Bowdich,<br />
1825; Grapsus testudinum Roux, 1828; Nautilograpsus<br />
major MacLeay, 1838; Nautilograpsus smithii<br />
MacLeay, 1838; Grapsus dirts Costa, 1838-<br />
1853; Planes linnaeana Bell, 1845.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Morocco: Off Cap Blanc du Nord, 33°-<br />
16'N, 09°10'W, surface, collected with hand net, 27 Mar<br />
1976, Orwersaagd Sta 140, 1
NUMBER 306 237<br />
the Northside of Jamaica." Chace (1951:81) gave<br />
this as the first reliable record of Planes minutus.<br />
Finally Linnaeus' reference, "Osbeck.iter.," is to<br />
Osbeck (1757:307), who mentioned Cancer minutus<br />
from the Sargasso Sea (Gras-Sjb) at 22°N in the<br />
Atlantic north of Ascension. Osbeck gave no morphological<br />
description of this species, but did so<br />
of Cancer pelagicus, which showed the latter to be<br />
Portunus sayi Gibbes; therefore, there can be little<br />
doubt that Osbeck's Cancer minutus is a Planes, and<br />
Chace (1951:81) also considered it as such.<br />
LECTOTYPE.—So far as we know no lectotype<br />
has ever been selected for Cancer minutus L. and in<br />
order to legalize the continued use of the epithet<br />
minutus for the North Atlantic Planes species, we<br />
now select as the lectotype of Linnaeus' species<br />
the specimen from the north coast of Jamaica<br />
figured by Sloane (1725, pi. 245, fig. 1).<br />
BIOLOGY.—Monod (1927:621) reported on four<br />
specimens found in Douala Bay near Souellaba,<br />
Cameroon, on a floating trunk of a tree (Aucoumea<br />
sp., family Burseraceae). The same author later<br />
(1956:426) examined eight specimens found near<br />
Cotonou, Dahomey, on floating weeds "Fucus<br />
natans," by which the collector doubtless meant<br />
Sargassum. Outside the tropical <strong>West</strong> <strong>African</strong> region<br />
the species has been reported from floating<br />
Sargassum weed, from marine turtles, and from<br />
pieces of wood and other floating objects; it is a<br />
true oceanic form.<br />
DISTRIBUTION.—There are very few records of<br />
the species from <strong>West</strong> Africa. Monod (1956:425,<br />
426) listed only four: Gambia River (Gambia),<br />
Monrovia (Liberia), Cotonou (Dahomey), and<br />
Souellaba (Cameroon). Later, Monod (1970:66)<br />
drew attention to Leach's (1818, in 1817-1818:<br />
414) record of the species from the "Gulph of<br />
Guinea." No other <strong>West</strong> <strong>African</strong> records are<br />
known to us.<br />
Chace (1951:80, fig. 8) indicated the specimens<br />
from Gambia, Liberia, and Cameroon as of uncertain<br />
identity, being either Planes minutus or P.<br />
cyaneus Dana; no material from these localities<br />
was seen by Chace and the data provided in the<br />
literature were insufficient to identify them with<br />
certainty. The specimens mentioned by Leach<br />
(1818, in 1817-1818) and Monod (1970) as Grapsus<br />
minutus cannot be identified, because too little<br />
is known about them.<br />
Quite different is the case with the specimens<br />
from Cotonou mentioned and figured by Monod<br />
(1956:425, fig. 583). Monod based the identification<br />
of his material on Chace's (1951) revision<br />
of the genus Planes, in which the differences between<br />
P. minutus and P. cyaneus were extensively<br />
dealt with. Furthermore, the Cotonou specimen<br />
figured by Monod shows the slender second pereiopod<br />
indicated by Chace as characteristic for P.<br />
minutus, and resembles that species also in other<br />
respects, so that there is no good reason to doubt<br />
Monod's identification. It is rather difficult therefore<br />
to understand Juanico (1976:146), who tried<br />
to show that Monod's identification is not reliable<br />
by stating on this account: "Monod (1956), dispone<br />
de un material de 8 ejemplares, pero no<br />
hace ninguna referencia a haberlos analizado<br />
respecto a los parametros que permiten distinguir<br />
una especie de otra [= P. minutus from P. cyaneus]."<br />
Juanico reached the conclusion that P.<br />
minutus is restricted to the North Atlantic, but did<br />
not define what he considered the line between<br />
the North and South Atlantic. Judging by the<br />
fact that he evidently considered Surinam, Pernambuco,<br />
and Dahomey to be in the South Atlantic,<br />
this line is not the equator. Therefore, we<br />
consider for now that both species of Planes occur<br />
in <strong>West</strong> <strong>African</strong> waters, with Monod's (1956)<br />
record of Planes minutus from Cotonou constituting<br />
the only reliable record of that species for the<br />
area.<br />
Outside the tropical <strong>West</strong> <strong>African</strong> region Planes<br />
minutus is known with certainty only from the<br />
North Atlantic between 11° and 52°N (see also<br />
Chace, 1951, fig. 8).<br />
Subfamily PLAGUSIINAE Dana, 1851<br />
Genus Percnon Gistel, 1848<br />
Acanthopus de Haan, 1833:5, 6 [invalid junior homonym of<br />
Acanthopus Klug, 1807 (Hymenoptera); a genus established<br />
without included nominal species; the first species assigned<br />
to the genus, by de Haan, 1835:29, 30, were Plagusia
238 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
clavimana Latreille, 1806, Cancer planissimus Herbst, 1804<br />
(as a synonym of P. clavimana), and Plagusia serripes Lamarck,<br />
1818. Type-species: Cancer planissimus Herbst, 1804,<br />
by selection by Rathbun, 1918:337; gender: masculine;<br />
name 465 on Official Index].<br />
Percnon Gistel, 1848:viii [substitute name for Acanthopus de<br />
Haan, 1833; type-species: Cancer planissimus Herbst, 1804;<br />
gender: neuter; name 345 on Official List}.<br />
Leiolophus Miers, 1876:46 [substitute name for Acanthopus de<br />
Haan, 1833; type-species: Cancer planissimus Herbst, 1804;<br />
gender: masculine].<br />
Liolophus Alcock, 1900:439 [emendation of Leiolophus Miers,<br />
1876; gender: masculine].<br />
* Percnon gibbesi (H. Milne Edwards, 1853)<br />
Acanthopus Gibbesi H. Milne Edwards, 1853:180.<br />
Percnon planissimum.—Monod, 1956:453, fig. 613.—Gauld<br />
and Buchanan, 1959:128.—Forest and Games, 1960:<br />
355.—Gauld, 1960:71.—Guinot and Ribeiro, 1962:72.—<br />
Ribeiro, 1964:17.—Forest and Guinot, 1966:93.—Zariquiey<br />
Alvarez, 1968:436, fig. 146a [Portugal]. [Not Cancer<br />
planissimus Herbst, 1804.]<br />
Percnon gibbesi.—Chapman and Santler, 1955:375.—Figueira,<br />
1960:11.<br />
SYNONYM.—Plagusia Delaunayi De Rochebrune,<br />
1883.<br />
MATERIAL EXAMINED.—Pillsbury Material: Fernando<br />
Poo: Sta 258, shore, \6, 1 juv (W).<br />
Annobon: Sta 271, shore, 3
NUMBER 306 239<br />
*Plagusia depressa (Fabricius, 1775)<br />
Plagusia depressa.—Buttikofer, 1890:487.—Johnston, 1906:<br />
862.—Monod, 1956:455, figs. 614-617.—Rossignol, 1957:<br />
95, 121 [key], fig. 9.—Figueira, 1960:11.—Forest and<br />
Games, 1960:356.—Gauld, 1960:71.—Guinot and Ribeiro,<br />
1962:72.—Rossignol, 1962:121.—Ribeiro, 1964:<br />
17.—Chace, 1966:647.—Forest and Guinot, 1966:93.—<br />
Hartmann-Schroder and Hartmann, 1974:24.<br />
Plagusia squamosa.—Stimpson, 1907:122.<br />
SYNONYMS.—Cancer squamosus Herbst, 1790; Plagusia<br />
sayi De Kay, 1844; Plagusia gracilis De Saussure,<br />
1858.<br />
MATERIAL EXAMINED.—Pillsbury Material: Nigeria:<br />
Sta 225, Lagos harbor, shore, 1$ ov (W).<br />
Annobon: Sta 271, shore, Id, 1$ ov (L). Sta 273, shore, 2$<br />
(1 ov) (W).<br />
Other Material: Madeira: No specific locality; U. S. Exploring<br />
Expedition, 1(5, 1? (W).<br />
Liberia: No specific locality, 1882, J. Buttikofer, let, 1?<br />
(L). No specific locality, 1890, J. Demery, 1$ (L).<br />
Ghana: Butre, 1840-1855, H. S. Pel, 1$ (L). Sekondi,<br />
1853, H. S. Pel, several specimens (L). Baya River, Elmina,<br />
27 Nov 1889, W. H. Brown, U. S. Eclipse Expedition, 16*<br />
(W). Accra, 1868-1869, M. Sintenis, 26*, 1$ (L).<br />
Cameroon: Kribi, among algae on rocks on a sandy<br />
beach, 11 Mar 1964, B. de Wilde-Duyfjes, 8
240 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Angola: Lobito, 1899, P. Kamerman, 1? (L).<br />
DESCRIPTION.—Rathbun, 1921:455; Chace and<br />
Hobbs, 1969:173.<br />
Figure: Monod, 1956, fig. 609.<br />
Male Pleopod: Monod, 1956, figs. 610-612 (Cape<br />
Verde Islands; Senegal); Ghace and Hobbs, 1969,<br />
fig. 58b-d (<strong>West</strong> Indies).<br />
Color: "Brun-rouge uniforme. Extremite des<br />
pattes brun fonce, extremites des pinces jaunatres"<br />
(Rossignol, 1957:94).<br />
MEASUREMENTS.—Our specimens have carapace<br />
widths ranging from 6 to 12 mm, those of<br />
ovigerous females being 8 and 9 mm.<br />
BIOLOGY.—This species is found under rocks in<br />
the intertidal zone. Off <strong>West</strong> Africa ovigerous<br />
females have been collected in March, May, and<br />
July-<br />
DISTRIBUTION.—The species inhabits both the<br />
eastern and the western Atlantic; it has also been<br />
reported from the Indo-<strong>West</strong> Pacific region, from<br />
East Africa to Polynesia (Campbell and Griffin,<br />
1966; Griffin, 1968b). In the western Atlantic it<br />
occurs from Bermuda and Florida to Brazil. In<br />
the eastern Atlantic its range extends from the<br />
Cape Verde Islands and Senegal to Angola. To<br />
the <strong>West</strong> <strong>African</strong> localities enumerated by Monod<br />
(1956) the following can be added:<br />
Cape Verde Islands: Baia da Murdeira, Sal (Guinot and<br />
Ribeiro, 1962; Ribeiro, 1964).<br />
Senegal: Dakar (Sourie, 1954a).<br />
Guinea: Conakry (Sourie, 1954a; Uschakov, 1970).<br />
Congo: Loango lagoon (Rossignol, 1957, 1962). Pointe-<br />
Noire (Rossignol, 1962).<br />
Angola: Lobito (Bott, 1964). Baia Farta and Baia de<br />
Santa Marta, Benguela (Guinot and Ribeiro, 1962).<br />
Genus Metagrapsus H. Milne Edwards, 1853<br />
Melagrapsus H. Milne Edwards, 1853:188 [type-species: Sesarma<br />
curvatum H. Milne Edwards, 1837, selection by Rathbun,<br />
1918:321; gender: masculine].<br />
REMARKS.—Serene and Soh (1970:397) restricted<br />
the genus Sarmatium Dana, 1851, to include<br />
only two species: the type, Sarmatium crassum<br />
Dana, 1851, and Sesarma germaini A. Milne Edwards,<br />
1869. The <strong>West</strong> <strong>African</strong> species, which is<br />
best known as Sarmatium curvatum (H. Milne Edwards,<br />
1837), according to these authors belongs<br />
to a distinct genus, for which the name Metagrapsus<br />
H. Milne Edwards, 1853, is available. Serene<br />
and Soh (1970:398) placed a second species in the<br />
genus Metagrapsus, viz., Metagrapsus pectinatus H.<br />
Milne Edwards, 1853. According to Rathbun<br />
(1918:322), who examined the type of Metagrapsus<br />
pectinatus, this species is synonymous with M. curvatus.<br />
Some authors, however, doubted the correctness<br />
of Rathbun's identification, mainly on<br />
zoogeographic grounds. As the type of M. pectinatus<br />
(said to be from Martinique) so far is the only<br />
specimen of the genus ever reported from the<br />
western Atlantic, there seems to be more reason<br />
to doubt the correctness of the locality label<br />
"Martinique" than Rathbun's identification. If<br />
Rathbun is correct, M. curvatus at present is the<br />
only species known of the genus Metagrapsus.<br />
* Metagrapsus curvatus<br />
(H. Milne Edwards, 1837)<br />
Sarmatium curvatum.—Capart, 1951:193.—Monod, 1956:449,<br />
fig. 608.—Rossignol, 1957:93, 122 [key], fig. 7.—Jordan,<br />
1957:198.—Humes, 1957:184, 187, 189.—Longhurst,<br />
1958:88.—Gauld, 1960:71.—Rossignol, 1962:121.—Uschakov,<br />
1970:448,455 [listed].—Powell, 1979:129.<br />
Sesarma curvatum.—Humes, 1957:186.<br />
Sarmatium.—Humes, 1957:188, 189.<br />
SYNONYMS.—Sesarma violacea Herklots, 1851;<br />
?Metagrapsus pectinatus H. Milne Edwards, 1853.<br />
MATERIAL EXAMINED.—Pillsbury Material: Nigeria:<br />
Sta 1, Lagos harbor, shore, 3
NUMBER 306 241<br />
koko, about 10 km west of Sapele, 05°N, 05°34.5'E, Oct-Dec<br />
1975, C. B. Powell, Ic5 (L). Between Brass and Port Harcourt,<br />
Niger delta, May-Aug 1960, H. J. G. Beets, 26, 1? (L).<br />
Congo: No specific locality, 1894, Dybowski, legator, 16\<br />
1$ (W).<br />
Zaire: Banana, mouth of the Congo River, Jul-Aug 1915,<br />
H. Lang, American Museum Congo Expedition, 4(5, 1$ (W).<br />
Malela, 8 Jul 1915, H. Lang, American Museum Congo<br />
Expedition, 5
242<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Episesarma De Man (1895b: 165). Type-species: Sesarmoides Serene and Soh (1970:392, 403).<br />
Sesarma taeniolatum Miers, 1877 (= Sesarma taeni- Type-species: Sesarma krausii De Man, 1887, by<br />
olata White, 1847, a nomen nudum), a subjective original designation; gender: masculine.<br />
junior synonym of Sesarma meden H. Milne Ed- Namlacium Serene and Soh (1970:392, 404).<br />
wards, 1853, selected by Holthuis (1978:24); gen- Type-species: Sesarma crepidatum Caiman, 1925, by<br />
der: neuter.<br />
Parasesarma De Man (1895b: 181). Type-species:<br />
original designation; gender: neuter.<br />
Cancer quadratus Fabricius, 1798 (not Fabricius,<br />
1787)(= Ocypode plicata Latreille, 1803), by selec-<br />
Subgenus Chiromantes Gistel, 1848<br />
tion by Rathbun, 1918:284); gender: neuter.<br />
REMARKS.—The name Chiromantes was pro-<br />
Perisesarma De Man (1895b: 208). Type-species: posed by Gistel (1848:x) as a replacement for the<br />
Sesarma {Perisesarma) eumolpe De Man, 1895, se- generic name Pachysoma de Haan, 1833, which is<br />
lected by Holthuis (1977a: 170); gender: neuter. preoccupied by Pachysoma MacLeay, 1821 (Co-<br />
Nanosesarma Tweedie (1950:310). Type-species: leoptera) and Pachysoma Geoffroy, 1828 (Mam-<br />
Sesarma andersonii De Man, 1887, by original desmalia).ignation; gender: neuter.<br />
De Haan (1833:5, circ. pi. 2, pi. 7: fig. 4, pi. 8:<br />
Beamum Serene and Soh (1970:389, 394). Type- fig. 3) introduced the name Pachysoma in the first<br />
species: Sesarma batavica Moreira, 1903, by original fascicle of his Crustacea volume of Ph. F. von<br />
designation; gender: neuter.<br />
Siebold's Fauna Japonica. In the text of this fascicle<br />
Neosesarma Serene and Soh (1970:389, 394). the name Pachysoma was given in a key to the<br />
Type-species: Sesarma gemmiferum Tweedie, 1936, subgenera of the genus Grapsus, and no species<br />
by original designation; gender: neuter.<br />
were mentioned there. However, on two of the<br />
Neoepisesarma Serene and Soh (1970:390, 395). plates of this first fascicle two species of the<br />
Type-species: Sesarma mederi H. Milne Edwards, subgenus Pachysoma were figured and mentioned<br />
1853, by original designation; gender: neuter. by name: Grapsus (Pachysoma) haematocheir de Haan<br />
Muradium Serene and Soh (1970:390, 396). (1833, pi. 7: fig. 4) and Grapsus (Pachysoma) quad-<br />
Type-species: Cancer tetragonus Fabricius, 1798, by ratus (Fabricius, 1798) (pi. 8: fig. 3). As these two<br />
original designation; gender: neuter.<br />
species are the only nominal species referred to<br />
Selatium Serene and Soh (1970:390, 397). Type- Pachysoma in the original publication of that<br />
species: Sesarma brockii De Man, 1887, by original subgenus, it is from among them that a type may<br />
designation; gender: neuter.<br />
be selected for the subgenus. Therefore, as Hol-<br />
Tiomanum Serene and Soh (1970:391, 398). thuis (1977a: 170) pointed out, the selection of<br />
Type-species: Sesarma indica H. Milne Edwards, Grapsus (Pachysoma) bidens de Haan, 1835, as the<br />
1837, by original designation; gender: neuter. type of Pachysoma, made by Fowler (1912:439) is<br />
Bresedium Serene and Soh (1970:391, 399). invalid. The first valid type selection for Pachysoma<br />
Type-species: Sesarma edwardsii brevipes De Man, de Haan, 1833 (and thus automatically for Chi-<br />
1889, by original designation; gender: neuter. romantes Gistel, 1848) is the one by Holthuis<br />
Pseudosesarma Serene and Soh (1970:391, 399). (1977a: 170), who selected Grapsus (Pachysoma) hae-<br />
Type-species: Sesarma edwardsi De Man, 1888, by matocheir de Haan, 1833, as the type of that<br />
original designation; gender: neuter.<br />
subgenus.<br />
Sesarmops Serene and Soh (1970:391, 400). As Grapsus (Pachysoma) haematocheir de Haan,<br />
Type-species: Sesarma impressa H. Milne Edwards, 1833, is also the type-species (by monotypy) of<br />
1837, by original designation; gender: masculine. the subgenus Holometopus H. Milne Edwards,<br />
Labuanium Serene and Soh (1970:392, 401). 1853, the name Chiromantes, being the senior of<br />
Type-species: Sesarma polita De Man, 1888, by the two, has to be used.<br />
original designation; gender: neuter.<br />
Although the transfer of a name from one
NUMBER 306 243<br />
taxon to another usually is harmful and to be<br />
avoided, the status of the taxonomy and nomenclature<br />
of the subgenera of Sesarma is at present<br />
in such a state of flux that it seems best to adhere<br />
strictly to the Code in the present case, at least<br />
until a revision of the Sesarma complex is achieved.<br />
The subgenus Chiromantes is represented in the<br />
eastern Atlantic by three species, all three of<br />
which occur in tropical <strong>West</strong> Africa. Sesarma roberti,<br />
erroneously reported from <strong>West</strong> Africa (p.<br />
226), also belongs in the present subgenus.<br />
The subgenus was referred to as Holometopus by<br />
Monod (1956) and all later authors, while they<br />
used the name Chiromantes for the subgenus here<br />
indicated with the name Perisesarma.<br />
Sesarma (Chiromantes) angolense De Brito<br />
Capello, 1864<br />
Sesarma angolensis.— Biittikofer, 1890:487.—Johnston, 1906:<br />
861.—Longhurst, 1958:88.—Gauld, 1960:71.<br />
Sesarma {Holometopus) angolensis.—Dartevelle, 1950:48.—<br />
Monod, 1956:445, fig. 605.—Rossignol, 1962:120.<br />
Sesarma {Holometopus) angolense.—Dartevelle, 1950:50.—Capart,<br />
1951:191, fig. 76.—Rossignol, 1957:92, 122 [key].—<br />
Jordan, 1957:198.—Humes, 1957:186, 187, 189.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Liberia: No specific locality, 1890, 1897, J.<br />
Demery, 2? (L). Grand Cape Mount near Robertsport, 1882,<br />
J. Biittikofer, 4(5, 3$ (L).<br />
Nigeria: Mayuku Creek near Ugbekoko, 10 km W of<br />
Sapele, Midwest State, ca. 05°54'N, 05°34.5'W, Oct, Oct-<br />
Dec, Dec 1975, Jan 1976, C. B. Powell, many specimens (L).<br />
Niger delta between Brass and Port Harcourt, May-Aug<br />
1960, H. J. G. Beets, 16, 29 (L).<br />
Angola: Lobito, 1899, P. Kamerman, 1
244<br />
of Sapele, 05°54'N, 05°37'E, in mangrove swamp, Oct-Dec<br />
1975, C. B. Powell, 28 specimens (1$ ov) (L).<br />
Cameroon: Kribi, among humid dead leaves on the bank<br />
of a forest stream not far from the sea, 8 Aug 1964, B. de<br />
Wilde-Duyfjes, Id, 2$ (L).<br />
Zaire: Banana, mouth of the Congo River, Jul-Aug 1915,<br />
H. Lang, American Museum Congo Expedition, Id, 19 (W).<br />
Malela, 8 Jun 1915, H. Lang, American Museum Congo<br />
Expedition, 226*, 109 (2 ov) (W). Zambi, 4 Jul 1915, H.<br />
Lang, American Museum Congo Expedition, 1 juv (W).<br />
DESCRIPTION.—Rathbun, 1921:449.<br />
Figures: Monod, 1956, figs. 606, 607; Rossignol,<br />
1957, fig. 6.<br />
Color: The specimens from Lagune Ebrie carried<br />
a note stating that in life the chelae are<br />
purplish blue with red tips. Additional color notes<br />
were provided by Rossignol (1957:91).<br />
BIOLOGY.—This species inhabits littoral estuarine<br />
areas, particularly mangroves and adjoining<br />
inland regions (see Rathbun, 1921:449, 450).<br />
Ovigerous females have been collected in<br />
March, June, July-August, and October through<br />
December (Rathbun, 1921; Rossignol, 1957; present<br />
paper).<br />
DISTRIBUTION.—Sesarma buettikoferi is found<br />
along the <strong>West</strong> <strong>African</strong> coast from Liberia to<br />
Angola. Monod (1956) listed the locality records<br />
known to him; to these the following can now be<br />
added:<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
59 paralectotypes (L, lectotype Crust. D. 150, paralectotypes<br />
Crust. D. 151).<br />
Nigeria: Harbor of Lagos, 13 Jun 1963, A. R. Longhurst,<br />
1
NUMBER 306 245<br />
<strong>West</strong> Africa (Guinea to Angola). To the localities<br />
listed by Monod (1956) the following can be<br />
added.<br />
Guinea: lie Marara, Rio Pongo (Uschakov, 1970).<br />
Sierra Leone: Kissy (Longhurst, 1958).<br />
Ghana: Near Butre (Gauld, 1960).<br />
Nigeria: Elechi Creek, Port Harcourt, 04°47'15"N, 06°<br />
48'45"E (Powell, 1979).<br />
Congo: Loya River and Djeno (Rossignol, 1957, 1962).<br />
Pointe-Noire (Rossignol, 1962).<br />
Zaire: Banana, mouth of the Congo River (Dartevelle,<br />
1950).<br />
Angola: Santo Antonio do Zaire (as Saint-Antoine) (Dartevelle,<br />
1950).<br />
Subgenus Perisesarma De Man, 1895<br />
REMARKS.—This subgenus was cited by Monod<br />
(1956) and most other authors as Chiromantes Gistel,<br />
1848. However, as shown by Holthuis (1977:<br />
170) the type-species of Chiromantes is Grapsus<br />
(Pachysomd) haematocheir De Haan, 1833; hereby<br />
Chiromantes Gistel, 1848, becomes a senior objective<br />
synonym of Holometopus H. Milne Edwards,<br />
1853, and should replace the latter name (p. 242).<br />
For the subgenus usually cited as Chiromantes the<br />
name Perisesarma is available.<br />
Three species of Perisesarma are known from the<br />
eastern Atlantic area, all three are restricted to<br />
tropical <strong>West</strong> Africa.<br />
Sesarma (Perisesarma) alberti Rathbun, 1921<br />
Sesarma (Chiromantes) alberti.—Dartevelle, 1950:48.—Capart,<br />
1951:189, fig. 75.—Monod, 1956:440, fig. 594.—Jordan,<br />
1957:198.—Rossignol, 1957:122 [key]; 1962:120.—Forest<br />
and Guinot, 1966:93.<br />
Sesarma (chiromantes) alberti.—Rossignol, 1957:90.<br />
Sesarma alberti.—Longhurst, 1958:88.—Powell, 1979:127.<br />
MATERIAL EXAMINED—Pillsbury Material: None.<br />
Other Material: Liberia: St. John River at upper Buchanan,<br />
mud in mangroves, collected at night, 24 Aug 1967, T. C.<br />
Rutherford, 1$ (W). Rock Spring, Monrovia, O. F. Cook<br />
and G. N. Collins, 9juv (W). St. Paul River lagoon at True<br />
Rubber Farm, 9 Oct 1953, G. C. Miller, 1
246<br />
mann-Schroder and Hartmann, 1974:19.—Powell, 1979:<br />
127.<br />
SYNONYM.—Sesarma africana H. Milne Edwards,<br />
1837.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Senegal: Senegal, legator M. Delambre,<br />
donor Mus. Paris, Ic3 syntype of Sesarma africanum H. Milne<br />
Edwards, dry (W).<br />
Liberia: Liberia, 1881, J. Buttikofer and J. A. Sala, 1
NUMBER 306 247<br />
Elechi Creek, Port Harcourt, 04°47'15"N, 06°58'45"E (Powell,<br />
1979).<br />
Congo: Loango (Pechiiel-Loesche, 1882). Mouths of the<br />
Songololo and Loya rivers (Rossignol, 1957, 1962). Songololo<br />
River near Pointe-Noire; mouth of Loeme River near Pointe-<br />
Noire (Humes, 1957).<br />
Zaire: Banana, mouth of the Congo River (Dartevelle,<br />
1950).<br />
Angola: Santo Antonio do Zaire (as Saint Antoine) (Dartevelle,<br />
1950). Between Cacuaco and Lobito-Benguela (Hartmann-Schroder<br />
and Hartmann, 1974).<br />
Sesarma (Perisesarma) kamermani De Man,<br />
1883<br />
Sesarma (Chiromantes) kamermani.—Monod, 1956:441, figs.<br />
595-600.—Rossignol, 1957:122 [key].<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Angola: Musserra, 1882, P. Kamerman, Id<br />
holotype (L. Crust. D. 166).<br />
DESCRIPTION.—De Man, 1883:165; Balss,<br />
1936:1.<br />
Figures: Balss, 1936, figs. A, B; Monod, 1956,<br />
figs. 595-600.<br />
MEASUREMENTS.—The carapace length of the<br />
holotype is 27 mm, its carapace width is 32 mm.<br />
DISTRIBUTION.—So far only two specimens,<br />
both males, have become known of this species.<br />
The type-specimen (in the Leiden Museum) originated<br />
from Musserra, Angola. The second specimen<br />
was reported upon by Balss (1936) and<br />
Monod (1956) and was collected at Banana,<br />
Zaire.<br />
Subfamily VARUNINAE<br />
H. Milne Edwards, 1853<br />
Genus Brachynotus de Haan, 1833<br />
Brachynotus de Haan, 1833:5, 7 [subgenus established without<br />
included nominal species; type-species: Goneplax sexdentatus<br />
Risso, 1827, by subsequent monotypy by de Haan, 1835:<br />
34; gender: masculine; name 1610 on Official List].<br />
Heterograpsus Lucas, 1846:18 [type-species: Heterograpsus<br />
sexdentatus Lucas, 1846, a subjective junior synonym and<br />
homonym of Goneplax sexdentatus Risso, 1827, by monotypy;<br />
gender: masculine].<br />
Shurebus Verany, 1846:7 [type-species: Shurebus genuensis Verany,<br />
1846, an objective: junior 1 synonym of Cleistotomaigemmellan<br />
Rizza, 1839, by monotypy; gender: masculine].<br />
Brachynotus atlanticus Forest, 1957<br />
Brachynotus sexdentatus.—Monod, 1933a:219, fig. 7.—Bouvier,<br />
1940, fig. 184A only [copied from Monod, 1933a, fig. 7].<br />
[Not Goneplax sexdentatus Risso, 1827.]<br />
Brachynotus sexdentatus lucasi.—Monod, 1956:428, 631, figs.<br />
589-592, 881 [not Heterograpsus lucasi H. Milne Edwards,<br />
1853 = Goneplax sexdentatus Risso, 1827].<br />
Brachynotus atlanticus Forest, 1957:505, figs. 2, 4, 6, 8, 10, 12,<br />
14.—Forest and Games, 1960:354, fig. 2.<br />
MATERIAL EXAMINED.—None.<br />
DESCRIPTION.—Forest, 1957:505.<br />
Figures: Monod, 1933a, fig. 7; Monod, 1956,<br />
figs. 589-592, 881; Forest, 1957, figs. 2, 4, 6, 8,<br />
10, 12, 14.<br />
Male Pleopod: Monod, 1956, figs. 591, 592<br />
(Mauritania); Forest, 1957, fig. 14 (Mauritania).<br />
DISTRIBUTION.—This species, which until recently<br />
was not distinguished from B. sexdentatus,<br />
is only known from the Atlantic coast of Morocco<br />
and from the Cap Blanc peninsula (Mauritania).<br />
The records are the following.<br />
Morocco: Numerous localities as far north as 34°53'N<br />
(Monod, 1933a; Forest and Gantes, 1960).<br />
Mauritania: Baie de l'Ouest, coast of Cap Blanc peninsula<br />
(Monod, 1956: Forest, 1957). Pointe de Cansado (Monod,<br />
1956). Baie de Cansado (Monod, 1956; Forest, 1957). Port-<br />
Etienne (Monod, 1956). Baie du Repos near Port-Etienne<br />
(Monod, 1956).<br />
The Mauritanian localities are all situated on<br />
the east coast of the Cap Blanc peninsula. The<br />
species is not known to occur farther south.<br />
Genus Euchirograpsus H. Milne Edwards, 1853<br />
Euchirograpsus H. Milne Edwards, 1853:175 [type-species:<br />
Euchirograpsus liguricus H. Milne Edwards, 1853, by monotypy;<br />
gender: masculine; name 300 on Official List].<br />
Euchirograpsus liguricus H. Milne Edwards,<br />
1853<br />
Euchirograpsus liguricus H. Milne Edwards, 1853:175.—Tvirkay,<br />
1975b: 105, figs. 1-3, 17, 23; 1976a:25 [listed], 39, figs.<br />
29-31.<br />
Euchirograpsus americanus.—A. Milne Edwards and Bouvier,<br />
1894:46, pi. 4: figs. 10-14; 1900:107.—Dollfus and
248<br />
BIOLOGY.—This species has been reported from<br />
bottoms of sand, sandy mud, mud, rock with<br />
gorgonians and/or sponges, and gravel at depths<br />
between 10 and 359 m, most of the records being<br />
from between 100 and 250 m.<br />
Few ovigerous females have been recorded<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Monod, 1927:216, figs. 1, 2.—Monod, 1933a:320; 1933b: from <strong>West</strong> Africa; the only record in the recent<br />
535.—Capart, 1951:184, fig. 72.—Monod, 1956:434, figs.<br />
592 bis, 882-884.—Rossignol, 1957:121 [key].—Forest and<br />
Games, 1960:355.—Rossignol, 1962:120.—Forest and<br />
Guinot, 1966:92.—Crosnier, 1970:1215 [listed], 1217. [Not<br />
Enchirograpsus americanus A. Milne Edwards, 1880].<br />
literature is that by Crosnier (1970) based on a<br />
specimen taken by the Undaunted in March and<br />
also recorded herein.<br />
DISTRIBUTION.—The range of this species extends<br />
from west of Portugal and Madeira and the<br />
MATERIAL EXAMINED.—PUlsbury Material: None. Azores south to South-<strong>West</strong> Africa; it also occurs<br />
Geronimo Material: Gabon: Sta 211, 100 m, 1
NUMBER 306 249<br />
EASTERN ATLANTIC GENERA.—Two, Cardisoma<br />
and Gecarcinus, each represented by one tropical<br />
species.<br />
EASTERN ATLANTIC SPECIES.—Two, both reported<br />
below. Neither species is represented in<br />
the Pillsbury collections.<br />
Genus Cardisoma Latreille, 1828<br />
Cardisoma Latreille, 1828b:685 [type-species: Cardisoma guanhumi<br />
Latreille, 1828, selected by H. Milne Edwards, 1838,<br />
in 1836-1844, pi. 20; gender: neuter].<br />
Perigrapsus Heller, 1862:522 [type-species: Perigrapsus excelsus<br />
Heller, 1862, a subjective junior synonym ofCancer camifex<br />
Herbst, 1794, by monotypy; gender: masculine].<br />
Discoplax A. Milne Edwards, 1867b:284 [type-species: Discoplax<br />
longipes A. Milne Edwards, 1867, by monotypy; gender:<br />
feminine].<br />
Cardiosoma S.I. Smith, 1869b: 16, 36 [unjustified emendation<br />
of Cardisoma Latreille, 1828; type-species: Cardisoma guanhumi<br />
Latreille, 1828; gender: neuter].<br />
Cardisoma armatum Herklots, 1851<br />
Cardisoma Guanhuni.—Studer, 1882:333, 353 [discussion] [not<br />
C. guanhumi Latreille, 1828].<br />
Cardisoma armatum.—Pechiiel-Loesche, 1882:299.—Biittikofer,<br />
1890:464, 487, fig. on p. 465.—Johnston, 1906:861.—<br />
Gruvel, 1913:169 [listed].—Capart, 1951:196, fig. 79.—<br />
Monod, 1956:458, fig. 618.—Rossignol, 1957:95, 120<br />
[key], pi. 2: fig. 2.—Humes, 1957:181,189.—Dubois, 1957:<br />
7, fig. 24.—Gauld, 1960:72.—Rossignol, 1962:121.—<br />
Guinot and Ribeiro, 1962:73.—De Leersnyder and<br />
Hoestlandt, 1963:211 [physiology].—Ribeiro, 1964:18.—<br />
De Leersnyder and Hoestlandt, 1966:43 [physiology].—<br />
Forest and Guinot, 1966:94.—Monod, 1967:180, pi. 17:<br />
fig. 1.—Uschakov, 1970:450, 455 [listed].—Bright and<br />
Hogue, 1972:17.—Tiirkay, 1973:86, figs. 1, 3, 7-8, map<br />
1.—Hartmann-Schroder and Hartmann, 1974:19.<br />
Cardisoma Guanhumi var. armatum.—Bouvier, 1911:226.<br />
Cardisoma guanhumi.—Gruvel, 1913:169 [listed] [not C. guanhumi<br />
Latreille, 1828].<br />
Cardiosoma armata.—Bruce-Chwatt and Fitz-John, 1951:117,<br />
118, 119.<br />
Gardiosoma.—Bruce-Chwatt and Fitz-John, 1951:120.<br />
Cardisoma.—Uschakov, 1970, fig. 4.<br />
Cardiosoma armatum.—Pauly, 1975:57.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Cape Verde Islands: Porto da Praia, Sao<br />
Tiago, coconut grove, North Pacific Exploring Expedition,<br />
16 (W).<br />
Senegal: Dakar, O. F. Cook, 16*, 1$ (W).<br />
Liberia: Grand Cape Mount, 1882, J. Biittikofer, 36",<br />
39 (L).<br />
Ghana: Accra, 1868-1869, M. Sintenis, lcJ (L). Elmina,<br />
1840-1855, H. S. Pel, 1$ holotype (L, Crust. D.58).<br />
Dahomey: Near Cotonou, Oct 1962, H. Hoestlandt, 16,<br />
19 (L).<br />
Nigeria: Lagos, 16 Aug 1926, A. S. Pearse, 19 (W). Adu,<br />
25 Aug 1926, A. S. Pearse, \6 fragmented (W). Niger delta<br />
between Brass and Port Harcourt, May-Aug 1960, H. J. G.<br />
Beets, 46, 59 (L).<br />
Cameroon: Batanga, A. J. Good, 13 Mar 1931, 16",<br />
19 (W).<br />
Zaire: Banana, Jul 1915, H. Lang, American Museum<br />
Congo Expedition, 18
250<br />
count of the species. To the references in Monod<br />
the following can be added:<br />
<strong>West</strong> Africa: No specific locality (Tiirkay, 1973).<br />
Cape Verde Islands: Baia de Sal-Rei, Boavista (Guinot<br />
and Ribeiro, 1962; Ribeiro, 1964).<br />
Senegal: No specific locality (Tiirkay, 1973). Marigots de<br />
Hann (Bouvier, 1911). Saint-Louis (Monod, 1967; Tiirkay,<br />
1973). Dakar (Monod, 1967). Rtombo [?](Tiirkay, 1973).<br />
Joal; Ngor near Dakar (Humes, 1957).<br />
Guinea: Conakry (Uschakov, 1970).<br />
Sierra Leone: Bunce River, opposite Kamatare Island,<br />
near Freetown (Humes, 1957).<br />
Liberia: No specific locality (Biittikofer, 1890; Johnston,<br />
1906).<br />
Ivory Coast: From market in Abidjan (Forest and Guinot,<br />
1966). Koumassi, near Abidjan (Humes, 1957). Abidjan<br />
(Tiirkay, 1973).<br />
Ghana: No specific locality (Gauld, 1960; Tiirkay, 1973).<br />
Kumasi and Sekondi (Tiirkay, 1973). Sakumo lagoon (Pauly<br />
1975). Accra; Osu Fisheries Station near Accra (Humes,<br />
1957).<br />
Dahomey: No specific locality (De Leersnyder and<br />
Hoestlandt, 1963, 1966).<br />
Nigeria: Lagos (Bruce-Chwatt and Fitz-John, 1951; Tiirkay,<br />
1973). Iru Fisheries Station near Lagos (Humes, 1957).<br />
Cameroon: Douala (Tiirkay, 1973).<br />
Fernando Poo: No specific locality (Tiirkay, 1973).<br />
Sao Tome: "environs de la ville" and Iogoiogo (Forest<br />
and Guinot, 1966).<br />
Gabon: No specific locality (Tiirkay, 1973).<br />
Congo: No specific locality (Rossignol, 1957). Baie de<br />
Pointe-Noire (Rossignol, 1962). Bahua [?] and Pointe-Noire<br />
(Tiirkay, 1973). Loango (Pechiiel-Loesche, 1882). Pointe-<br />
Noire; Songololo River near Pointe-Noire (Humes, 1957).<br />
Zaire: Banana (Dubois, 1957; Tiirkay, 1973).<br />
Angola: Luanda (Tiirkay, 1973). Santo Antonio do Zaire;<br />
Mo^amedes (Guinot and Ribeiro, 1962). Between Cacuaco<br />
and Lobito-Benguela (Hartmann-Schroder and Hartmann,<br />
1974).<br />
Genus Gecarcinus Leach, 1814<br />
Gecarcinus Leach, 1814:430 [type-species: Cancer ruricola Lin-<br />
naeus, 1758, by selection by H. Milne Edwards, 1838, in<br />
1836-1844, pi. 21; gender: masculine].<br />
Geocarcinus Miers, 1886:xiv, 216, 219 [invalid emendation of<br />
Gecarcinus Leach, 1814; type-species: Cancer ruricola Lin-<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
EASTERN ATLANTIC GENERA.—One, Neotroglocarnaeus,<br />
1758; gender: masculine].<br />
cinus, represented in the tropical fauna.<br />
Johngarthia Tiirkay, 1970:335, 341, 343 [type-species: Gecarcinus<br />
planatus Stimpson, 1860, by original designation; EASTERN ATLANTIC SPECIES.—One, Neotroglocar-<br />
gender: feminine].<br />
cinus balssi, reported below. It is not represented<br />
in the Pillsbury collections.<br />
Gecarcinus weileri (Sendler, 1912)<br />
Pelocarcinus weilen Sendler, 1912:191, figs, 4, 5.<br />
Gecarcinus lagostoma.—Capart, 1951:198, fig. 80.—Monod,<br />
1956:461 [not fig. 619 = G. lagostoma sensu stricto].—<br />
Rossignol, 1957:120 [key].—Forest and Guinot, 1966:94.<br />
[Not G. lagostoma H. Milne Edwards, 1837.]<br />
Gecarcinus (Johngarthia) weilen.—Tiirkay, 1973:96, figs. 15-<br />
17, map 2.<br />
Johngarthia weileri.—Tiirkay, 1976c:69, 70, 71 [lectotype designated].<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Fernando Poo: No specific locality, Feb<br />
1959, J. A. Warners, 1$ dry (L).<br />
DESCRIPTION.—Tiirkay, 1973:97.<br />
Figures: Tiirkay, 1973, figs. 15-17.<br />
MEASUREMENTS.—Our specimen has a carapace<br />
width of 60 mm.<br />
BIOLOGY.—This is a terrestrial species from the<br />
Gulf of Guinea, largely restricted to the offshore<br />
islands but also known from the mainland in<br />
Cameroon. Capart (1951) recorded an ovigerous<br />
female in January.<br />
DISTRIBUTION.—<strong>West</strong> Africa, where it has been<br />
taken from two localities in Cameroon and from<br />
the offshore islands of the Gulf of Guinea. Monod<br />
(1956) saw no material. Records in the literature<br />
since 1956 include:<br />
<strong>West</strong> Africa: No specific locality (Tiirkay, 1973).<br />
Cameroon: Bibundi (Tiirkay, 1973; 1976c).<br />
Fernando Poo: No specific locality (Tiirkay, 1973).<br />
Principe: Ilheu Caro^o (Forest and Guinot, 1966).<br />
Sao Tome: No specific locality (Tiirkay, 1973). Ribeiro<br />
do Peixe (Forest and Guinot, 1966). Ilheu Gago Coutinho<br />
(as Hot das Rollas) (Forest and Guinot, 1966; Tiirkay, 1973).<br />
Annobon: No specific locality (Forest and Guinot, 1966;<br />
Tiirkay, 1973).<br />
Family HAPALOCARCINIDAE Caiman, 1900<br />
Lithoscaptes A. Milne Edwards, 1862b: F 10.<br />
HAPALOCARCINIDAE Caiman, 1900:3, 49.<br />
Genus Neotroglocarcinus Fize and Serene, 1957<br />
Neotroglocarcinus Fize and Serene, 1957:134 [type-species:
NUMBER 306 251<br />
Troglocarcmus monodi Fize and Serene, 1955, by original<br />
designation; gender: masculine].<br />
Neotroglocarcinus balssi (Monod, 1956)<br />
Troglocarcmus balssi Monod, 1956:463, 632, figs. 620-627.—<br />
Longhurst, 1958:88.—Gauld, 1960:72.—Monod, 1963,<br />
fig. 37 [no locality].—Crosnier, 1969:535.<br />
Neotroglocarcinus Balssi—Fize and Serene, 1957:135, 136, 141-<br />
143.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Canary Islands: Ponta de Matorra, Fuertaventura,<br />
14 km from Morro, 20-25 m, in Phyllangia mouchezi<br />
(Lacaze-Duthiers, 1897), 27 Jan 1975, M. Grasshoff and F.<br />
Engelhardt, 1 damaged specimen (MP).<br />
<strong>West</strong> Africa: No specific locality, in Asterosimilia marchadi<br />
(Chevalier, 1966), A. R. Longhurst, 1$ ov (BMNH).<br />
Ivory Coast: Off Abidjan, 62 m, 23 Aug 1968, C.R.O.,<br />
Abidjan, 1? (MP).<br />
DESCRIPTION.—Monod, 1956:463.<br />
Figures: Monod, 1956, figs. 620-627.<br />
Male Pleopod: Monod, 1956, fig. 623 (Ghana).<br />
MEASUREMENTS.—Our specimens have carapace<br />
lengths of 3.2 to 3.8 mm; that of the ovigerous<br />
female is 3.8 mm.<br />
REMARKS.—As the single adult female from<br />
Sierra Leone listed by Monod (1956:632) was<br />
collected by A. R. Longhurst, and the single<br />
specimen reported upon by Longhurst (1958:88)<br />
was identified by Monod (see Longhurst, 1958:<br />
5), it seems likely that they are the same specimen<br />
and that an error crept into the ecological data<br />
reported for either: Monod reported the specimen<br />
as an epibiont of Cidaris cidaris (Linnaeus) from<br />
68 m depth, while Longhurst listed it as an<br />
epibiont of Eucidaris tribuloides (Lamarck) from<br />
200 m. (Under Eucidaris tribuloides, however,<br />
Longhurst (1958:101) only mentioned two specimens,<br />
both from 25-88 m depth, and under<br />
Cidaris cidaris 6 specimens from depths between<br />
88 and 132 m.) Longhurst (1958:7) reported that<br />
his collection finally was deposited in the British<br />
Museum (Natural History). Manning examined<br />
a specimen of this species of the British Museum<br />
collection, viz., the above-mentioned ovigerous<br />
female labeled "<strong>West</strong> Africa" and collected by<br />
Longhurst; this specimen is said to be taken from<br />
the coral Asterosimilia marchadi (Chevalier). The<br />
fact that both Cidaris and Eucidaris are very unlikely<br />
hosts for the species, and the conflicting<br />
information in the ecological data suggest that<br />
these should be regarded with the utmost reserve,<br />
and the possibility is not precluded that both the<br />
Monod (1956) and the Longhurst record pertain<br />
to the ovigerous female now in the British Museum,<br />
and that this specimen was collected in<br />
Asterosimilia marchadi in a depth of 68 m, at<br />
08°25'N, 14°18'W (off Freetown, Sierra Leone)<br />
on a sandy bottom on 22 Feb 1956 {Cape St. Mary<br />
StaMBl/A3).<br />
BIOLOGY.—This is a sublittoral species usually<br />
associated with corals. It has been taken with<br />
Dendrophyllia, in 44 m (Gauld, 1960); on rocks<br />
with gorgonians, probably associated with Phyllangia,<br />
in 10 m (Crosnier, 1969); on Eucidaris<br />
tribuloides (?) in 200 m (Longhurst, 1958); on<br />
Cidaris cidaris (?) in 68 m (Monod, 1956); and<br />
with Phyllangia mouchezi and Asterosimilia marchadi.<br />
Ovigerous females have been recorded in January<br />
(Crosnier, 1969; present paper); ova measured<br />
0.4 mm in the ovigerous female we examined.<br />
DISTRIBUTION.—<strong>West</strong> Africa, from localities in<br />
the Canary Islands, off <strong>West</strong> Africa proper from<br />
localities between Sierra Leone and the Congo,<br />
and from Ilheu Gago Coutinho (as I. das Rolas),<br />
near Sao Tome in the Gulf of Guinea (Balss,<br />
1922) in depths between 10 and 200 m. It has not<br />
previously been recorded from the Canary Islands<br />
or the Ivory Coast. Monod's specimens came from<br />
Ghana; since 1956 the species has been recorded<br />
from the following:<br />
Sierra Leone: No specific locality, 200 m (Longhurst,<br />
1958); 08°25'N, 14°18'W, 68 m (Monod, 1956).<br />
Ghana: Off Accra, 44 m (Gauld, 1960).<br />
Congo: Pointe-Noire, ca. 10 m (Crosnier, 1969).<br />
Family HYMENOSOMATIDAE MacLeay, 1838<br />
HYMENOSOMIDAE MacLeay, 1838:68 [corrected to Hymenosomatidae<br />
by Stebbing, 1905:49].<br />
HYMENICINAE Dana, 1851c:290.<br />
EASTERN ATLANTIC GENERA.—Two, Elamena
252<br />
and Hymenosoma, both represented by <strong>West</strong> <strong>African</strong><br />
species.<br />
EASTERN ATLANTIC SPECIES.—Two, both listed<br />
below. This family was not represented in the<br />
Pillsbury collections.<br />
Genus Elamena H. Milne Edwards, 1837<br />
Elamena H. Milne Edwards, 1837:33 [type-species: Hymenosoma<br />
mathaei Desmarest, 1825, by monotypy; gender: feminine].<br />
Trigonoplax H. Milne Edwards, 1853:224 [type-species: Ocypode<br />
(Elamene) unguiformis de Haan, 1839, by monotypy;<br />
gender: feminine].<br />
Elamena (Trigonoplax) gordonae Monod, 1956<br />
Elamena (Trigonoplax) gordonae Monod, 1956:469, figs. 629-<br />
637 [between Conakry, Guinea and Monrovia, Liberia,<br />
30-40 m; Sierra Leone]; 1963, fig. 40 [no locality].<br />
Elamena gordonae.—Uschakov, 1970:455 [listed; Guinea].<br />
DISTRIBUTION.—<strong>West</strong> Africa, from the localities<br />
cited above.<br />
Genus Hymenosoma Desmarest, 1825<br />
Hymenosoma Desmarest, 1825:163 [type-species: Hymenosoma<br />
orbiculare Desmarest, 1825, by selection by H. Milne Edwards,<br />
1842, in 1836-1844, pi. 35: fig. 1; gender: neuter].<br />
Leachium MacLeay, 1838:68 [type-species: Hymenosoma orbiculare<br />
Desmarest, 1825, by original designation; gender:<br />
neuter].<br />
Centridion Gistel, 1848:viii [substitute name for Leachium<br />
MacLeay, 1838; type-species: Hymenosoma orbiculare Desmarest,<br />
1825; gender: neuter].<br />
Hymenosoma orbiculare Desmarest, 1825<br />
Hymenosoma orbiculare.—Capart, 1951:61, fig. 18 [Angola;<br />
South-<strong>West</strong> Africa].—Monod, 1956:468, fig. 628 [Gabon<br />
(?); South-<strong>West</strong> Africa].—Penrith and Kensley, 1970a:<br />
209, 232 [South-<strong>West</strong> Africa].<br />
DISTRIBUTION.—Known with certainty from localities<br />
in Angola, South-<strong>West</strong> Africa, and South<br />
Africa. The record from Gabon is questionable.<br />
Family MAJIDAE Samouelle, 1819<br />
MAIADAE Samouelle, 1819:88 [corrected to Majidae by Neumann,<br />
1878:5].<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
MACROPODIADAE Samouelle, 1819:90.<br />
INACHIDAE MacLeay, 1838:56 [given preference over Macropodiadae<br />
Samouelle, 1819, by International Commission<br />
on Zoological Nomenclature, Opinion 763, 1966; name<br />
400 on Official List].<br />
EPIALTIDAE MacLeay, 1838:56.<br />
HUENIDAE MacLeay, 1838:56.<br />
EURYPODIDAE MacLeay, 1838:56.<br />
MITHRACIDAE MacLeay, 1838:56.<br />
LEPTOPODIADAE Bell, 1844:1.<br />
MACROCHEIRINAE Dana, 1851a:427.<br />
PISINAE Dana, 1851a:428 [name 368 on Official List, date<br />
and citation erroneous there].<br />
LIBININAE Dana, 1851a:429.<br />
PRIONORHYNCHINAE Dana, 1851a:429.<br />
MICIPPINAE Dana, 1851a:429.<br />
CHORININAE Dana, 1851a:429.<br />
PYRINAE Dana, 1851a:430.<br />
OTHONINAE Dana, 1851a:430.<br />
SALACINAE Dana, 1851a:430.<br />
CYCLACINAE Dana, 185la:431.<br />
TYCHIDAE Dana, 1851a:431.<br />
CRIOCARCININAE Dana, 185 la:431.<br />
CAMPOSCINAE Dana, 1851a:431.<br />
AMATHINAE Dana, 1851a:431.<br />
STENORHYNCHINAE Dana, 1851a:432.<br />
ACHAEINAE Dana, 1851a:432.<br />
PERICERIDAE Dana, 1851a:432.<br />
PARAMICIPPINAE Dana, 1851a:432.<br />
INACHOIDINAE Dana, 1851a:432.<br />
MENAETHINAE Dana, 1851a:433.<br />
STENOCIOPINAE Dana, 1851a:433.<br />
ONCININEA Dana, 1852b:77.<br />
ONCINOPIDAE Stimpson, 1858d:222.<br />
LEPTOPINAE Stimpson, 1871a: 109.<br />
NAXIINAE Stimpson, 1871a: 114.<br />
COLLODINAE Stimpson, 1871a: 119.<br />
ANOMALOPINAE Stimpson, 1871a: 124.<br />
ACANTHONYCHINAE Stimpson, 1871a: 127.<br />
IXIONINAE Neumann, 1878:10.<br />
ACANTHOPHYRINAE Neumann, 1878:10.<br />
PICROCERINAE Neumann, 1878:12.<br />
PODOCHELINAE Neumann, 1878:13.<br />
CYPHOCARCININAE Neumann, 1878:15.<br />
EURYNOMINAE Neumann, 1878:17.<br />
MICRORHYNCHINAE Miers, 1879a:651.<br />
SCHIZOPHRYSINAE Miers, 1879a:659.<br />
LISSOIDA Alcock, 1895:161.<br />
BLASTIDAE Stebbing, 1902:2.<br />
MAMAIIDAE Stebbing, 1905:22.<br />
OPHTHALMIINAE Balss, 1929:6.<br />
HYASTENIINAE Balss, 1929:8, 14.<br />
MACROCOELOMINAE Balss, 1929:8, 16, 20.<br />
OREGONHNAE Garth, 1958:134.
NUMBER 306 253<br />
EASTERN ATLANTIC GENERA.—Twenty, of which<br />
the following 15 are represented by tropical species:<br />
Acanthonyx, Achaeus, Apiomithrax, Calypsachaeus,<br />
new genus, Capartiella, new genus, Dorhynchus,<br />
Ergasticus, Eurynome, Herbstia, Inachus, Macropodia,<br />
Maja, Micropisa, Pisa, and Stenorhynchus. The<br />
other genera are as follows:<br />
Anamathia Smith (1885:493). Substitute name<br />
for Amathia Roux (1828, pi. 3), an invalid junior<br />
homonym of Amathia Lamouroux, 1812; type-species:<br />
Amathia rissoana Roux, 1828, by monotypy;<br />
gender: feminine; name 1606 on Official List.<br />
Hyas Leach (1814:431). Type-species: Cancer araneus<br />
Linnaeus, 1758, by monotypy; gender: masculine.<br />
Hyastenus White (1847b:56). Type-species: Hyastenus<br />
sebae White, 1847, by monotypy; gender:<br />
masculine.<br />
Lissa Leach (1815b:69). Type-species: Cancer<br />
chiragra Fabricius, 1775, by monotypy; gender:<br />
feminine; name 1302 on Official List.<br />
Rochinia A. Milne Edwards (1875, in 1873-<br />
1881:86). Type-species: Rochinia gracilipes A.<br />
Milne Edwards, 1875, by monotypy; gender: feminine;<br />
name 1647 on Official List.<br />
EASTERN ATLANTIC SPECIES.—Sixty-six, of<br />
which 49 occur in tropical waters. Monod (1956)<br />
recorded the following species.<br />
Name in Monod<br />
Maja squinado<br />
Maja verrucosa<br />
Maja goltziana<br />
Eurynome aspera<br />
Herbstia rubra<br />
Herbstia rubra<br />
Pisa tetraodon<br />
Pisa nodipes<br />
Pisa gibbsi<br />
Pisa carinimana<br />
Micropisa ovata<br />
Apiomithrax violaceus<br />
Apiomithrax bocagei<br />
Acanthonyx lunulatus<br />
Dorhynchus thomsoni<br />
Ergasticus clouei<br />
Inachus angolensis<br />
Inachus dorsettensis<br />
Inachus guentheri<br />
Current Name<br />
Maja squinado<br />
Maja crispata<br />
Maja goltziana<br />
Eurynome aspera*<br />
Herbstia rubra<br />
Herbstia condyliata*<br />
Pisa tetraodon<br />
Pisa nodipes<br />
Pisa armata*<br />
Pisa carinimana*<br />
Micropisa ovata<br />
Apiomithrax violaceus*<br />
Apiomithrax bocagei<br />
Acanthonyx lunulatus<br />
Dorhynchus thomsoni<br />
Ergasticus clouei<br />
Inachus angolensis<br />
Inachus nanus, new species*<br />
Inachus guentheri<br />
Inachus phalangium<br />
Inachus thoracicus<br />
Inachus aguiari<br />
Inachus leptochirus<br />
Physachaeus (?) longipes<br />
Achaeus crane hi<br />
Achaeus fores ti<br />
Achaeus sp.<br />
Achaeus monodi<br />
Macropodia gilsoni<br />
Macropodia macrocheles<br />
Macropodia rostrata<br />
Macropodia straeleni<br />
Stenorhynchus seticomis<br />
Inachus phalangium<br />
Inachus biceps, new species*<br />
Inachus aguiarii<br />
Inachus leptochirus<br />
Capartiella longipes*<br />
Achaeus cranchii<br />
Achaeus fores ti*<br />
Achaeus trifalcatus<br />
Achaeus monodi<br />
Macropodia gilsoni*<br />
Macropodia macrocheles*<br />
Macropodia spinulosa*<br />
Macropodia straeleni*<br />
Stenorhynchus lanceolatus*<br />
Representatives of 24 species were taken by the<br />
Pillsbury, eight of them previously undescribed.<br />
The following species occur outside of the tropical<br />
region:<br />
Achaeus gracilis O. Costa, 1839 (= Achaeus gordonae<br />
Forest and Zariquiey Alvarez, 1955). Forest<br />
and Zariquiey Alvarez (1955:68, figs. 2, 4, 6, 8)<br />
pointed out that two species of Achaeus could be<br />
recognized in the Mediterranean, A. cranchii<br />
Leach, 1817, and a second species that they<br />
named Achaeus gordonae. In their account Forest<br />
and Zariquiey Alvarez noted that O. Costa (1839,<br />
in Costa and Costa, 1838-1871:25) in his Fauna<br />
del Regno di Napoli, named a small crab, Macropodia<br />
gracilis, which has the general aspect of an Achaeus<br />
and that Costa also had recognized A. cranchii.<br />
Forest and Zariquiey Alvarez (1955:66) considered<br />
Costa's M. gracilis to be unidentifiable ("espece<br />
douteuse") and they did not synonymize it<br />
with A. cranchii. An examination of Costa's account<br />
and figure (pi. 3: fig. lA,b) suggests to us<br />
that he was dealing with an Achaeus, one in which<br />
the rostral teeth are almost contiguous and the<br />
hepatic lobes of the carapace are poorly developed,<br />
characters which Forest and Zariquiey Alvarez<br />
used to separate A. gordonae from A. cranchii.<br />
In addition, in Costa's figure 1A the length/width<br />
ratio of the illustrated specimen is 1.44; this ratio<br />
was given by Forest and Zariquiey Alvarez as<br />
1.16 to 1.38 in A. cranchii, 1.32 to 1.55 in A.<br />
gordonae. Thus this ratio in A. gracilis is larger than<br />
that of A. cranchii but fits well within the range<br />
reported for A. gordonae. We believe that Achaeus
254<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
gordonae must be considered a synonym of Macro- Macropodia longirostris (Fabricius, 1775). Medipodia<br />
gracilis O. Costa, 1839. Achaeus gracilis occurs terranean; sublittoral, between 4 and 50 m<br />
in the Mediterranean and adjacent Atlantic; sub- (Zariquiey Alvarez, 1968).<br />
littoral, to about 20 m (Zariquiey Alvarez, 1968). Macropodia rostrata (Linnaeus, 1761). Northeast-<br />
Anamathia rissoana (Roux, 1828). Azores and ern Atlantic, from Norway southward at least to<br />
eastern Mediterranean; sublittoral to about 400 Mediterranean; sublittoral; tropical <strong>West</strong> Africa<br />
m (Zariquiey Alvarez, 1968).<br />
records are referable to M. spinulosa (Zariquiey<br />
Eurynome spinosa Hailstone, 1835. Eastern Atlan- Alvarez, 1968; Christiansen, 1969; Forest, 1978).<br />
tic, from Scandinavia and British Isles southward Macropodia tenuirostris (Leach, 1814). Northeast-<br />
to NW Spain, Azores, Mediterranean; sublittoral, ern Atlantic, from the Faroes southward to Por-<br />
from about 180 to about 400 m (Zariquiey Altugal; sublittoral, to more than 150 m (Zariquiey<br />
varez, 1968; Christiansen, 1969).<br />
Alvarez, 1968; Christiansen, 1969; Forest, 1978).<br />
Hyas araneus (Linnaeus, 1758). Eastern Atlantic Pisa corallina (Risso, 1816). Mediterranean; sub-<br />
from Barents Sea southward to NW France; also littoral, usually in less than 10 m (Zariquiey<br />
western Atlantic; sublittoral, shallow water to Alvarez, 1968).<br />
more than 300 m (Christiansen, 1969).<br />
Pisa muscosa (Linnaeus, 1758). Mediterranean;<br />
Hyas coarctatus Leach, 1815. Eastern Atlantic sublittoral, from 4-5 to about 40 m (Zariquiey<br />
from Barents Sea and Arctic southward to NW Alvarez, 1968).<br />
France; also western Atlantic, northern Pacific; Rochinia carpenteri (Thomson, 1873). Eastern At-<br />
sublittoral, between 1 and 500 m (Christiansen, lantic, from Iceland and the Faroes southward to<br />
1969).<br />
NW Africa, Azores; sublittoral, from about 180<br />
Hyastenus hilgendorfi De Man, 1887. An Indo- to about 1300 m (Zariquiey Alvarez, 1968; Chris-<br />
<strong>West</strong> Pacific immigrant into the eastern Meditertiansen, 1969).<br />
ranean, now known from the coasts of Israel We believe that the following species, tenta-<br />
(Lewinsohn and Holthuis, 1964) and Egypt (Ratively recorded by Monod (1956) from <strong>West</strong> Afmadan<br />
and Dowidar, 1976).<br />
rica, are based on erroneously labeled specimens<br />
Inachus communissimus Rizza, 1839. Mediterra- (each is known from a single specimen thought to<br />
nean and adjacent Atlantic, at least as far as be <strong>West</strong> <strong>African</strong>) and should not be included in<br />
Portugal; sublittoral, 15 to 24 m (Zariquiey Al- the <strong>West</strong> <strong>African</strong> fauna:<br />
varez, 1968).<br />
Libinia erinacea A. Milne Edwards, 1879. A west-<br />
Lissa chiragra (Fabricius, 1775). Mediterranean, ern Atlantic species recorded from "<strong>West</strong> Africa"<br />
sublittoral, between 20 and 40 m (Zariquiey Al- as Libinia dubia by Streets (1870:105) and subsevarez,<br />
1968).<br />
quently recorded by Rathbun (1925:321). Monod<br />
Macropodia czernjawskii (Brandt, 1880). Medit- (1956:514, fig. 705), who examined the specimen,<br />
teranean; sublittoral, between 10 and 30 m (Zar- identified it with L. erinacea.<br />
iquiey Alvarez, 1968).<br />
Notolopas brasiliensis Miers, 1886. A western At-<br />
Macropodia deflexa Forest, 1978. Northeastern lantic species reported by Monod (1956:513, figs.<br />
Atlantic, from southern England to Portugal; 703, 704), based on a specimen collected off<br />
sublittoral, shallow water to 20 m (Forest, 1978). South-<strong>West</strong> Africa in 1900.<br />
Macropodia intermedia Bouvier, 1940. (See page Rochinia gracilipes A. Milne Edwards, 1875. A<br />
300).<br />
western Atlantic species included by Monod<br />
Macropodia linaresi Forest and Zariquiey Al- (1956:516, figs. 706-708), on the basis of a single<br />
varez, 1964. Eastern Atlantic, from southern England<br />
and France to Spain, Mediterranean; sublittoral,<br />
between 30 and 80 m (Zariquiey Alvarez,<br />
1968; Forest, 1978).<br />
specimen from Gabon collected by Heurtel.<br />
Much of the material from Gabon in this collection<br />
apparently has been erroneously labeled.<br />
REMARKS.—In our account of this family we
NUMBER 306 255<br />
have departed from our usual format in presenting<br />
synonymies for several Mediterranean-North<br />
Atlantic species which may not be represented in<br />
the tropical fauna. The occurrence of three such<br />
both coasts of the Americas (Rathbun, 1925;<br />
Garth, 1958); A. lunulatus (Risso, 1816) (with A.<br />
brevifrons A. Milne Edwards, 1869, as a synonym),<br />
from the eastern Atlantic (see p. 256, and Zari-<br />
species, Pisa armata, Acanthonyx lunulatus, and Herbsquiey Alvarez, 1968 for Mediterranean refertia<br />
condyliata, is well documented, but in the cases ences); A. sanctaehelenae Chace, 1966, from Saint<br />
of the latter two species older records may well Helena; A. simplex Dana, 1852, from the Pacific;<br />
pertain to other species recognized herein. and three species from the western Indian Ocean:<br />
The occurrence of several other northern spe- A. consobrinus A. Milne Edwards, 1862, A. limbatus<br />
cies off tropical <strong>West</strong> Africa should be verified: A. Milne Edwards, 1862, and A. elongatus Miers,<br />
Achaeus cranchii, Inachus aguiarii, I. dorsettensis (trop1877<br />
(Stephensen, 1945; Barnard, 1950; Guinot,<br />
ical records of which are referred to /. norms, new 1967a).<br />
species, p. 291), /. leptochirus (tropical records are Acanthonyx closely resembles Dehaanius Macreferred<br />
to /. biceps, new species, p. 285), /. phal- Leay, 1838, the only difference mentioned in the<br />
angium, I. thoracicus, Macropodia intermedia, M. lonliterature<br />
being the presence of a seven-segmented<br />
gipes, and Pisa nodipes.<br />
abdomen in the male in the latter, and an abdomen<br />
with six or less somites in the former. Ste-<br />
Subfamily EPIALTINAE MacLeay, 1838<br />
phensen (1945) and Barnard (1950) both suggest<br />
that this character is unreliable. The status of the<br />
Genus Acanthonyx Latreille, 1828 two genera requires clarification.<br />
According to accounts in the literature (cited<br />
Acanthonyx Latreille, 1828a:698 [type-species: Maia lunulata<br />
Risso, 1816, by monotypy; gender: masculine; name 1603<br />
on Official List, there erroneously dated 1827].<br />
Gonosoma Costa, 1844:69 [type-species: Gonosoma viridis Costa,<br />
1844 (= Acanthonyx viridis Costa, 1838), a subjective junior<br />
above), the four Indo-<strong>West</strong> Pacific species of Acanthonyx<br />
can be distinguished from those occurring<br />
in the Atlantic as follows: A. simplex lacks dorsal<br />
spines or tubercles on the carapace; A. dentatus<br />
synonym of Maia lunulata Risso, 1816, by monotypy; gen- has a strong tooth on the orbital margin; A.<br />
der: neuter].<br />
consobrinus has a tooth on the orbital margin and<br />
Peltinia Dana, 1851d:272 [type-species: Peltinia scutiformis<br />
Dana, 1851, a subjective junior synonym of Acanthonyx<br />
petiverii H. Milne Edwards, 1834, by present selection;<br />
gender: feminine].<br />
four lateral projections on the carapace; and A.<br />
limbatus has much stronger lateral projections on<br />
the carapace.<br />
In his account of A. sanctaehelenae, Chace (1966:<br />
REMARKS.—Acanthonyx is dated from 1827 in 13) provided good comparative illustrations of<br />
the Official List of Generic Names in Zoology. So far the three species then known from the Atlantic,<br />
as we can determine, this is an error that origi- A. lunulatus, A. petiverii, and A. sanctaehelenae.<br />
nated in Neave, Nomenclator Zoologicus, in which Material of Acanthonyx available to us from<br />
1827 is given. Other genera named in the same <strong>West</strong> <strong>African</strong> localities includes one new species<br />
article by Latreille are correctly dated 1828 in from the Pillsbury collections, a second new species<br />
Neave.<br />
from Point-Noire, and A. brevifrons from the Cape<br />
Until now Acanthonyx included seven species, as Verde Islands. We recognize four species from the<br />
follows: A. petiverii H. Milne Edwards, 1834, from eastern Atlantic.<br />
Key to Eastern Atlantic Species of Acanthonyx<br />
1. Carapace with only 2 lateral lobes. [Rostral teeth short. 8-13 tubercles<br />
ventrally on dactyli of pereiopods] A. brevifrons<br />
Carapace with 3 lateral lobes 2
256<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
2. Orbital margin with blunt tooth or projection. [Size very small, adults with<br />
carapace lengths of less than 4 mm. 6-8 tubercles ventrally (usually 6; 3<br />
or 4 in juveniles) on dactyli of walking legs] . . . . A. minor, new species<br />
Orbital margin smooth, evenly concave 3<br />
3. Rostral teeth strongly depressed. Rostral sinus V-shaped. Merus of claw<br />
with 1 tubercle. [Size very small, adults with carapace lengths of<br />
less than 5 mm. 4-6 tubercles ventrally on dactyli of walking legs]<br />
A. depressifrons, new species<br />
Rostral teeth not markedly depressed. Rostral sinus U-shaped. Merus of<br />
claw with 2 tubercles. [Size moderate to large, adults with carapace<br />
lengths of 15 mm or more. Males with 11-17 and females with 8-12<br />
tubercles ventrally on dactyli of walking legs] A. lunulatus<br />
Acanthonyx brevifrons A. Milne Edwards,<br />
1869<br />
FIGURES 60a, 61<br />
Acanthonyx brevifrons A. Milne Edwards, 1869:353.—A. Milne<br />
Edwards and Bouvier, 1894:12; 1900:152.—Bouvier,<br />
1940:349.<br />
?Acanthonyx lunulatus.—Miers, 1886:43.—Barrois, 1888:9. [See<br />
p. 258, references to A. lunulatus from Cape Verde Islands;<br />
not Acanthonyx lunulatus (Risso, 1816)].<br />
?Acanthonyx lunulatus var. brevifrons.—Chapman and Santler,<br />
1955:375.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Cape Verde Islands; between Ilheu Branco<br />
and Ilheu Raso, 110-180 m, sand and rock, Talisman, 27 Jul<br />
1883, 1?, ljuv(W).<br />
DESCRIPTION.—Carapace (Figure 61a) pearshaped,<br />
length in midline 1/5 longer than maximum<br />
breadth. Dorsal surface of carapace smooth,<br />
lacking distinct pubescence (in material studied).<br />
Hepatic lobe scarcely produced anteriorly. One<br />
prominent rounded lobe on branchial region,<br />
lateral margin between hepatic and branchial<br />
lobes straight, subparallel or slightly convergent<br />
posteriorly. Rostral teeth, preorbital, hepatic, and<br />
branchial lobes each with terminal tuft of stout<br />
setae. Rostral sinus (Figure 61£,
NUMBER 306 257<br />
FIGURE 60.—Carapaces of <strong>West</strong> <strong>African</strong> species of Acanthonyx in lateral view: a, A. brevifrons A.<br />
Milne Edwards, female, cl 8.8 mm, Cape Verde Islands; b, A. depressifrons, new species, holotype,<br />
male, cl 4.0 mm, Pointe-Noire; c, A. lunulatus (Risso), female, cl 6.8 mm, Tunis; d, A. minor, new<br />
species, paratype, male, cl 2.6 mm, Pilhbury Sta 271.<br />
Cape Verde Islands by A. Milne Edwards (1869);<br />
no depth data were given. Miers (1886:43) recorded<br />
a small female of A. lunulatus from Sao<br />
Vicente and noted that it approached the "species<br />
or variety Acanthonyx brevifrons A. Milne Edwards,<br />
in the form of the front, but there are indications<br />
of three antero-lateral teeth, and the carapace, as<br />
in Acanthonyx lunulatus, bears several tufts of setae."<br />
Miers' material, which could not be located, may<br />
be referable to A. lunulatus. A. Milne Edwards and<br />
Bouvier (1894, 1900) gave additional differences<br />
between this species and A. lunulatus, but did not<br />
give figures. Balss (1914) identified material from<br />
Annobon with this species, but in 1922 he synonymized<br />
A. brevifrons with A. lunulatus. We suspect<br />
that the material reported by Balss from Annobon<br />
actually belongs to a new species, A. minor (p.<br />
261). Bouvier (1940:348) suggested that A. brevifrons<br />
should be considered as a race of A. lunulatus,<br />
and Chapman and Sander (1955) identified material<br />
from the Azores as A. lunulatus var. brevifrons.<br />
Monod (1956) synonymized A. brevifrons with A.<br />
lunulatus, and figured a male of the latter species<br />
from Dakar. In his list of material, Monod noted<br />
that his figured specimen, a large male (cl 15<br />
mm), had an obsolete intermediate tooth on the<br />
carapace and corresponded to "f. brevifrons". The<br />
figured specimen appears to be a typical lunulatus,<br />
as does the specimen from Mauritania figured<br />
earlier by Monod (1933b, fig. 7D). None of the<br />
material of A. lunulatus that we have examined<br />
approaches our material of A. brevifrons in short-<br />
FIGURE 61.—Acanthonyx brevifrons A. Milne Edwards, Cape<br />
Verde Islands. Female, cl 8.8 mm: a, carapace; b, front,<br />
ventral view; c, cheliped; d, fifth pereiopod. Juvenile, cl 5.4<br />
mm: e, front, ventral view;/, chela.
258<br />
ness of rostral teeth or in the total suppression of<br />
the second lateral tooth on the carapace.<br />
Bouvier (1940) noted that A. lunulatus occurred<br />
to a depth of about 20 m; it is usually taken in<br />
shallow water on algae. In the Mediterranean it<br />
often occurs on the algal genus Cystoseira. Two of<br />
the records for A. brevifrons given by A. Milne<br />
Edwards and Bouvier (1900) are from depths of<br />
75 m and 110-180 m (our material is from the<br />
latter collection). It seems likely that A. brevifrons<br />
occurs in much deeper water than does A. lunulatus,<br />
and it is possible that both occur together in<br />
the Cape Verde Islands. All of the material from<br />
the Cape Verde Islands, the Azores, and Madeira<br />
(below) should be reexamined to determine which<br />
species occurs in the localities.<br />
BIOLOGY.—Little information is available on<br />
the ecology of this species. Our specimens were<br />
taken by the Talisman on sand and rock in 110-<br />
180 m; no information was given for the type. A.<br />
Milne Edwards and Bouvier (1900) also recorded<br />
material from 75 m but did not record the habitat.<br />
In 1894 these same authors recorded the<br />
species from algae and rocks in 10 m. Shallower<br />
water records for this species, as well as those<br />
based on material collected in the alga Cystoseira,<br />
a favorite habitat for A. lunulatus, may be based<br />
on this latter species.<br />
DISTRIBUTION.—Acanthonyx brevifrons has been<br />
reported from localities in the Azores, Madeira,<br />
and the Cape Verde Islands. At least some of the<br />
following records may be referable to A. lunulatus;<br />
all should be verified. Records in the literature<br />
include:<br />
Azores: No specific locality (A. Milne Edwards and Bouvier,<br />
1900; Bouvier, 1940; Barrois, 1888). E coast of Ilha das<br />
Flores, 39°26'30"N, 33°29'15"W of Paris (= 31°09'15"W of<br />
Greenwich), intertidal, and Graciosa, Ilheu da Praia, 39°-<br />
03'15"N, 30°18'15"W of Paris (= 27°48'15"W of Greenwich),<br />
beach, 10 m (A. Milne Edwards and Bouvier, 1894).<br />
Pasteleiro, Feteira, and Almoxarife [?], Ilha do Faial; and<br />
Madalena, Ilha do Pico (Chapman and Santler, 1955).<br />
Madeira: No specific locality (Chapman and Santler,<br />
1955).<br />
Cape Verde Islands: Sao Vicente (A. Milne Edwards,<br />
1869; Miers, 1886). Channel between Sao Vicente and Santo<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Antao, 75 m, and between Ilheu Branco and Ilheu Raso,<br />
110-180 m (A. Milne Edwards and Bouvier, 1900).<br />
Acanthonyx depress!frons, new species<br />
FIGURES 60i, 62<br />
MATERIAL EXAMINED.—Pilkbwy Material. None.<br />
Other Material: Congo: Pointe-Noire, 5-10 m, in lobster<br />
nets, 24 Jan 1967, J. Marteau, 26 (larger male is holotype),<br />
4$ (L, W).<br />
DESCRIPTION.—Carapace (Figure 62a) smooth,<br />
length 1.20-1.35 times greatest width, greatest<br />
width at level of anterolateral tooth. Surface with<br />
2 protogastric, 1 gastric, and 1 cardiac tubercle<br />
in midline, lateral branchial tubercle also present.<br />
All dorsal tubercles with 1 or 2 broad setae.<br />
Rostral sinus (Figure 62a,b) V-shaped, rostral<br />
teeth noticeably depressed in lateral view (Figure<br />
606), teeth blunt. Carapace, posterior to base of<br />
rostrum, with usual 2 diverging rows of curved,<br />
hook-shaped hairs. Preorbital tooth blunt,<br />
rounded, much shorter than rostral teeth. Orbital<br />
margin curving evenly to anterior margin of first<br />
anterolateral tooth, which is not markedly set off<br />
as in A. lunulatus. No tooth present on orbital<br />
margin as in A. minor. First anterolateral tooth<br />
much larger than second and third, which are<br />
distinct and subequal in size. Carapace lacking<br />
subhepatic tubercle. Eyes small.<br />
Basal segment of antennal peduncle (Figure<br />
62b) unarmed. Antennal peduncle extending to<br />
or beyond apices of rostral teeth.<br />
Chelipeds (Figure 62c) small in both sexes.<br />
Fingers about as long as palm, cutting edges<br />
dentate, gape slight. Palm smooth. Carpus with<br />
low dorsal crest; carpus about as long as palm,<br />
much shorter than merus. Merus with single<br />
blunt tubercle on basal half.<br />
Dactyli of walking legs (Figure 62d) broad,<br />
opposable margins with 4-6 pairs of teeth. Second<br />
pereiopod with dactyl shorter than propodus.<br />
Propodus about 3 times as long as high, outer<br />
surface with disc-like projection extending over<br />
base of dactylus; posterior margin of propodus<br />
with broad tubercle, ornamented with large setae,<br />
near midline. Carpus about as long as propodus,
NUMBER 306 259<br />
a<br />
FIGURE 62.—Acanthonyx depressifrons, new species. Female paratype, cl 3.6 mm, Pointe-Noire: a,<br />
carapace. Holotype, male, cl 4.0 mm, Pointe-Noire: b, front, dorsal view; c, cheliped; d, fifth<br />
pereiopod; e, abdomen;/, gonopod.<br />
much shorter than merus. Third and fourth pereiopods<br />
shorter than, but otherwise similar to,<br />
second. Fifth pereiopod shortest. Dactylus about<br />
as long as propodus. Propodus stout, slightly more<br />
than 2 times as long as wide; tubercles on posterior<br />
margin low, with slender setae and club-like<br />
hairs. Carpus shorter than propodus. Merus<br />
longer than propodus.<br />
Male abdomen (Figure 62*) narrow. Telson<br />
triangular, length and width subequal. Sixth somite<br />
expanded distally. Male pleopod as illustrated<br />
(Figure 62/).<br />
MEASUREMENTS.—Carapace lengths of males<br />
3.5 to 4.0 mm, of females 3.3 to 4.3 mm.<br />
REMARKS.—Acanthonyx depressifrons differs from<br />
the other three eastern Atlantic species of the<br />
genus in having the rostrum strongly depressed<br />
and in having only one tubercle on the merus of<br />
the cheliped. It further differs from A. brevifrons in<br />
having three distinct lateral lobes on the carapace<br />
and fewer tubercles on the dactyli of the walking<br />
legs; from A. lunulatus in having the rostral sinus<br />
V-shaped and fewer tubercles on the dactyls of<br />
the pereiopods; and from A. minor, new species, in<br />
lacking a projection on the orbital margin and in<br />
having its greatest width at the level of the anterolateral<br />
tooth of the carapace in both sexes and<br />
at all sizes. Like A. minor, and perhaps A. brevifrons,<br />
A. depressifrons is a much smaller species than A.<br />
lunulatus, which can attain 15 mm or more in<br />
carapace length.<br />
Acanthonyx depressifrons is a much smaller species<br />
than A. petiverii or A. sanctaehelenae, both of which<br />
as adults may attain a carapace length of 18 mm<br />
(Rathbun, 1925; Chace, 1966). It also differs from<br />
A. petiverii in having the rostral sinus V-shaped<br />
rather than broadly U-shaped and from A. sanctaehelenae<br />
in having one rather than two lobes or<br />
tubercles on the outer margin of the merus of the<br />
chelipeds.<br />
TYPE-LOCALITY.—Off Pointe-Noire, Congo.<br />
DISPOSITION OF TYPES.—The holotype is the
260 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
larger male, carapace length 4.0 mm (USNM<br />
127191); three female paratypes (USNM 169535)<br />
also have been deposited in the collections of the<br />
<strong>Smithsonian</strong> <strong>Institution</strong>. A male and a female<br />
paratype have been deposited in the Rijksmuseum<br />
van Natuurlijke Histoire, Leiden.<br />
ETYMOLOGY.—The specific epithet is from the<br />
Latin and refers to the characteristic depressed<br />
front of this species.<br />
DISTRIBUTION.—Known only from the typelocality,<br />
Point-Noire, Congo, where it was taken<br />
at a depth of 5-10 m.<br />
Acanthonyx lunulatus (Risso, 1816)<br />
FIGURES 60C, 63<br />
Acanthonyx lunulatus.—Capart, 1951:84, fig. 26.—Monod,<br />
1956:517, figs. 709, 710.—Rossignol, 1957:78, 116<br />
[key].—Forest and Games, 1960:356.—Gauld, 1960:72.—<br />
Rossignol, 1962:122.—Guinot and Ribeiro, 1962:76.—<br />
Ribeiro, 1964:20.—Chace, 1966, fig. 13a-d [Mediterranean].—Forest<br />
and Guinot, 1966:106.—Zariquiey Alvarez,<br />
1968:466, figs. 7d, 153e,f, 154i,j [Spain; references].—Kensley,<br />
1970:181.—Penrith and Kensley,<br />
1970b:252, 260.—Uschakov, 1970:455 [listed].<br />
SYNONYMS.—Maia glabra Latreille, 1836; Acanthonyx<br />
viridis O. Costa, 1838; Gonosoma viridis O.<br />
Costa, 1844.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Madeira: SE coast near Agua de Pena,<br />
32°41'N, 16°46'W, 0-25 m, diving, 9 Mar 1976, Onversaagd<br />
Sta 27, 1$ (L).<br />
REMARKS.—Monod (1956) summarized earlier<br />
records for this species and provided a good illustration<br />
of a large male from Dakar. He reported<br />
FIGURE 63.—Acanthonyx lunulatus (Risso), female, cl 6.8 mm, Tunis: a, carapace; b, front, dorsal<br />
view; c, fifth pereiopod; d, distal segments of fifth pereiopod, enlarged.
NUMBER 306 261<br />
numerous specimens from localities in Mauritania,<br />
Senegal, Ghana, and Cameroon, and noted<br />
that its range in the Atlantic extended from<br />
Portugal to the equatorial region.<br />
We have examined material of A. lunulatus from<br />
the Gulf of Tunis, Messina and Trapani, Sicily,<br />
Naples, Nice, and Cadaques, Spain (all W) and<br />
have provided here some sketches of a young<br />
female from the Gulf of Tunis (Figures 60c, 63)<br />
to help distinguish this species from the three<br />
others occurring off <strong>West</strong> Africa.<br />
Our discovery of two previously undescribed<br />
species of Acanthonyx in <strong>West</strong> <strong>African</strong> waters and<br />
our recognition of A. brevifrons from the Cape<br />
Verde Islands and other localities indicate that<br />
earlier records of A. lunulatus may have been based<br />
on one or more of the four species now known to<br />
occur off <strong>West</strong> Africa. The specimens reported in<br />
Monod, the literature records cited by him, and<br />
the records given below all need to be verified.<br />
DISTRIBUTION.—Mediterranean Sea and adjacent<br />
Atlantic, from Portugal to South-<strong>West</strong> Africa,<br />
generally in shallow water, littoral and subtidal.<br />
Records since 1956 include the following.<br />
Morocco: Rabat, Temara, Skhirat, Oued Cherrat, David,<br />
Tillet [?], Sidi Moussa, Agadir, and Foum Assaka (all Forest<br />
and Games, 1960).<br />
Cape Verde Islands: Matiota, Sao Vicente, beach and<br />
shore, and Tarrafal do Monte Trigo, Santo Antao (Guinot<br />
and Ribeiro, 1962; Ribeiro, 1964).<br />
Guinea: No specific locality (Uschakov, 1970).<br />
Ivory Coast: Lagoon of Abidjan, 05°16'12"N, 04°00'-<br />
20"W (Forest and Guinot, 1966).<br />
Ghana: Tenkpobo to Dixcove (Gauld, 1960).<br />
Sao Tome: Ilheu Macao (as Hot dos Cocos), 3-8 m, and<br />
Sant'Ana, shore (Forest and Guinot, 1966).<br />
Congo: Pointe Indienne (Rossignol, 1957). Baie de<br />
Loango and near Pointe-Noire (Rossignol, 1962).<br />
Angola: Baia do Lobito, shore; Praia das Conchas, Mocamedes,<br />
shore; and Baia das Vacas, Benguela, shore (all<br />
Guinot and Ribeiro, 1962).<br />
South-<strong>West</strong> Africa: Rocky Point, 18°59'S, 12°29'E (Kensley,<br />
1970; Penrith and Kensley, 1970b). Kunene River<br />
mouth, 17°15'S, 11°45'E, and Mowe Point, 19°23'S, 12°-<br />
42'E (Kensley, 1970).<br />
Mowe Point is the southernmost record for<br />
Acanthonyx on the <strong>West</strong> <strong>African</strong> coast.<br />
* Acanthonyx minor, new species<br />
FIGURES 60rf, 64<br />
?Acanthonyx brevifrons.—Balss, 1914:101 [not A. brevifrons A.<br />
Milne Edwards, 1869].<br />
MATERIAL EXAMINED.—Pillsbury Material: Annobon:<br />
Sta 271, shore, 6c5 (includes holotype), 4$ (1 ov) (L, W). Sta<br />
273, shore, 1
262<br />
a<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 64.—Acanthonyx minor, new species, Pillsbury Sta 271. Female paratype, cl 3.8 mm: a,<br />
carapace; b, front, ventral view; c, cheliped; d, second pereiopod; e, fifth pereiopod. Male<br />
paratype, cl 2.6 mm: /, carapace; g, abdomen; h, first pleopod. Male paratype, cl 1.7 mm: i,<br />
carapace.<br />
Fingers almost as long as palm (measured along<br />
upper surface). Cutting edges dentate, with only<br />
slight gape proximally. Palm smooth. Carpus<br />
globular, smooth, bearing no carinae or tubercles,<br />
shorter than palm, much shorter than merus.<br />
Latter smooth except for 2 blunt tubercles on<br />
basal half.<br />
Walking legs similar to those of other Acanthonyx<br />
species, but dactylus appearing more slender,<br />
with tip less strongly curved, terminating in slender<br />
apex, posterior margin with 7 or 8 small but<br />
distinct teeth. Second pereiopod (first walking<br />
leg) dactylus about as long as propodus (measured<br />
dorsally). Latter 4 times as long as high,<br />
h
NUMBER 306 263<br />
outer surface ending in disc-like projection covering<br />
base of dactylus. Posterior margin of propodus<br />
with low tubercle or prominence near midlength<br />
ornamented with strong, stiff hairs. Carpus<br />
as long as propodus, much shorter and narrower<br />
than merus. First walking leg longer than remainder,<br />
latter of similar shape but decreasing in size<br />
posteriorly. Fifth pereiopod (Figure 64*) shortest.<br />
Dactylus slightly longer than propodus, more<br />
robust than that of second leg. Propodus about<br />
2.5 times as long as wide. Tubercle on posterior<br />
margin low but distinct because of tuft of hairs.<br />
Carpus distinctly shorter than propodus, merus<br />
about as long as propodus.<br />
Male abdomen (Figure 64g) narrow. Telson<br />
tongue-shaped. Sixth somite broad proximally,<br />
distinctly narrowing at telson. Abdomen of largest<br />
females very broad, reaching to bases of legs;<br />
sixth somite widest.<br />
First male pleopod as shown in Figure 64h. It<br />
does not differ significantly from that of A. sanctaehelenae<br />
Chace from Saint Helena (Chace, 1966,<br />
fig. 12f).<br />
MEASUREMENTS.—Carapace lengths of males<br />
1.7 to 3.4 mm, of non-ovigerous females 1.8 to 3.8<br />
mm, of the ovigerous female 3.7 mm.<br />
REMARKS.—Acanthonyx minor, like A. depressifrons,<br />
is a very small species, as the name implies. It can<br />
be distinguished from all of the Atlantic species<br />
now known by the distinct projection on the<br />
orbital margin; that margin is evenly concave in<br />
the other species.<br />
It is likely that the two specimens reported by<br />
Balss (1914:101) from Annobon under the name<br />
A. brevifrons belong to the present species; Balss<br />
indicated that they were juveniles. Later, Balss<br />
(1922:72) identified this material with A. lunulatus,<br />
synonymizing A. brevifrons with that species.<br />
TYPE-LOCALITY.—Annobon Island, 01°25'S,<br />
05°38'E, equatorial Guinea.<br />
DISPOSITION OF TYPES.—The holotype (Crust.<br />
D. 31762), a male from Pillsbury Sta 271 with a<br />
carapace length of 3.4 mm, and seven paratypes<br />
are in the Rijksmuseum van Natuurlijke Historie,<br />
Leiden; two male and one female paratypes are<br />
in the collection of the <strong>Smithsonian</strong> <strong>Institution</strong>.<br />
ETYMOLOGY—The specific epithet is from the<br />
Latin and refers to the diminutive size of this<br />
species.<br />
BIOLOGY.—The present material all originates<br />
from Annobon. It was collected at two fish poison<br />
stations on the shore of Annobon. The first (Sta<br />
271) was situated on the NE shore between Punta<br />
Yoyo and Punta Pedrinha, the coast here was<br />
rocky with rubble and a small sandy beach;<br />
collecting was done in the surge where the water<br />
was rather turbid. The second station (Sta 273)<br />
was on the north shore of Annobon near Islote<br />
Piramide on a rocky and sandy shore. Because of<br />
the high surf, collecting was difficult, but most of<br />
the fishes and invertebrates killed at this station<br />
were gathered at the outer edge of the surf zone<br />
in about 5 to 8 feet (1.5 to 2.4 m) of water over<br />
rocks.<br />
DISTRIBUTION.—Known only from the typelocality,<br />
Annobon Island, equatorial Guinea.<br />
Subfamily INACHINAE MacLeay, 1838<br />
REMARKS.—The extensive Pillsbury collections<br />
of inachine crabs has necessitated a much more<br />
detailed coverage of this subfamily than we had<br />
originally expected. Eight genera of this subfamily<br />
occur off <strong>West</strong> Africa. Two of these are characterized<br />
and named herein. Representatives of<br />
six genera are included in the Pillsbury material;<br />
Dorhynchus and Ergasticus, each represented in the<br />
eastern Atlantic by one species, were not collected.<br />
37 species of inachine crabs are now known<br />
to occur in the eastern Atlantic, a surprising<br />
increase in recognized species since 1956 when<br />
Monod recognized 22 species, 19 from tropical<br />
<strong>West</strong> <strong>African</strong> waters; we recognize 29 tropical<br />
species below.<br />
Because the Pillsbury collections included what<br />
appear to be representatives of two new genera,<br />
we have examined representatives of Achaeopsis,<br />
Dorhynchus, Ergasticus, Physachaeus, and several related<br />
genera in order to better understand generic<br />
characters and relationships. As a result, we believe<br />
that Dorhynchus Thomson, 1873, is distinct<br />
from Achaeopsis Stimpson, 1858. We have in-
264<br />
eluded a diagnosis of Dorhynchus (p. 279), under<br />
which our conclusions are discussed.<br />
We also believe that three interrelated "major"<br />
morphological features of inachine crabs are subject<br />
to misinterpretation: the supraorbital spine,<br />
the postorbital spine(s), and whether or not the<br />
eyes are retractile.<br />
The supraorbital spine of Dorhynchus (Barnard,<br />
1950, fig. 4d) or of Macropodia gilsoni probably<br />
should be interpreted as an outgrowth of the<br />
supraorbital eave rather than as being homologous<br />
with the intercalated spine or lobe in the<br />
Majinae. Balss (1929) stressed the importance of<br />
the intercalated spine in majid classification but<br />
his conclusions do not appear to have gained<br />
general acceptance. Garth (1958:37), however,<br />
includes in his key to American genera of Inachinae<br />
the genus Achaeopsis, which can be distinguished<br />
by the presence of a spine intercalated<br />
between pre- and postorbital spines. Griffin<br />
(1966b: 35) suggested that "if the supraorbital<br />
spine found in Achaeopsis species is regarded as<br />
homologous with the intercalated spine of other<br />
majids, Achaeopsis becomes widely separated from<br />
these genera ...." We agree with Griffin, for<br />
Achaeopsis and the closely related Dorhynchus<br />
(which Griffin considered a synonym of Achaeopsis),<br />
are otherwise similar to Macropodia, differing<br />
primarily in having the rostral spines divergent<br />
rather than appressed. Further, the <strong>West</strong> <strong>African</strong><br />
Macropodia gilsoni and M. intermedia, which have a<br />
supraorbital spine but otherwise closely resemble<br />
the other species of the genus, would have to be<br />
separated into a distinct genus, a taxon which, to<br />
us, would be unnecessary and unnatural. We<br />
suspect that the supraorbital spine, like that of<br />
Dorhynchus and Macropodia gilsoni, which clearly<br />
arises from the supraorbital eave, should not be<br />
considered homologous to the intercalated spine.<br />
In the case of M. gilsoni, it is one of several specific<br />
characters that can be used to distinguish that<br />
species.<br />
The question of homologies of the postorbital<br />
or postocular spine in the subfamily Inachinae is<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
perhaps more troublesome. The term has been<br />
used for a spine arising from the posterior margin<br />
of the orbit (Rathbun, 1925, fig. 1, for example),<br />
as well as for any spine that may be situated on<br />
the lateral margin of the carapace between the<br />
orbital margin and the hepatic lobe, i.e., posterior<br />
to the orbit (Griffin, 1966a, fig. 1, upper left<br />
figure). If this latter spine, which is situated between<br />
the orbit and the hepatic lobe, is considered<br />
to be a displaced, "true" postorbital spine like<br />
that of Inachus, then, again, Macropodia gilsoni<br />
would have to be generically distinct from M.<br />
rostrata, which lacks it. We believe that this latter<br />
spine should not be considered to be a true<br />
postorbital spine; we propose to call this the<br />
nuchal spine to differentiate it from the postorbital.<br />
Among the eastern Atlantic genera of Inachinae,<br />
a postorbital spine is found only in Inachus<br />
and Ergasticus; in Dorhynchus, Achaeopsis, Stenorhynchus,<br />
and in some species of Macropodia, there is a<br />
nuchal spine but no postorbital spine. Indeed, the<br />
eastern Atlantic species of Macropodia show a complete<br />
range of development of the nuchal spine.<br />
Finally, the question of whether the eye is<br />
retractile or not may be one of semantics but it is<br />
a feature that can be difficult to interpret. Among<br />
the eastern Atlantic genera of Inachinae, all of<br />
which have movable eyes, the eye in some appears<br />
to be retractile, that is, it can be moved into a<br />
position adjacent to the postocular spine. This<br />
occurs only in two genera, Inachus and Ergasticus.<br />
In Dorhynchus, in which the eye is said to be<br />
retractile to the sides of the carapace (Rathbun,<br />
1925:27, definition of Achaeopsis) (and which also<br />
has been described as having a postocular spine<br />
but which we interpret as a nuchal spine), the eye<br />
in material available to us is no more retractile<br />
than in our material of Macropodia. In Macropodia<br />
the eye is described as not retractile (Christiansen,<br />
1969:110), that is, the eye in representatives of<br />
the two genera folds backward about to the same<br />
level. The eye can approach the carapace in<br />
Dorhynchus, but, we suspect, only because the carapace<br />
is broader in that genus and the angle
NUMBER 306 265<br />
formed by a line extending from the orbit to the<br />
hepatic lobe is quite different in Dorhynchus than<br />
it is in Macropodia. We would describe the eye as<br />
retractile only when it can be folded back into a<br />
postocular process like that of Inachus.<br />
Finally, although Ergasticus was assigned to the<br />
subfamily Pisinae by Bouvier (1940) and by Zariquiey<br />
Alvarez (1968), we can see no reason to<br />
Key to Eastern Atlantic Genera of Inachinae<br />
exclude it from the Inachinae, where it was placed<br />
by Balss (1957) and Monod (1956). The basal<br />
antennal article is extremely slender and the orbit<br />
is incomplete, with a supraorbital eave divided<br />
into spines, a large, postorbital spine, but no<br />
ventral orbital margin. We have not examined<br />
the male pleopod, which does not appear to have<br />
been illustrated.<br />
1. Rostrum undivided, elongate (as long as or longer than postrostral carapace),<br />
with spinules laterally. Basal antennal article convex ventrally,<br />
not longitudinally sulcate. [Abdomen of 6 somites in male, 5 in female]<br />
Stenorhynchus<br />
Rostrum distinctly bilobed or very short, anteriorly truncate. Basal antennal<br />
article flattened or longitudinally channelled ventrally 2<br />
2. Orbit with distinct spine or cupped process on posterior margin against<br />
which eye can be folded 3<br />
Orbit lacking distinct spine or process on posterior margin. [Carapace,<br />
between orbit and hepatic lobe, smooth, tuberculate, or with erect spine<br />
(nuchal). Abdomen of 6 somites in both sexes] 4<br />
3. Supraorbital spines present. Postorbital spine slender, not a cupped process.<br />
Rostrum composed of 2 long, slender, divergent spines. First walking leg<br />
not markedly longer than second. Abdomen of 7 somites in male, 6 in<br />
female Ergasticus<br />
Supraorbital margin unarmed dorsally. Postorbital spine a broad, cupped<br />
process. Rostrum short, of 2 blunt lobes or broad, triangular spines. First<br />
walking leg much longer than second, noticeably enlarged. Abdomen of<br />
6 somites in both sexes Inachus<br />
4. Rostral spines slender, divergent, upturned, extending anteriorly slightly<br />
beyond end of antennal peduncle. [Distinct supraorbital spine present.<br />
Antennular fossae grading into rostrum, lacking anterior rim. Nuchal<br />
spine present. Merus of pereiopods with distal spine(s). Dactyli of<br />
posterior 2 pereiopods not falciform] Dorhynchus<br />
Rostral spines usually short, separate, triangular or with spiniform apices,<br />
or composed of 2 appressed spines of varying length 5<br />
5. Front truncate, simple or obscurely bilobed, overreached slightly by interantennular<br />
septum. Carapace lacking spines dorsally. [Antennular fossae<br />
with anterior rim. Interantennular septum not produced ventrally into<br />
beak. None of dactyli falciform] Capartiella, new genus<br />
Front produced into 2 spines or spined lobes. Carapace with spines or<br />
tubercles dorsally. [Interantennular spine may be visible at base of rostral<br />
teeth] 6
266<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
6. Interantennular septum distinct but not strongly produced ventrally,<br />
scarcely if at all visible in lateral view below level of basal antennal<br />
segment. [Rostrum bilobed. Antennular fossae lacking anterior rim,<br />
grading into rostrum. Chelae of male inflated. Dactyli of posterior 2 or<br />
3 pereiopods falciform] Achaeus<br />
Interantennular septum produced ventrally into distinct, prominent beak,<br />
visible in lateral view, extending well below level of basal antennal<br />
segment<br />
Rostrum produced into 2 distinctly separate spines, spinulous laterally,<br />
truncated and spinulous apically. Chelae compressed, palm cristate<br />
above and below in both sexes. Orbital margin spinulous above. [Basal<br />
article of antenna lined with spines on inner and outer margins. Dactyli<br />
of posterior 2 pereiopods falciform] Calypsachaeus, new genus<br />
Rostrum produced into 2 smooth, unarmed spines, contiguous throughout<br />
their length. Chelae inflated, especially in adult males. Orbital margins<br />
unarmed above or with 1 or 2 dorsal spines, not lined with spinules.<br />
[Basal article of antenna, if armed, with 1 line or irregular row of<br />
tubercles or spines] Macropodia<br />
Genus Achaeus Leach, 1817<br />
Achaeus Leach, 1817, in 1815-1875, legend of pi. 22C [typespecies:<br />
Achaeuscranchii'Leach, 1817,bymonotypy; gender:<br />
masculine; name 1605 on Official List].<br />
DIAGNOSIS.—Carapace pyriform to elongate triangular,<br />
length usually greater than width, somewhat<br />
narrowed behind orbit as a neck, smooth or<br />
ornamented with tubercles, spinules, or, rarely,<br />
with long dorsal spine on cardiac or gastric region<br />
or both (as in A. monodi sensu stricto). Rostrum<br />
very short, of 2 acute or rounded lobes or of 2<br />
spinules. Orbit dorsally with only narrow supraorbital<br />
eave, laterally spinulated or smooth;<br />
postorbital spine absent, nuchal lobe or spine, if<br />
present, not prominent. Eyestalks long, nonretractile,<br />
cornea obliquely subterminal, slightly<br />
ventral, large, ovoid. Interantennular septum not<br />
produced into ventrally directed beak. Basal antennal<br />
article extremely slender, smooth, channelled<br />
longitudinally, weakly tuberculate, spinulous,<br />
or armed with lateral spines, spines not<br />
arranged in 2 rows flanking longitudinal channel.<br />
Merus of third maxilliped ovate, not notched<br />
distally, narrower than ischium, palp articulating<br />
at summit. Chelipeds usually spinous, greatly<br />
enlarged with inflated palm in males. Walking<br />
legs long and slender, dactyli of posterior 2 or 3<br />
pairs usually strongly falcate and ventrally spinulated.<br />
Abdomen of 6 somites in both sexes,<br />
male abdomen widest at middle of third somite.<br />
Male first pleopod bluntly pointed, aperture located<br />
subterminally in groove (modified from<br />
Griffin, 1966b).<br />
REMARKS.—Achaeus now includes more than 30<br />
species (Serene, 1968, listed 25 Indo-<strong>West</strong> Pacific<br />
species) occurring in the eastern Atlantic and in<br />
the Indo-<strong>West</strong> Pacific, from South Africa to Japan<br />
(Griffin, 1966b, 1968a; Griffin and Yaldwyn,<br />
1965).<br />
The Pillsbury collections, comprising four species,<br />
bring to six the number of nominal species<br />
known from tropical <strong>West</strong> Africa, between Senegal<br />
and Angola. Two of the species taken by the<br />
Pillsbury are described here as new, and the other<br />
two are redescribed. One species, originally assigned<br />
to Achaeus and described from the Calypso<br />
collections in 1966 (Forest and Guinot, 1966), is<br />
transferred to another genus, Calypsachaeus.<br />
The seventh eastern Atlantic species, A. gracilis<br />
(O. Costa, 1839) (= A. gordonae Forest and Zariquiey<br />
Alvarez, 1955), occurs in the Mediterranean<br />
and so far has not been recorded from<br />
Atlantic localities. Monod (1956:542, figs. 767-<br />
770) supplemented the original description.
NUMBER 306 267<br />
It seems very likely that several additional<br />
species of Achaeus from <strong>West</strong> Africa remain to be<br />
recognized and described. Monod (1956:539, figs.<br />
747-766) almost certainly included two species in<br />
his account of A. cranchii: A. cranchii sensu stricto<br />
from European waters and an undescribed species<br />
from Senegal. The male pleopods of the two<br />
species, as figured by Monod (1956, figs. 760-762<br />
from Senegal and figs. 763-766 from Spain,<br />
France, and the Canary Islands) are quite different.<br />
Monod's account of Achaeus monodi (1956:548,<br />
figs. 782-810), which he, as well as Forest and<br />
Guinot (1966:110), considered to be an extremely<br />
variable species, may be represented in Monod's<br />
account by three distinct species: (1) A. monodi<br />
sensu stricto (fig. 782, reproduced from Capart,<br />
1951, pi. 1: fig. 11); a broad, pyriform species<br />
from the Congo and Angola with spiniform rostral<br />
spines and slender, erect spines dorsally on<br />
the gastric and cardiac regions of the carapace;<br />
(2) a slender, elongate-triangular species from<br />
Senegal with broad, triangular rostral spines and<br />
no dorsal spines (figs. 783-785); and (3) a second<br />
species from Senegal with appressed rostral spines<br />
(figs. 786-789), similar to those of the South<br />
<strong>African</strong> A. spinosissimus Griffin, 1968(b). None of<br />
these species was represented in the Pillsbury collections.<br />
Our generic diagnosis, based on Griffin<br />
(1966b), has been modified to include A. monodi,<br />
sensu stricto, with its erect dorsal spines. That<br />
species should be restudied, however, because it<br />
is different enough from the other species of<br />
Achaeus for Capart (1951) to have assigned it to<br />
Podochela.<br />
The relative proportions of the carapace, even<br />
though they may vary within a species with size<br />
and sex, provide a helpful recognition character<br />
for species of Achaeus. They allow the immediate<br />
distinction between slender species, like A. turbator,<br />
new species (in which the carapace width varies<br />
from 0.64 to 0.72 times the carapace length), from<br />
the broader species, like A. foresti (in which the<br />
carapace width varies from 0.86 to 0.90 times the<br />
length). Key characters employed for species not<br />
in the collections have been extracted from the<br />
literature: Forest and Zariquiey Alvarez, 1955,<br />
for A. cranchii and A. gracilis (as A. gordonae); Forest<br />
and Guinot, 1966, for A. trifalcatus; and Forest,<br />
1968, for A. foresti. A similar feature to illustrate<br />
the relative proportions of the carapace, the relation<br />
of the width at the hepatic lobes to the<br />
distance between the summit of the gastric prominence<br />
and the anterior margin of the rostral<br />
spines, also proves to be helpful. In A. turbator,<br />
new species, for example, the width of the carapace<br />
at the hepatic lobes is less than the distance<br />
from the gastric prominence to the anterior margin<br />
of the rostral spines, whereas in A. buderes, new<br />
species, the hepatic width is far greater. The<br />
relative length of the neck also provides a helpful<br />
recognition character: the neck is almost absent<br />
in A. buderes, new species, whereas it is quite<br />
distinct and elongate in A. turbator, new species.<br />
The shape of the rostral spines or projections also<br />
is important and appears to be characteristic. In<br />
A. trifalcatus the rostrum is made up of two<br />
rounded, unarmed lobes; in A. turbator, new species,<br />
it is made up of two rounded lobes, each of<br />
which has an anterior spinule. There is a definite<br />
shoulder on each side of the apical spinule; and<br />
in A. cranchii and A. gracilis the rostrum is made<br />
up of two slender, separate spines, tapering evenly<br />
from the base to the apex and lacking a distinct<br />
shoulder posteriorly. Note that in A. cursor (A.<br />
Milne-Edwards and Bouvier, 1900:161, pi. 21:<br />
figs. 15, 16, and pi. 22: figs. 1-3), considered a<br />
synonym of A. cranchii by Forest and Zariquiey<br />
Alvarez, 1955, there may be a distinct shoulder<br />
at the base of the rostrum. The proportions of the<br />
segments of the pereiopods also appears to be<br />
important. Forest (1968), for example, pointed<br />
out that in A. foresti the dactylus of the fourth<br />
pereiopod, measured across its arc, is equal to the<br />
length of the ischium and merus together; in A.<br />
monodi, according to the figure of the holotype<br />
(Capart, 1951, fig. 31), the dactylus, measured<br />
across its arc, is less than half as long as the merus<br />
alone. The size of the eye (compare Figures 65<br />
and 66A of the eye in A. foresti and A. buderes, new<br />
species) appears to be important. Finally, the<br />
relative height of the gastric and cardiac promi-
258 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
nences and whether or not they are produced into The key to eastern Atlantic Achaeus given below<br />
erect spines seems to be important; in none of the should be used with caution, for it does not take<br />
eastern Atlantic species are these spines so well into account the "variants" identified with A.<br />
developed as in A. monodi sensu stricto. cranchii and A. monodi by Monod (1956).<br />
Key to Eastern Atlantic Species of Achaeus<br />
1. Rostrum unarmed, scarcely bilobed. Posterior 3 pereiopods with falciform<br />
dactyli. [Carapace lacking erect dorsal spines. Width between hepatic<br />
lobes greater than distance between gastric prominence and anterior<br />
margin. Carapace width 0.70 to 0.83 times carapace length]<br />
A. trifalcatus<br />
Rostrum with apical spinules or spines. At most posterior 2 pereiopods with<br />
falciform dactyli 2<br />
2. Rostrum consisting of 2 sharp spines, tapering evenly from base to<br />
apex 3<br />
Rostrum consisting of 2 lobes, each with apical spinule (i.e., rostrum<br />
posteriorly with distinct shoulder) 4<br />
3. Fissure between rostral spines narrow. Orbits usually unarmed dorsally,<br />
occasionally with tubercle or spinules. Posteromedian margin of carapace<br />
unarmed. Cardiac protuberance rounded, lower than gastric protuberance.<br />
Width between hepatic lobes equal to distance between gastric<br />
prominence and anterior margin. [Carapace width 0.65 to 0.79 times<br />
carapace length in males, 0.75 to 0.86 times carapace length in females]<br />
A. gracilis<br />
Fissure between rostral spines wide, U- or V-shaped. Orbits armed dorsally.<br />
Posteromedian margin of carapace usually tuberculate or lightly spinulose.<br />
Cardiac protuberance conical, as high as gastric protuberance.<br />
Width between hepatic lobes greater than distance between gastric<br />
prominence and anterior margin. [Carapace width 0.73 to 0.87 times<br />
carapace length in males, 0.75 to 0.95 times carapace length in females]<br />
A. cranchii<br />
4. Carapace with erect spines on both cardiac and gastric regions. [Width<br />
between hepatic lobes greater than distance from gastric protuberance<br />
to anterior margin. Carapace width greater than 0.8 times carapace<br />
length. Only 1 pereiopod with falciform dactylus (?)] A. monodi<br />
Carapace without erect spines on either cardiac and gastric regions .... 5<br />
5. Supraorbital margin spinulate. Posterior margin of carapace with strong<br />
spines posterolaterally, smaller spinules and tubercles posteriorly. Eyes<br />
slender, diameter of stalk about one-fourth length. [Width between<br />
hepatic lobes greater than distance from gastric protuberance to anterior<br />
margin. Carapace width 0.86 to 0.90 times carapace length] . A. foresti<br />
Supraorbital margin unarmed. Posterior margin of carapace unarmed,<br />
with at most low, inconspicuous tubercles posterolaterally. Eyes stout,<br />
diameter about half length 6
NUMBER 306<br />
6. Carapace stout, pyriform, greatest width 0.67 to 0.81 (mean 0.75) times<br />
carapace length. Width at hepatic lobes greater than distance from<br />
gastric protuberance to anterior margin. Hepatic lobe with apical spine<br />
and several subapical spinules. Neck very short, stout<br />
A. buderes, new species<br />
Carapace slender, elongate triangular, greatest width 0.64 to 0.72 (mean<br />
0.68) times carapace length. Width at hepatic lobes subequal to or less<br />
than distance from gastric protuberance to anterior margin. Hepatic<br />
lobe angled apically, unarmed. Neck very long, tapering anteriorly<br />
A. turbator, new species<br />
*Achaeus buderes, new species<br />
FIGURE 65<br />
MATERIAL EXAMINED.—Pillsbury Material: Ghana:<br />
Sta 17, 48 m, fine sand and green mud, 1
270<br />
spines distally and 1 or 2 tubercles or spinules<br />
proximally. Antennal segments unarmed.<br />
Antennular fossae large, longitudinally suboval,<br />
margins of fossae smooth. Basal antennular<br />
segment with line of tubercles ventrally. Interantennular<br />
process slender, terminating posteriorly<br />
in rounded right angle. Anterior process of<br />
epistome very thin.<br />
Epistome (Figure 65c) broader than long,<br />
widening posteriorly, most of surface smooth,<br />
with sharp spinule anterolateral to opening of<br />
antennal gland. Pterygostomian region with<br />
sharp spine along lateral border.<br />
Third maxillipeds not meeting in midline,<br />
hairy, not heavily spined. Ischium with 2 parallel<br />
lines of sharp denticles, mesial margin tuberculate<br />
or with small spines, lacking large spines. Merus<br />
with line of sharp denticles on surface, several<br />
small spines on mesial margin. Palp with slender<br />
dactylus, shorter than stout carpus and propodus<br />
combined. Carpus and propodus each with distal<br />
spine mesially.<br />
Chelipeds slender in adult female, merus not<br />
extending beyond orbit. Ischium with line of<br />
spines ventrally. Merus with line of spines, some<br />
large, on ventrolateral border, 2 widely separated<br />
and 1 distal spine dorsally, and 1 distal spine<br />
mesially. Carpus with proximal outer spine, 2<br />
spines on oblique dorsal line, and 1 inner distal<br />
spine, flanked proximally by sharp tubercle.<br />
Chela shorter than remainder of cheliped, fingers<br />
longer than palm. Palm with row of spines dorsally<br />
and ventrally, inner and outer surfaces<br />
smooth. Fingers flattened, lacking prominent<br />
gape (in $), cutting edges crenulate, lacking enlarged<br />
tooth on movable finger. Chela largely<br />
naked, with few long and short hairs scattered<br />
over surface.<br />
Walking legs short, slender, merus and propodus<br />
of first walking leg each shorter than carapace,<br />
with curled hairs arising singly on dorsal<br />
surface of carpus and propodus, longer hairs scattered<br />
over surfaces of both segments. First walking<br />
leg longest, remainder decreasing in length posteriorly.<br />
Proximal segments of walking legs unarmed,<br />
merus lacking conspicuous distal dorsal<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
spine. Dactylus of first walking leg straight proximally,<br />
slightly curved distally, unarmed ventrally,<br />
more than half as long as propodus, latter<br />
slightly shorter than merus. Dactylus of second<br />
walking leg shorter than that of first, more evenly<br />
curved, with 1 subdistal tooth ventrally; dactylus,<br />
measured across arc, more than half as long as<br />
propodus, latter shorter than merus. Dactylus of<br />
third walking leg falciform, with 3 large distal<br />
and 3 smaller proximal ventral teeth, length,<br />
measured across arc, slightly more than half propodus<br />
length, latter 4/5 as long as merus. Dactylus<br />
of fourth walking leg (Figure 65d) falciform,<br />
with 5 or 6 large distal and several smaller proximal<br />
teeth ventrally, length, measured across arc,<br />
more than half that of propodus, latter shorter<br />
than merus. Sternum unarmed. Male too damaged<br />
to determine details of abdomen and gonopod.<br />
MEASUREMENTS.—Carapace length of male 4.7<br />
mm, of non-ovigerous females 3.4 to 4.0 mm, of<br />
ovigerous females 3.0 to 7.0 mm.<br />
REMARKS.—Achaeus buderes, with its stout, pyriform<br />
carapace with a very short neck, most closely<br />
resembles A. foresti. As in that species, there is a<br />
spinulose patch on the anterior margin of the eye,<br />
the hepatic lobes are armed, and there are spines<br />
and tubercles laterally on the carapace. This new<br />
species differs from A. foresti in having much<br />
stouter eyes, with the diameter of the stalk about<br />
half its length, having the supraorbital margin<br />
unarmed, having the posterior margin of the<br />
carapace unarmed, and in having a much shorter<br />
dactylus on the fourth pereiopods. In A. foresti<br />
that dactylus, measured across its arc, is longer<br />
than the ischium and merus combined, whereas<br />
in A. buderes it is slightly longer than half the<br />
length of the merus.<br />
TYPE-LOCALITY.—Off Ghana, 05°10'N, 00°-<br />
25'W to 05°08'N, OO°28'W, in 42 m (Pillsbury Sta<br />
23).<br />
DISPOSITION OF TYPES.—The holotype (Crust.<br />
D. 23903), an ovigerous female, is in the Rijksmuseum<br />
van Natuurlijke Historie, Leiden. One<br />
lot of paratypes is also deposited there and two<br />
lots of paratypes are in the National Museum of
NUMBER 306 271<br />
Natural History, <strong>Smithsonian</strong> <strong>Institution</strong>, Washington,<br />
D.C.<br />
ETYMOLOGY.—The specific epithet is derived<br />
from the Greek prefix bou-, large, and dere, neck.<br />
BIOLOGY.—This species was taken by the Pillsbury<br />
in depths between 35 and 51m, usually on<br />
soft bottom, fine sand and green mud, with bryozoans.<br />
Ovigerous females were taken in May.<br />
DISTRIBUTION.—Known only from type-locality,<br />
in 35 to 51 m.<br />
Achaeus cranchii Leach, 1817<br />
Achaeus cranchi.—Monod, 1956:539, 632, figs. 747-766 [Europe,<br />
Mediterranean, Canary Islands, Senegal, Sierra<br />
Leone].—Longhurst, 1958:89 [Sierra Leone].—Forest and<br />
Games, 1960:357 [Morocco].<br />
Achaeus cranchii.—Zariquiey Alvarez, 1968:474, fig. 160e,f<br />
[Spain; references].—Christiansen, 1969:109, fig. 45, map<br />
38 [North Atlantic].<br />
SYNONYM.—?Achaeus cursor A. Milne Edwards<br />
and Bouvier, 1898.<br />
DISTRIBUTION.—Eastern Atlantic from the British<br />
Isles southward to Sierra Leone, including the<br />
Azores and the Canary Islands; Mediterranean.<br />
Littoral to at least 68 m.<br />
* Achaeus foresti Monod, 1956<br />
FIGURE 66<br />
Achaeus foresti Monod, 1956:545, figs. 771-776.—Forest,<br />
1968:10%, figs. 1-6.<br />
MATERIAL EXAMINED.—PUlsbury Material: Liberia:<br />
Sta 68, 70 m, broken shell, 1$ (?).<br />
Ghana: Sta 17, 48 m, fine sand and green mud, 16* (?).<br />
DESCRIPTION.—Forest, 1968:1096.<br />
Male Pleopod: Forest, 1968, fig. 6 (Ghana).<br />
MEASUREMENTS.—Carapace length of male 2.8<br />
mm, of female 3.7 mm. The only other known<br />
specimen, the holotype, is an ovigerous female<br />
with a carapace length of 5 mm.<br />
REMARKS.—Forest (1968) redescribed this species<br />
from two specimens collected by the PUlsbury.<br />
His figures have been reproduced here (Figure<br />
66) for the sake of completeness and to facilitate<br />
the recognition of this species and the other species<br />
from the PUlsbury collections.<br />
The holotype, in the Museum national<br />
d'Histoire naturelle, Paris, is in poor condition.<br />
As Forest (1968:1098) pointed out, all that remains<br />
of the type is the carapace on which the<br />
apices of the rostrum have been broken. Thus,<br />
these rostral spinules are shown by Forest (1968,<br />
fig. 1) but the rostrum appears unarmed in the<br />
illustrations given by Monod (1956, figs. 771—<br />
773); this latter figure also does not show the<br />
marginal spinules on the basal portion of the<br />
rostrum. Monod (1956, fig. 776) illustrates one of<br />
the pereiopods of the type on which the merus is<br />
h<br />
FIGURE-66.—Achaeus foresti Monod. Female, cl 3.7 mm, PUlsbury<br />
Sta 68: a, carapace; b, eye; c, d, right cheliped, viewed<br />
from different angles. Male, cl 2.8 mm, PUlsbury Sta 17: e,<br />
right cheliped; /, fifth pereiopod; g,h, gonopod in anterior<br />
and posterior views. (All from Forest, 1968, figs. 1-6.)
272<br />
considerably longer than the propodus and the<br />
dactylus is subequal to the propodus; he may<br />
have shown the second pereiopod, for in the other<br />
species of Achaeus reported here that is the only<br />
pereiopod with the dactylus unarmed. Forest<br />
(1968) described the second pereiopod as having<br />
the merus shorter than the propodus and the<br />
dactylus 0.7 times as long as the propodus; his<br />
account of the third pereiopod indicates that the<br />
merus is slightly longer than the propodus and,<br />
as on the second pereiopod, the dactylus is 0.7<br />
times as long as the propodus. Finally, the strong<br />
spines near the opening of the antennal gland<br />
mentioned by Forest in the Pillsbury specimens<br />
are not shown in Monod's original illustration of<br />
the species. We have not had an opportunity to<br />
reexamine the two Pillsbury specimens.<br />
BIOLOGY.—Almost nothing is known of the<br />
biology of this species, represented by three specimens,<br />
one taken in 40 m off Senegal, one in<br />
70 m on broken shell off Liberia, and one from<br />
48 m on fine sand and green mud off Ghana. The<br />
only ovigerous female so far known was collected<br />
in March.<br />
DISTRIBUTION.—Known from the following<br />
three localities off <strong>West</strong> Africa.<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Achaeus trifalcatus Forest and Guinot, 1966<br />
Achaeus sp.—Monod, 1956:547, figs. 777-781 [Annobon].<br />
Achaeus trifalcatus Forest and Guinot, 1966:110, figs. 15-17,<br />
[Principe, Sao Tome, Annobon].<br />
DISTRIBUTION.—Known only from the offshore<br />
islands of the Gulf of Guinea, Principe, Sao Tome,<br />
and Annobon, in depths from 0-6 to 37 m..<br />
* Achaeus turbator, new species<br />
FIGURE 67<br />
MATERIAL EXAMINED.—Pillsbury Material: Nigeria:<br />
Sta 248, 33 m, 96* (includes holotype), 2$ ov (L, W).<br />
DESCRIPTION.—Carapace (Figure 67a,c) elongate<br />
triangular, width 0.64-0.72 (mean 0.68)<br />
times length, strongly narrowed anteriorly,<br />
slightly constricted behind orbits, neck appearing<br />
elongate, branchial regions swollen, regions well<br />
defined. Surface appearing smooth, not markedly<br />
granular.<br />
Rostrum (Figure 67a,c) consisting of 2 very<br />
short, slender spinules with distinct posterior or<br />
basal shoulder, arising from truncate lobe or from<br />
2 lobes separated by very shallow median emar-<br />
Senegal: S of lie de la Madeleine, near Dakar, 40 m gination. Rostral spines directed anteriorly, not<br />
(Monod, 1956).<br />
upturned in lateral view (Figure 676).<br />
Liberia: 04°23'N, 08°05.5'W to 04°24'N, 08°07.5'W, Supraorbital eave (Figure 67a,b) smooth or<br />
70 m (Forest, 1968).<br />
lightly tuberculate, not armed with spinules.<br />
Ghana: 05°35'N, 00°10'E to 05°36'N, 00°11.5'E, 48 m<br />
Postorbital margin smooth, unarmed. Eyestalks<br />
(not 42 m)(Forest, 1968).<br />
stout, width about half length, expanded distally,<br />
anterior margin sinuous, lacking well-marked<br />
Achaeus monodi (Capart, 1951) projection. Narrow process on anterior margin of<br />
Podochela monodi Capart, 1951:95, fig. 31, pi. 1: figs. 8, 11, pi. stalk extending to rounded tubercle over cornea.<br />
2: figs. 17, 18, 20 [Cabinda, Zaire, Angola].<br />
Latter large, oval, obliquely terminal.<br />
Achaeus monodi—Monod, 1956:548, figs. 782-810 [Senegal, Hepatic region moderately expanded, pro-<br />
Guinea, Sierra Leone, Gabon].—Longhurst, 1958:89 duced into acute or subacute unarmed lobe, an-<br />
[Sierra Leone].—Rossignol, 1962:122 [Congo].—Forest terior margin almost perpendicular to body line,<br />
and Guinot, 1966:109 [Guinea, Ivory Coast, Nigeria, Principe,<br />
Sao Tome].—Le LoeufT and Intes, 1968, table 1 occasionally with lateral tubercle but lacking ap-<br />
[Ivory Coast].<br />
ical spine. Width at hepatic lobes subequal to or<br />
less than distance from gastric protuberance to<br />
DISTRIBUTION.—<strong>West</strong> Africa, from Senegal to anterior margin.<br />
Angola, including the offshore islands of the Gulf Dorsal surface of carapace with 2 prominences<br />
of Guinea, Principe, and Sao Tome; sublittoral, on midline, gastric prominence an inflated tuber-<br />
0-4 to 100 m.<br />
cle, smaller and lower than cardiac prominence,
NUMBER 306 273<br />
a ZL1<br />
FIGURE 67.—Achaeus turbator, new species. Ovigerous female paratype, cl 6.0 mm, Pillsbuty Sta<br />
248: a, carapace, dorsal view; b, carapace, lateral view. Male paratype, cl 6.6 mm, Pillsbury Sta<br />
248: c, carapace, dorsal view; d, epistome; e, chela; /, dactylus of fifth pereiopod; g, abdomen;<br />
h, gonopod.<br />
neither spined dorsally. Protogastric region with<br />
low, unarmed tubercle. Branchial region smooth,<br />
with low mesobranchial tubercle centered dorsally.<br />
Epibranchial swelling, if present, low, inconspicuous.<br />
Low swelling present on metabranchial<br />
region above posterior pereiopods. Posterior<br />
and posterolateral margins unarmed, tubercles<br />
present anteriorly on margin above chelipeds.<br />
Basal antennal article (Figure 67d) usually<br />
smooth, lightly tuberculate or with 1 or 2 low<br />
spinules in some specimens. Antennal segments<br />
unarmed.<br />
Antennular fossae large, longitudinally subovate,<br />
lateral margins of fossae irregularly crenulate<br />
along border with basal antennal segment. Basal<br />
antennular segment bearing irregular line of tubercles<br />
ventrally. Interantennular process slender,<br />
terminating posteriorly in thin, blunt, obtuse,<br />
triangular lobe. Anterior process of epistome very<br />
slender.<br />
Epistome (Figure 61d) appearing elongate, actually<br />
broader than long, widening posteriorly,<br />
most of surface smooth, with single sharp tubercle<br />
present anterolateral to opening of antennal
274<br />
gland. Pterygostomian region with sharp tubercle<br />
on lateral border.<br />
Third maxillipeds not meeting in midline,<br />
hairy, not heavily spined. Ischium with 2 parallel<br />
lines of small tubercles, mesial margin tuberculate,<br />
erect spines present on margin in some specimens.<br />
Merus largely smooth, with few surface<br />
tubercles, narrow, unarmed mesially or with tubercles<br />
or small spines. Palp with slender dactylus,<br />
shorter than stout carpus and propodus combined;<br />
carpus and propodus each with distal spine<br />
mesially.<br />
Chelipeds (Figure 67
NUMBER 306 275<br />
ETYMOLOGY.—The name is from the Latin,<br />
turbator, meaning disturber or trouble-maker.<br />
DISTRIBUTION.—Known only from the typelocality,<br />
off Nigeria.<br />
Genus Calypsachaeus, new genus<br />
TYPE-SPECIES.—Achaeus calypso Forest and<br />
Guinot, 1966.<br />
ETYMOLOGY.—The name is derived from the<br />
name of the vessel Calypso in combination with<br />
the generic name Achaeus, gender of the name is<br />
masculine.<br />
DIAGNOSIS.—Carapace subtriangular, longer<br />
than broad, narrowed behind orbit, largely<br />
smooth dorsally, margins spinulous. Rostrum of<br />
2 elongate, anteriorly truncated lobes, apex and<br />
margins spinulous. Orbit with only supraorbital<br />
eave above, lined with long spinules. Postorbital<br />
spine absent. Eyestalks long, nonretractile, cornea<br />
obliquely subterminal, slightly ventral, large,<br />
ovoid. Interantennular septum produced ventrally<br />
into cupped process with distal sharp spine,<br />
visible in dorsal view. Basal antennal article<br />
slender, channelled longitudinally, depression<br />
flanked on each side by line of tubercles, with<br />
strong distolateral spine. Merus of third maxilliped<br />
ovate, not notched distally, narrower than<br />
ischium, palp articulating at summit. Chelipeds<br />
spinous, slightly enlarged in adult male, palm<br />
strongly compressed, cristate dorsally and ventrally.<br />
Walking legs long and slender, dactyli of<br />
posterior 2 falciform, ventrally spinulated. Abdomen<br />
of 6 free somites in both sexes, male<br />
abdomen widest at third somite. Male first pleopod<br />
bluntly pointed, aperture terminal in groove.<br />
REMARKS.—Calypsachaeus superficially resembles<br />
Achaeus, differing in the form of the rostrum,<br />
which is much better developed, in the strong<br />
development of the interantennular septum, in<br />
the ornamentation of the basal antennal segment,<br />
and in the form of the chelae, on which the palms<br />
are strongly compressed and cristate dorsally and<br />
ventrally. The male pleopod of Calypsachaeus (Forest<br />
and Guinot, 1966, fig. \9c-d) is similar to that<br />
of Achaeus in overall shape, but opens terminally<br />
rather than subterminally. Achaeus calypso is the<br />
only species we assign to this genus.<br />
* Calypsachaeus calypso (Forest and Guinot,<br />
1966), new combination<br />
FIGURE 68<br />
Achaeus calypso Forest and Guinot, 1966:113, figs. 18, 19.<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 68, 70 m, broken shell, 2
276<br />
orbital eave, completely lined with denticles or<br />
strong spinules. Distinct postorbital spine absent.<br />
Eyestalks slender, elongate, anterior or anteroventral<br />
margin with prominent, thin, blade-like, obtuse<br />
projection, as wide as or wider than stalk,<br />
with anterior spinules. Narrow process from anterior<br />
margin extending to rounded, setiferous<br />
tubercle over cornea. Latter broad, oval,<br />
obliquely terminal.<br />
Hepatic region well defined, strongly projecting<br />
laterally, smooth or with apical spine or with<br />
apical row of spinules flanked anteriorly by spinular<br />
crest on hepatic lobe. Carapace width at<br />
hepatic lobes subequal to or greater than distance<br />
from gastric prominence to anterior margin.<br />
Dorsal surface of carapace with 2 prominences<br />
in midline, gastric slender, usually an erect spine,<br />
often obsolete or broken, much the higher, cardiac<br />
prominence a low, obtusely rounded, conical lobe.<br />
Branchial regions smooth, unarmed, subregions<br />
not well marked. Lateral and posterior margins<br />
of carapace completely lined with tubercles or<br />
erect spinules.<br />
Basal antennal article longitudinally sulcate,<br />
with inner and outer lines of spinules, fewer in<br />
smaller specimens, outer line terminating in enlarged<br />
distolateral tooth. Free segments of antennal<br />
peduncle variably tuberculate or spinulose.<br />
Antennular fossae elongate-triangular, margins<br />
unarmed, lacking anterior rim, each fossa grading<br />
anteriorly into rostral spine. Basal antennular<br />
segment with patch or line of spiniform tubercles.<br />
Interantennular process strongly produced (Figure<br />
68/>), projecting well beyond level of basal<br />
antennal segment, expanded laterally to form<br />
broad, anteriorly cupped lobe, curved anteriorly,<br />
apically acute or spiniform. Inner, dorsal margin<br />
of cupped lobe grading evenly into inner surfaces<br />
of rostral horns. Anterior process of epistome thin,<br />
rounded, blade-like, projecting ventrally.<br />
Epistome subrectangular, broader than long,<br />
most of surface smooth, with spinulose crest or<br />
patch of spinules lateral to opening of antennal<br />
gland. Pterygostomian region with stalked projection,<br />
apically spinulose, on lateral border.<br />
Third maxillipeds not meeting in midline,<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
hairy, heavily spined. Exopod lined with spinules.<br />
Ischium with 2 divergent rows of sharp denticles,<br />
mesial margin spinulose, tuberculate. Merus with<br />
line of sharp denticles on surface, spined mesially,<br />
with spinules laterally and distally. Palp with<br />
slender dactylus, shorter than stout carpus and<br />
propodus combined. Carpus and propodus each<br />
with distal spine mesially.<br />
Chelipeds slender in both sexes, longer and<br />
slightly inflated in males, merus extending beyond<br />
eye in males, not extending to eye in females<br />
and juveniles. Ischium very spinous, largest spine<br />
set distoventrally. Merus with line of erect spines<br />
on sharp ventral border, outer surface smooth,<br />
upper border cristate, smooth or with line of<br />
tubercles, inner surface with some lines of tubercles.<br />
Palm strongly compressed, cristiform dorsally<br />
and ventrally, upper margin with spaced<br />
spines and tubercles, lower margin smooth, inner<br />
surface of lower margin with line of spines and<br />
denticles, outer surface smooth, inner surface tuberculate.<br />
Fingers longer than palm, flattened,<br />
lacking prominent gape, cutting edges crenulate,<br />
lacking enlarged proximal tooth on movable finger.<br />
Chelae largely naked, with few hooked hairs<br />
on outer surface, upper and lower borders with<br />
few long, stiff hairs interspersed between spines.<br />
Walking legs long and slender (merus of second<br />
pereiopod longer than carapace, propodus subequal<br />
to or slightly longer than carapace), with<br />
curled hairs arising in groups on dorsal surface of<br />
distal 4 segments and longer hairs scattered over<br />
surface, especially on propodus and dactylus. Second<br />
pereiopod longest, remainder decreasing in<br />
length posteriorly. Merus of walking legs with<br />
distal arc of spinules or low tubercles dorsally.<br />
Dactylus of second pereiopod straight proximally,<br />
curved distally, unarmed ventrally, more than<br />
half as long as propodus, latter shorter than<br />
merus. Dactylus of third pereiopod as long as that<br />
of second, slightly more curved distally, unarmed<br />
ventrally, more than half as long as propodus,<br />
latter subequal to merus. Dactylus of fourth pereiopod<br />
falciform, ventral margin with numerous<br />
triangular teeth, some swollen, proximally tuberculate,<br />
length across arc subequal to that of
NUMBER 306 277<br />
propodus, latter shorter than merus. Dactylus of<br />
fifth leg falciform, armature similar to that of<br />
fourth, length across arc subequal to that of<br />
propodus, latter shorter than merus.<br />
Male abdomen narrowed distally, apex<br />
broadly rounded, with median tubercle on each<br />
somite, sixth somite with 2 subapical tubercles or<br />
spinules. Male pleopod as figured by Forest and<br />
Guinot (1966, fig. 19c,d).<br />
Sternum granular in both sexes, in male ornamented<br />
with anteriorly convex arc of strong<br />
spines and tubercles, with 1 enlarged spine near<br />
base of each cheliped.<br />
MEASUREMENTS.—Carapace lengths of males<br />
5.7 to 9.0 mm, of females 4.8 to 9.3 mm, of<br />
ovigerous females 8.6 to 9.3 mm. The male holotype<br />
has a carapace length of 10.0 mm (Forest<br />
and Guinot, 1966).<br />
REMARKS.—This striking species can readily be<br />
recognized by its relatively smooth carapace, eyes<br />
with their flattened anterior projection, broad<br />
rostral spines, well-developed interantennular<br />
spine, flattened chelae, and the spinous ventral<br />
surface.<br />
BIOLOGY.—This species occurs on the shelf in<br />
the Gulf of Guinea in depths between 30 and 70<br />
m. The Pillsbury specimens were taken in 48 m on<br />
fine sand and green mud, in 51 m on rough<br />
bottom, and in 70 m on broken shell. The holotype<br />
was taken by the Calypso in 32 m on mud<br />
with Area, and Forest and Guinot (1966) also<br />
recorded material taken on mud in 30, 50, and<br />
70 m. Ovigerous females were collected in May.<br />
DISTRIBUTION.—Gulf of Guinea, where it has<br />
been recorded from localities between Liberia<br />
and Cameroon in depths between 30 and 70 m.<br />
Records in the literature, all from Forest and<br />
Guinot (1966), include the following:<br />
Ivory Coast: No specific locality, 30, 50, and 70 m.<br />
Nigeria: Off the mouths of the Niger River, 04°03'N,<br />
06°12'E, 32 m.<br />
It has not been recorded previously from Liberia,<br />
Ghana, Dahomey, or Cameroon.<br />
Genus Capartiella, new genus<br />
TYPE-SPECIES.—Achaeus longipes Capart, 1951.<br />
ETYMOLOGY.—We take great pleasure in dedi-<br />
cating this genus to A. Capart. His pioneering<br />
work on <strong>West</strong> <strong>African</strong> crabs published in 1951<br />
added much to our knowledge of the brachyuran<br />
fauna there. Gender of the generic name is masculine.<br />
DIAGNOSIS.—Carapace pyriform, length<br />
slightly greater than width, scarcely narrowed<br />
behind the orbit as a neck, lacking long spines or<br />
sharp tubercles. Rostrum a single short rounded<br />
projection, depressed anteriorly, overreached by<br />
strong interantennular spine. Orbit consisting<br />
above of narrow supraorbital eave, postorbital<br />
spines absent. Eyestalks long, nonretractile, movable.<br />
Cornea terminal, slightly ventral. Interantennular<br />
septum produced into distinct spine.<br />
Basal article of antennule slender, longitudinally<br />
sulcate, ornamented with low, rounded tubercles<br />
and distal blunt angular projection. Merus of<br />
third maxilliped subovate, not notched distally,<br />
narrower than ischium, palp articulating anterointernally.<br />
Chelipeds short, slender in both sexes.<br />
Walking legs long, slender, dactyli elongate,<br />
arched, with triangular ventral spines. Abdomen<br />
of 6 somites in both sexes. Male first pleopod with<br />
blunt apex, aperture terminal.<br />
REMARKS.—In his original description, based<br />
on a single non-ovigerous female, Capart (1951:<br />
63) noted that "e'est avec grande hesitation que<br />
je place cette curieuse espece dans le genre<br />
Achaeus...." He further noted that C. longipes<br />
could be separated from other species of Achaeus<br />
by its simple rostrum and by the shape of the<br />
dactyls of the walking legs. Monod (1956), who<br />
had no additional material, transferred the species<br />
to Physachaeus Alcock, with reservations, noting<br />
the superficial resemblance of C. longipes to P.<br />
ctenurus Alcock. Subsequent authors recording the<br />
species have followed Monod in placing the species<br />
in Physachaeus.<br />
Forest and Guinot (1966:108, 109), in recording<br />
the first males to be collected, interpreted the<br />
rostrum as being formed of two very short lobes<br />
which were overreached ventrally by the interantennular<br />
septum, i.e., the rostrum comprised<br />
two fused lobes rather than a single one. They<br />
also pointed out differences in the eyes and the
278<br />
basal antennal segment of C. longipes and those of<br />
species of Physachaeus, based on accounts in the<br />
literature, and noted (p. 109) "il est possible que<br />
Ton soit amene par la suite a creer pour elle un<br />
nouveau genre."<br />
We have been able to compare our material<br />
with a male and female Physachaeus ctenurus (13°-<br />
17'15"N, 93°10'25"E, 185 fm (339 m), Investigator,<br />
USNM 42757) and from this comparison conclude<br />
that the resemblance between C. longipes<br />
and P. ctenurus is superficial. In Physachaeus the<br />
rostrum is clearly bifid, the eyes are short and<br />
fixed, with a terminal cornea, there are large<br />
gastric and cardiac spines on the carapace, the<br />
basal article of the antenna bears a long spine,<br />
and there are but 5 somites in the abdomen of<br />
the female. The condition of the eyes in C. longipes,<br />
long and movable, versus those of P. ctenurus,<br />
would be enough to exclude C. longipes from<br />
Physachaeus.<br />
Capartiella most closely resembles Achaeus<br />
Leach, Achaeopsis Stimpson, Dorhynchus Studer,<br />
and the American Podochela Stimpson (definition<br />
of the latter genus in Rathbun, 1925). It resembles<br />
Achaeus, Achaeopsis, and Dorhynchus and differs<br />
from Podochela in having the abdomens of each<br />
sex composed of 6 somites. In Podochela, which<br />
also has an unarmed carapace and usually has a<br />
simple but anteriorly produced rostrum, the abdomen<br />
is composed of 6 somites in the male, 5 in<br />
the female. The legs of Capartiella, like those of<br />
Achaeopsis, are very long, slender, setose, with the<br />
dactyl slender, not strongly recurved, but Capartiella<br />
lacks the spines on the carapace. Capartiella<br />
shares many features with Achaeus, but differs in<br />
having a single rostrum, slender chelipeds that<br />
are not inflated or enlarged in adult males, and<br />
slender dactyli on the walking legs. The male<br />
pleopod of Capartiella longipes (figured by Forest<br />
and Guinot, 1966, fig. 14), resembles that of some<br />
species of Achaeus figured by Griffin (1970, figs.<br />
13-15) in general configuration, and differs<br />
strongly from the pleopods of Podochela, which are<br />
apically acute with a subterminal opening<br />
flanked by a rounded flap, as shown by Garth<br />
(1958, pi. H).<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
* Capartiella longipes (Capart, 1951),<br />
new combination<br />
FIGURES 69, 70<br />
Achaeus ? longipes Capart, 1951:62, fig. 19, pi. 1: fig. 1, pl. 2:<br />
figs. 21,22.<br />
Physachaeus (?) longipes.—Monod, 1956:537, fig. 746.—Forest,<br />
1959:15.—Rossignol, 1962:122.—Forest and Guinot,<br />
1966:108, fig. 14.—Crosnier, 1967:340.<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 68, 70 m, broken shell, 29 (L).<br />
Ivory Coast: Sta 42,62-75 m, mud with brown, branched<br />
Foraminifera, 16\ 19 (L, W). Sta 46, 38-42 m, mud with<br />
dense Julltenella, 1$ ov (L). Sta 60, 79-82 m, coral or rock, 26<br />
(W).<br />
Ghana: Sta 16, 46 m, mud with Foraminifera, shells, 16*,<br />
29 ov (L). Sta 23, 42 m, foliate brown to orange bryozoans,<br />
39 ov (L, W). Sta 24, 35-37 m, dark red bryozoans, 19 (W).<br />
Sta 28, 49-53 m, 16\ 19 ov (L).<br />
Nigeria: Sta 239, 73 m, 19 (W). Sta 240, 37 m, 16* (L).<br />
Sta 241, 59-63 m, mud and shell, 19 (L). Sta 248, 33 m, ld\<br />
19 (W).<br />
Other Material: Congo: Off Pointe-Noire, 04°48'S, 11°-<br />
39'E, 54-56 m, sandy mud, 23 Sep 1965, Ombango, A. Crosnier,<br />
26, 49 (1 ov) (W).<br />
. Cabinda: SW of Landana, 50 m, 23 May 1959, Ombango,<br />
A. Crosnier, 19 ov (W).<br />
DESCRIPTION.—Capart, 1951:63.<br />
Male Pleopod: Forest and Guinot, 1966, fig. 14<br />
(Ghana).<br />
MEASUREMENTS.—Carapace lengths of males<br />
6.0 to 11.1 mm, of non-ovigerous females 4.0 to<br />
13.9 mm, of ovigerous females 6.0 to 13.0 mm.<br />
REMARKS.—We have provided here sketches of<br />
C. longipes, as well as Capart's original figures<br />
(Figures 69, 70) of it, apparently one of the more<br />
common species on the shallow part of the shelf<br />
in the Gulf of Guinea. All of our specimens were<br />
completely covered with a dense coat of algae<br />
and mud, so dense that their shape was scarcely<br />
discernible. This may have caused the species to<br />
be overlooked in earlier collections, for the shelf<br />
from Ghana to Nigeria has been well sampled in<br />
the last 25 years.<br />
BIOLOGY.—This species apparently lives on soft<br />
bottoms in depths of no more than 82 m. The<br />
Pillsbury specimens were taken in depths of 33 to<br />
82 m, usually on mud bottom with foraminifera
NUMBER 306 279<br />
FIGURE 69.—Capartiella longipes (Capart) (from Capart, 1951, fig. 19.)<br />
or Bryozoa; one sample was taken on broken shell<br />
and one on coral or rock (which terminated the<br />
haul; a rock outcrop may have been encountered<br />
on an otherwise soft bottom). Capart (1951) reported<br />
the species from a depth of 60 m in green<br />
mud. The specimens reported by Forest and<br />
Guinot (1966) came from mud, sand, and compact<br />
sand (sable construit) in 67-75 m and from<br />
mud in 50 m. Crosnier (1967) found the species<br />
on mud and sandy mud in depths between 35<br />
and 58 m.<br />
Ovigerous females have been collected in February,<br />
May, June, and September.<br />
DISTRIBUTION.—Tropical <strong>West</strong> Africa, where it<br />
has been taken at a few localities between Senegal<br />
and Angola; sublittoral, in depths between 33<br />
and 82 m. Records include the following:<br />
Senegal: 12°55.5'N, 17°33'W, 65-75 m (Forest, 1959;<br />
Forest and Guinot, 1966).<br />
Ivory Coast: No specific locality (Forest and Guinot,<br />
1966).<br />
Ghana: 04°40'N, 02°08'W to 04°39'N, 02°05'W, 50 m<br />
(Forest and Guinot, 1966).<br />
Dahomey: No specific locality, 35-38 m (Crosnier, 1967).<br />
Cabinda: W of Landana, 50 m (Rossignol, 1962).<br />
Angola: 10 mi NxW of Luanda, 08°37'S, 13°12'E, 60 m<br />
(Capart, 1951).<br />
Genus Dorhynchus Thomson, 1873<br />
Dorhynchus Thomson, 1873:175 [type-species: Dorhynchus thomsoni<br />
Thomson, 1873, by monotypy; gender: masculine;<br />
name 1619 on Official List].<br />
Lispognathus A. Milne Edwards, 1881, in 1873-1881:349<br />
[type-species: Lispognathus furcillatus A. Milne Edwards,<br />
1881, by monotypy; gender: masculine].<br />
DIAGNOSIS.—Carapace triangular, pyriform,<br />
longer than broad, expanded behind orbit, with<br />
dorsal spines, margins spinulous. Rostrum of 2
280<br />
a<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 70.—Capartiella longipes (Capart). Male, cl 10.0 mm, Pillsbury Sta 248: a, front, dorsal<br />
view; b, front, oblique anterior view; c, front, ventral view; d, abdomen. Female, cl 12.0 mm,<br />
Pillsbury Sta 248: e, third maxilliped. Male, cl 11.1 mm, Congo:/, propodus and dactylus of<br />
fifth pereiopod.<br />
slender, sharp spines usually divergent anteriorly.<br />
Orbits with supraorbital eave provided with supraorbital<br />
spine(s), preorbital spine absent.<br />
Postorbital spine absent, nuchal spine present<br />
between hepatic lobe and orbit. Eyestalks short,<br />
movable but nonretractile, cornea terminal, large,<br />
subglobular. Interantennular septum produced<br />
into sharp spine, usually visible in dorsal view.<br />
Basal antennal article slender, channelled longitudinally,<br />
channel flanked by inner and outer<br />
rows of spines, with enlarged terminal spine distolaterally.<br />
Merus of third maxilliped spinous,<br />
ovate or subrectangular, not notched distally,<br />
narrower than ischium, palp articulating at summit.<br />
Chelipeds spinous, slenderer in female, palm<br />
inflated. Walking legs long and slender, dactyli<br />
elongate, not falciform, posterior 2 with low teeth<br />
ventrally. Abdomen of 6 somites in both sexes.<br />
Male first pleopod bluntly pointed, with distinct<br />
subterminal lappet.<br />
REMARKS.—Although the Pillsbury collections<br />
included no representatives of this genus, our<br />
recognition herein of some new genera in this<br />
subfamily necessitated examination of representatives<br />
of all of the genera of Inachinae in the<br />
eastern Atlantic. In the case of Dorhynchus, considered<br />
by some (Rathbun, 1925; Barnard, 1950;<br />
Griffin, 1966b) but not others (Bouvier, 1940;<br />
Monod, 1956; Zariquiey Alvarez, 1968; and<br />
Christiansen, 1969) to be synonymous with<br />
Achaeopsis Stimpson (1858d:218; type-species: A.<br />
spinulosa Stimpson, 1858, by monotypy; gender:<br />
feminine; name 1604 on Official List), comparison<br />
of material of D. thomsoni (off Morocco, 1105 m,<br />
Talisman, USNM 22974, 3d, cl 7.3-7.5 mm, 3$, cl<br />
5.8-7.5 mm) with a specimen of A. spinulosa<br />
(South Africa, 36°40'S, 21°26'E, 200 m, USNM:<br />
Id, cl 12 mm) leads us to suggest that the two<br />
should be regarded as distinct but closely related<br />
genera.<br />
Dorhynchus differs from Achaeopsis in several features:<br />
the rostral spines are much longer; the<br />
interantennular spine is much larger, projects<br />
further ventrally, and its apex usually is visible<br />
between the rostral spines in dorsal view; the<br />
walking legs are longer and slenderer: in D. thorn-
NUMBER 306 281<br />
soni the merus of the second pereiopod is longer<br />
than the carapace whereas in A. spinulosa it is<br />
shorter than the carapace; and the dactyli of the<br />
pereiopods are slender and similar on all legs in<br />
D. thomsoni, whereas they are falciform in the last<br />
two legs of A. spinulosa. Also, as Barnard (1950:<br />
25) pointed out, A. spinulosa generally occurs in<br />
shallower water than does D. thomsoni.<br />
If this interpretation is correct and Achaeopsis<br />
and Dorhynchus are retained as separate genera,<br />
then two other species assigned to Achaeopsis sensu<br />
lato by Griffin (1966b), Achaeopsis rostrata Sakai,<br />
1932, from Japan and Stenorhynchus ramusculus<br />
Baker, 1906, from South Australia and New Zealand,<br />
both of which have slender legs, should be<br />
transferred to Dorhynchus, and Achaeopsis should<br />
be restricted to include only the type-species, A.<br />
spinulosa.<br />
Although Dorhynchus thomsoni is now considered<br />
to be widely distributed ["virtually cosmopolitan,"<br />
Griffin (1966b:35)], we agree with Bouvier<br />
(1940:351), Monod (1956:523), and Guinot<br />
(1967a:289, footnote), all of whom question<br />
whether the populations from different areas are<br />
conspecific. In the collection of the <strong>Smithsonian</strong><br />
<strong>Institution</strong>, for example, there is one lot from<br />
Chiloe Island, off Chile (USNM 156225), in<br />
which the rostrum is spinulose ventrally, rather<br />
than smooth, and in which there are three distal<br />
spines on the meri of the pereiopods, rather than<br />
one as shown for D. thomsoni by Christiansen<br />
(1969:106, fig. 44). Material of Dorhynchus thomsoni<br />
should be examined from throughout its range to<br />
determine the relationship of the various populations.<br />
Dorhynchus thomsoni Thomson, 1873<br />
Dorynchus thomsoni Thomson, 1873:174, 175, fig. 34.<br />
Dorynchus Thomsoni.—Filhol, 1885a: 56.<br />
Lispognathus Thompsoni.—Filhol, 1885a, fig. 1.<br />
Dorhynchus thomsoni.—Monod, 1956:522 [references].—Zariquiey<br />
Alvarez, 1968:467 [Spain; references].—Christiansen,<br />
1969:106, fig. 44, map 37 [North Atlantic].<br />
DISTRIBUTION.—Eastern Atlantic, W and S of<br />
Iceland and the Faroes southward to the Cape<br />
Verde Islands; deep water species, in depths between<br />
100 and 2100 m. The occurrence of this<br />
species outside of the eastern Atlantic requires<br />
verification.<br />
Genus Ergasticus Studer, 1883<br />
Ergasticus Studer, 1883:7 [type-species: Ergasticus clouei Studer,<br />
1883, by monotypy; gender: masculine; name 1620<br />
on Official List].<br />
Ergasticus clouei Studer, 1883<br />
Ergasticus Clouei Studer, 1882:335 [nomen nudum]; 1883:8.<br />
Ergasticus clouei.—Monod, 1956:523.—Peres, 1964:20.—Zariquiey<br />
Alvarez, 1968:463, fig. 155b [Spain; references].—<br />
Tiirkay, 1976a:25 [listed], 39, fig. 32 [Portugal, in part].<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Morocco: Off Cap de Mazagan, 33°39'N,<br />
08°46'W, 500 m, Agassiz trawl, 20 Mar 1976, Onversaagd Sta<br />
151, 1$ (L).<br />
Cape Verde Islands: 16°52'N, 27°30'W of Paris (25° 10'W<br />
of Greenwich), 400-580 m, sand and rock, 29 Jul 1883,<br />
Talisman Sta 111, 16", 1? ov (W).<br />
DESCRIPTION.—A. Milne Edwards and Bouvier,<br />
1900:140.<br />
Figure: A. Milne Edwards and Bouvier, 1900,<br />
pi. 21: figs. 1-7.<br />
MEASUREMENTS.—Carapace length of nonovigerous<br />
females 8-10 mm, of ovigerous female<br />
8 mm.<br />
BIOLOGY.—Ergasticus clouei is a deepwater species,<br />
occurring in depths between 70 m and<br />
1241 m; more than 90% of the depth records in<br />
the literature are from depths between 300 and<br />
750 m.<br />
Ovigerous females have been recorded in July<br />
and August (A. Milne Edwards and Bouvier,<br />
1899, 1900; Bouvier, 1922).<br />
DISTRIBUTION.—Eastern Atlantic from the Bay<br />
of Biscay to the Cape Verde Islands, including<br />
Madeira and the Azores; Mediterranean. Deepwater,<br />
in depths between 70 and 1241 m. Monod<br />
(1956) reported no specimens. Other <strong>West</strong> <strong>African</strong><br />
records include the following:<br />
Morocco: Bane de Spartel, 35°54'N, 06°14'W, 210 m
282<br />
(Peres, 1964). 33°12.6'N, 09°15.2'W, 500 m, and 33°10.5'N,<br />
19°17.5'W, 170-345 m (Tiirkay, 1976a).<br />
Cape Verde Islands: 15°40'N, 23°06'W, 38 fm (70 m)<br />
(Studer, 1882).<br />
Genus Inachus Weber, 1795<br />
Inachus Weber, 1795:93 [type-species: Cancer scorpio Fabricius,<br />
1779, a subjective junior synonym of Cancer dorsettensis<br />
Pennant, 1777, by subsequent designation by H. Milne<br />
Edwards, 1840, in 1836-1844, pi. 34: fig. 2; gender: masculine;<br />
name 1698 on Official List].<br />
Macropus Latreille, 1803a:27 [an invalid junior homonym of<br />
Macropus Shaw, 1790; type-species: Cancer phalangium Fabricius,<br />
1775, by monotypy; gender: masculine; name<br />
1777 on Official Index].<br />
Leptopodia Leach, 1814:431 [type-species: Cancer phalangium<br />
Fabricius, 1775, by monotypy; gender: feminine].<br />
Pseudocollodes Rathbun, 1911:247 [type-species: Pseudocollodes<br />
complectens Rathbun, 1911, by monotypy; gender: masculine].<br />
REMARKS.—Griffin (1974:18) correctly synonymized<br />
Pseudocollodes Rathbun, 1911 with Inachus<br />
Weber, 1795, but, as pointed out later (p.<br />
290), we suspect that Inachus complectens (Rathbun),<br />
known from the western Indian Ocean and<br />
South Africa (Barnard, 1950; Griffin, 1974) is not<br />
conspecific with /. dorsettensis (Pennant).<br />
As is the case with most of the <strong>West</strong> <strong>African</strong><br />
spider crabs, the species of Inachus appear to<br />
require much further study. Four species are represented<br />
in the Pillsbury collections and three of<br />
these are undescribed; two of the species previously<br />
were identified with European species.<br />
Monod (1956) recorded seven species from<br />
<strong>West</strong> Africa: Inachus angolensis Capart (the only<br />
endemic species to be listed), /. dorsettensis (Pennant),<br />
/. guentheri (Miers), /. phalangium (Fabricius),<br />
/. thoracicus (Roux), /. aguiarii De Brito<br />
Capello, and /. leptochirus Leach. Monod's accounts<br />
of /. dorsettensis and /. leptochirus, both<br />
relatively well-known European species, appear<br />
to be based on material of the two new species<br />
described herein. Monod's account of /. guentheri<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
(Miers), which unfortunately is not represented<br />
in the Pillsbury collections, does not agree with the<br />
account of that species in Barnard (1950:27): the<br />
rostral teeth and the spination of the carapace<br />
appear to be quite different in the Gulf of Guinea<br />
and South <strong>African</strong> populations now referred to<br />
this species. Our studies of the Pillsbury collections<br />
have suggested that relatively few <strong>West</strong> <strong>African</strong><br />
species identified by earlier workers with European<br />
or Mediterranean species actually are conspecific<br />
with those species. We suspect, therefore,<br />
that available <strong>West</strong> <strong>African</strong> material referred to<br />
/. phalangium, I. thoracicus, and /. aguiarii, should<br />
be carefully reexamined.<br />
The identity of the dwarf forms of Inachus<br />
dorsettensis reported by Doflein (1904) and Odhner<br />
(1923) from deep water off <strong>West</strong> Africa cannot<br />
be determined with certainty without examining<br />
their material; they could have been dealing with<br />
the new species /. grallator or /. nanus, both of<br />
which occur in relatively deep water and are<br />
smaller than European Atlantic specimens of /.<br />
dorsettensis sensu stricto. We have tentatively referred<br />
them to /. grallator.<br />
The identity of the juvenile female from Angola<br />
reported by Odhner (1923:19) as Inachus sp.<br />
also cannot be determined with certainty on the<br />
basis of his account. He may have been dealing<br />
with /. biceps, new species, or with /. guentheri sensu<br />
lato.<br />
The following key distinguishes the <strong>West</strong> <strong>African</strong><br />
species of Inachus. The key would be complete<br />
for the eastern Atlantic by adding the Mediterranean<br />
/. communissimus (Rizza), which would key<br />
out with /. nanus, new species, /. dorsettensis, which<br />
would key out with /. angolensis and /. grallator,<br />
new species, and /. leptochirus, which would key<br />
out with /. biceps, new species. We decided against<br />
adding these species to the key because of the<br />
uncertain status of the Mediterranean population<br />
of /. dorsettensis (see Inachus grallator, new species,<br />
p. 287).<br />
Key to Tropical <strong>West</strong> <strong>African</strong> Species of Inachus<br />
1. Gastric areas of carapace with transverse row of 4 tubercles anterior to<br />
gastric spine 2<br />
Gastric area of carapace with at most 2 tubercles anterior to gastric<br />
spine 4
NUMBER 306 283<br />
2. Interantennular spine short, not visible between rostral spines in dorsal<br />
view /. nanus, new species<br />
Interantennular spine large, well developed, visible between rostral spines<br />
in dorsal view [often overreaching rostral spines] 3<br />
3. Cardiac and branchial regions of carapace with erect spines<br />
/. grallator, new species<br />
Cardiac and branchial regions of carapace lacking erect spines<br />
/. angolensis<br />
4. Rostral spines appressed for most of their length [white sternal callosity<br />
absent in male] /. phalangium<br />
Rostral spines separate for all of their length 5<br />
5. Branchial regions of carapace each with erect dorsal spine 6<br />
Branchial regions of carapace each with obtuse swelling or rounded tubercle<br />
dorsally, lacking erect dorsal spine 7<br />
6. Male lacking white sternal callosity /. guentheri<br />
Male with prominent, simple sternal callosity /. biceps, new species<br />
7. Walking legs elongate, first (= second pereiopod) more than 4 times as<br />
long as carapace in males. Sternal callosity in males only .. /. thoracicus<br />
Walking legs short, first (= second pereiopod) less than 4 times as long as<br />
carapace in males. Sternal callosity present in both sexes .... /. aguiarii<br />
Inachus aguiarii De Brito Capello, 1876<br />
Inachus aguiari.—Monod, 1956:533, figs. 736-739 [Guinea].<br />
—Forest and Guinot, 1966:107 [Spanish Sahara].<br />
Inachus aguiarii.—Zariquiey Alvarez, 1968:473, fig. 158c,d<br />
[Spain; references].<br />
DISTRIBUTION.—Eastern Atlantic, from Portugal,<br />
Madeira, and Guinea; Mediterranean. Sublittoral,<br />
in depths between 20 and 200 m.<br />
* Inachus angolensis Capart, 1951<br />
Inachus angolensis Capart, 1951:72, fig. 22, pi. 1: fig. 7, pi. 2:<br />
fig. 10.—Monod, 1956:524, figs. 711-714.—Rossignol,<br />
1957:77, 116 [key], pi. 1: fig. 1.—Gauld, 1960:72.— Rossignol,<br />
1962:122.—Crosnier, 1964:34.—Forest, 1965b:394<br />
[discussion].—Forest and Guinot, 1966:106.—Le Loeuff<br />
and Intes, 1968:31, 46, table 1, figs. 51, 63.—Maurin,<br />
1968a:62; 1968b:486, 489, 491.—Le Loeuff and Intes,<br />
1969:66.—Crosnier, 1970:1215 [listed], 1218.<br />
Inachus.—Maurin, 1968b, figs. 4, 9.<br />
?Inachus mauritanicus.—Maurin, 1968b:484 [not Inachus mauritanicus<br />
Lucas, 1846 = /. communissimus Rizza, 1839].<br />
MATERIAL EXAMINED.—Pillsbury Material: Ivory<br />
Coast: Sta 42, 62-75 m, mud with brown, branched Foraminifera,<br />
2(5, 2$ (L). Sta 49, 73-77 m, 1$ (L). Sta 50, 128-<br />
192 m, 19 (L). Sta 62, 46 m, brown, branched and foliate<br />
Foraminifera, 5(5, 1? (L). Sta 63, 64 m, sandy mud with<br />
shells, 3c5, 1$ ov (L). Sta 64, 68 m, 1(5 (L).<br />
Ghana: Sta 16, 46 m, mud with Foraminifera, shells, 19<br />
(L). Sta 22, 51 m, rough bottom, 19 (W). Sta 28, 49-53 m,<br />
2(5, 49 (L).<br />
Nigeria: Sta 239,73 m, 2(5 (L). Sta 241, 59-63 m, mud<br />
and shell, 79 (6 ov) (L).<br />
Cameroon: Sta 259, 59 m, mud and broken shell, 26(5, 219<br />
(3 ov) (W).<br />
Undaunted Material: Angola: Sta 94, 90 m, 1(5, 19 (L). Sta<br />
96, 162 m, 1(5, 19 (L). Sta 103, 90 m, 19 (L).<br />
Other Material: Congo: Off Pointe-Noire, 150 m, slightly<br />
sandy mud, 27 Feb 1967, A. Crosnier, 1(5 (W).<br />
DESCRIPTION.—Capart, 1951:72.<br />
Figures: Capart, 1951, fig. 22, pi. 1: fig. 7;<br />
Monod, 1956, figs. 711-714.<br />
Male Pleopod: Capart, 1951, pi. 2: fig. 10 (Angola);<br />
Monod, 1956, fig. 714 (Senegal).<br />
MEASUREMENTS.—Our specimens have carapace<br />
lengths of 7 to 23 mm; the ovigerous females<br />
have carapace lengths of 12 to 16 mm. The largest
284<br />
specimen recorded by Capart (1951) was 23 mm<br />
long.<br />
REMARKS.—Inachus angolensis, one of five Atlantic<br />
species now known to have a transverse line of<br />
four tubercles on the gastric region of the carapace,<br />
is one of three of those species in which the<br />
interantennular spine is strongly developed, being<br />
clearly visible in dorsal view. It is the only one in<br />
this group of species lacking erect spines on the<br />
cardiac and branchial regions.<br />
In /. angolensis the branchial margin of the<br />
carapace is continuous with the free epineral<br />
margin as described by Forest (1965a) for /. dorsettensis<br />
(Pennant). The dactylus of the chela is<br />
provided with 1 or 2 large basal teeth. The second<br />
pereiopod may be as much as 5 times as long as<br />
the carapace in adult males, and the merus of<br />
that leg may be up to 1.5 times as long as the<br />
carapace. The dactylus of the fifth pereiopod<br />
lacks subapical spinules as in the Mediterranean<br />
/. communissimus Rizza (Forest, 1965a:393).<br />
Monod (1956) suggested that /. angolensis might<br />
be a southern form of /. mauritanicus Lucas [a<br />
junior synonym of/, communissimus Rizza, according<br />
to Forest (1965a) who examined the types of<br />
/. mauritanicus], but /. angolensis differs in numerous<br />
features and, as pointed out by Forest (1965a)<br />
and Forest and Guinot (1966), can be distinguished<br />
by the long interantennular spine alone.<br />
BIOLOGY.—This species apparently prefers soft<br />
bottoms in moderate depths on the shelf and<br />
upper slope. Most of the specimens collected by<br />
the Pillsbury were taken at depths between 46 and<br />
77 m, but one sample came from 128-192 m; the<br />
samples were taken on mud with Foraminifera<br />
and shells and brown, branching and foliate Foraminifera<br />
in 46 m, rough bottom in 51 m, mud<br />
with brown, branched Foraminifera in 62-75 m,<br />
mud and broken shell in 59 m, and sandy mud<br />
with shells in 64 m. The specimen recorded here<br />
from Pointe-Noire was taken in slightly sandy<br />
mud in 150 m. Capart (1951) recorded material<br />
on green mud in 60-110 m, on black, brown mud<br />
in 100 m, on sand and mud in 240 m, and on<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
green sandy mud in 250-300 m. Monod (1956)<br />
did not specify bottom type, but recorded material<br />
in depths between 82 and 350 m.<br />
Crosnier (1964) characterized /. angolensis as a<br />
cold water species, living at depths greater than<br />
50 m. Maurin (1968b) found this species off<br />
Spanish Sahara on muddy detritic bottom in 50-<br />
90 m and off Mauritania on mud or sandy mud<br />
in 90-100 m and on fine detritic sand, occasionally<br />
muddy, in 200-400 m. Le Loeuff and Intes<br />
(1968:46) note that "c'est une espece d'eaux<br />
froides (19°5 a 15°2) et salees (35,4 %o a 35,72<br />
%o)"; they found the species in 80 to 200 m.<br />
Capart (1951) recorded it in 240 m with a bottom<br />
temperature of 10.85°C. Forest and Guinot<br />
(1966) found /. angolensis on sand and compact<br />
sand [sables construit] in 65-75 m and on mud in<br />
50 m.<br />
Ovigerous females have been recorded in February,<br />
March, April, May, June, September, and<br />
October (Capart, 1951; Monod, 1956; Crosnier,<br />
1970; Pillsbury).<br />
DISTRIBUTION.—<strong>West</strong> Africa, from localities between<br />
Spanish Sahara and southern Angola in<br />
depths between 46 and at least 350 m. Capart's<br />
(1951) material came from off the Congo and<br />
Angola and Monod (1956) reported material<br />
from Senegal; since 1956 it has been recorded<br />
from the following localities.<br />
Spanish Sahara: Between Cabo Corbeiro and Cabo<br />
Blanco, 50-90 m (Maurin, 1968b).<br />
Mauritania: Bane d'Arguin, 90-150 m (Maurin, 1968a),<br />
and 40-60, 60-70, and 90-100 m (Maurin, 1968b). Off<br />
Tamzak (as Tamxat), 200-400 m (Maurin, 1968b).<br />
Senegal: Saint-Louis, 75-85 m (Maurin, 1968b).<br />
12°55.5'N, 17°33'W, 65-75 m (Forest and Guinot, 1966).<br />
Ivory Coast: No specific locality, 60+ m (Le Loeuff and<br />
Intes, 1969), 80-200 m (Le Loeuff and Intes, 1968).<br />
Ghana: Off Accra, 60 m (Gauld, 1960). 04°40'N, 02°-<br />
08'W to 04°39'N, 02°05'W, 50 m (Forest and Guinot, 1966).<br />
Cameroon: No specific locality, depths greater than 50 m<br />
(Crosnier, 1964).<br />
Congo: Off Pointe-Noire (Rossignol, 1957). W of Pointe-<br />
Noire (Rossignol, 1962).<br />
Angola: 16°27'S, 11°35'E, 90 m; 16°41'S, 11°21'E, 162<br />
m; and 17°06'S, 11°35'E, 90 m (Crosnier, 1970).
NUMBER 306 285<br />
*Inachus biceps, new species<br />
FIGURES 71, 72<br />
Inachus leptochirus.—Monod, 1956:535, 632, figs. 740-745.—<br />
Longhurst, 1958:89.—Gauld, 1960:72.—Forest and<br />
Guinot, 1966:107. [Not Inachus leptochirus Leach, 1817.]<br />
MATERIAL EXAMINED.—Pillsbury Material: Ghana:<br />
Sta 17, 48 m, fine sand and green mud, 16* (L). Sta 23, 42 m,<br />
foliate brown to orange bryozoans, 6c5, 39 (2 ov) (L). Sta 24,<br />
35-37 m, dark red bryozoans, 14(5 (includes holotype), 14$<br />
(13 ov) (L, W). Sta 26, 27m, shell bottom (scallops), 16\ 2$<br />
(1 ov) (L).<br />
Nigeria: Sta 248, 33 m, Ic5, 2$ ov (L).<br />
Other Material: Dahomey: OffGrand-Popo, 30 m, Petersen<br />
grab, 23 Feb 1964, Guinean Trawling Survey, Tr 34, Sta 2:<br />
16\l$ov(L).<br />
DESCRIPTION.—Carapace (Figure l\a,d)<br />
broader than long. Gastric region with pair of<br />
tubercles anterior to erect gastric spine (Figure<br />
72a). Cardiac region with low, rounded or obtuse<br />
dorsal projection, lower than gastric spine,<br />
FIGURE 71.—Inachus biceps, new species. Ovigerous female,<br />
Ghana: a, dorsal view; b, carapace, lateral view; c, front,<br />
ventral view. Male, Ghana: d, dorsal view; e, first pleopod;<br />
/, apex of first pleopod; g, sternum. (From Monod, 1956, figs<br />
740-745.)<br />
flanked posteriorly on midline by lower prominence.<br />
Branchial regions each with dorsal tubercle,<br />
lateral branchial margin tuberculate. Hepatic<br />
region irregularly tuberculate anteriorly. Rostrum<br />
very short, teeth obtusely rounded, each<br />
usually with minute apical tubercle. Eyes large,<br />
cornea often extending laterally beyond broad<br />
postocular spine. Interantennular spine a very<br />
small, obtuse lobe, not visible in dorsal view,<br />
scarcely or not at all visible in lateral view (Figure<br />
72a). Posterolateral border of carapace continuous<br />
with free epimeral margin. Basal antennal<br />
segment (Figure 12a,b) with row of tubercles,<br />
some sharp, on mesial and lateral margins, anterolateral<br />
angle of basal segment lacking prominent<br />
spine.<br />
Chelipeds (Figure 12c) enlarged in adult males,<br />
about 1.5 times as long as carapace, merus and<br />
propodus markedly inflated in adult. Chelipeds<br />
in females slender, subequal to or slightly longer<br />
than carapace. Merus and carpus spiny in both<br />
sexes. Palm with row of large tubercles proximally<br />
on outer face in males. Palm depth 1/2 length in<br />
females, 2/3 length in males. Fingers subequal to<br />
or slightly longer than palm. Cutting edges of<br />
fingers unarmed basally in females, dactylus with<br />
2 prominent proximal teeth in males.<br />
First walking leg 4 or slightly more than 4 times<br />
as long as carapace, merus, propodus, and dactylus<br />
each longer than carapace in males, relatively<br />
shorter in females. Dactylus unarmed, sinuous<br />
distally, apex gently curved, length subequal<br />
to or slightly greater than that of propodus. Second<br />
to fourth walking legs decreasing in size<br />
posteriorly, fourth about twice carapace length in<br />
females, slightly longer in males. Second walking<br />
leg scarcely overreaching propodus of first by tip<br />
of dactylus. Latter sinuous, gently curved distally,<br />
shorter than propodus, with 1 or 2 subdistal teeth<br />
ventrally. Third walking leg not extending to end<br />
of propodus of second, overreaching carpus of<br />
second by distal third of propodus. Dactylus more<br />
curved than that of preceding leg, shorter than<br />
propodus, with 2 subdistal teeth ventrally. Fourth<br />
walking leg rather short, not extending to end of<br />
propodus of third. Dactylus short, more strongly
286<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 72.—Inachus biceps, new species, paratype, male, cl 8.3 mm, Pillsbury Sta 24: a, front,<br />
lateral view; b, front, ventral view; c, chela; d, abdomen.<br />
curved than those of preceding legs but not sickleshaped,<br />
length across arc subequal to that of<br />
propodus, armed ventrally with 1 or 2 large,<br />
subdistal teeth and several teeth proximally.<br />
Male abdomen (Figure 72
NUMBER 306 287<br />
Inachus biceps can be distinguished readily from<br />
the other <strong>West</strong> <strong>African</strong> species which have two<br />
gastric tubercles and a sternal callosity in the<br />
male. The new species differs from /. aguiarii De<br />
Brito Capello in several features: the interantennular<br />
spine is smaller; there is a dorsal tubercle<br />
rather than a spine on the branchial region; and<br />
the sternal callosity is subcircular in shape. In /.<br />
aguiarii the sternal callosity is developed in both<br />
sexes, rather than only in the males as in /. biceps.<br />
Neither of the other <strong>West</strong> <strong>African</strong> species with<br />
two gastric tubercles, /. phalangium (Fabricius) or<br />
/. guentheri (Miers), have the sternal callosity developed<br />
in either sex.<br />
TYPE-LOCALITY.—Off Ghana, 04°56'N, 00°-<br />
47.5'W to 04°56'N, 00°50'W, in 35 to 37 m<br />
(Pillsbury Sta 24).<br />
DISPOSITION OF TYPES.—The holotype (Crust.<br />
D. 31770), a male, cl 9.4 mm, cb 10.3 mm, from<br />
Pillsbury Sta 24, is in the Rijksmuseum van Natuurlijke<br />
Historie, Leiden. Paratypes are in the<br />
collections of the National Museum of Natural<br />
History, <strong>Smithsonian</strong> <strong>Institution</strong>, and the Rijksmuseum<br />
van Natuurlijke Historie at Leiden.<br />
ETYMOLOGY.—The specific epithet is from the<br />
English derivative of the Latin musculus biceps,<br />
alluding to the inflated cheliped.<br />
BIOLOGY.—Inachus biceps is a sublittoral species,<br />
occurring in depths between 27 and 300 m. Only<br />
two of the 16 depth records available are from<br />
relatively deep water, namely off Sierra Leone in<br />
220-240 m (Monod, 1956) and 300 m (Longhurst,<br />
1958); the remainder of the records are from<br />
depths of less than 60 m. Material from the two<br />
deepest records should be reexamined. The Pillsbury<br />
collected the species off Ghana, where it<br />
appears to be relatively common (28 specimens<br />
at Sta 24), on fine sand and green mud and on<br />
bottom with bryozoans or shell. It was taken by<br />
the Calypso on shells, and mud with Area.<br />
Ovigerous females have been collected in February,<br />
May, and November (Monod, 1956; Forest<br />
and Guinot, 1966; present paper).<br />
DISTRIBUTION—Inachus biceps is a tropical <strong>West</strong><br />
<strong>African</strong> species, known from a few localities between<br />
Senegal and Nigeria in depths between 27<br />
and 300 m. The following records are in the<br />
literature:<br />
Senegal: S of Goree, 40-41 and 46-50 m (Monod, 1956).<br />
Sierra Leone: No specific locality, 220-240 m (Monod,<br />
1956), 300 m (Longhurst, 1958).<br />
Ghana: Off Accra, 43, 44, and 51 m (Monod, 1956) and<br />
44-51 m (Gauld, 1960).<br />
Nigeria: Off the mouths of the Niger River, 04°03'N,<br />
06°12'E, 32 m (Forest and Guinot, 1966).<br />
It has not been recorded previously from Dahomey,<br />
although that is well within its known<br />
range.<br />
Inachus dorsettensis (Pennant, 1777)<br />
Cancer Dorsettensis Pennant, 1777:8, pi. 9A: fig. 18.<br />
Inachus dorsettensis.—A. Milne Edwards and Bouvier, 1899:45<br />
[key], 46 [Azores]; 1900:143 [?part; Ilhas Desertas; Spanish<br />
Sahara].—Balss, 1922:72 [listed].—Bouvier, 1922:79 [Canary<br />
Islands].—Monod, 1933b:503 [p. 48 on separate;<br />
listed].—Capart, 1951:70 [part; Spanish Sahara].—<br />
Maurin, 1968a:31 [Morocco]; 1968b:489 [Mauritania].—<br />
Zariquiey Alvarez, 1968:472, figs, 157f, 159b, 160b [Spain;<br />
references].—Christiansen, 1969:100, fig. 41, map 34<br />
[North Atlantic ].—Tiirkay, 1976a:26 [listed], 40, fig. 34<br />
[Portugal, Morocco].<br />
SYNONYMS.—ICancer dodecos Linnaeus, 1767;<br />
Macropus parvirostris Risso, 1816; Doclea fabriciana<br />
Risso, 1827.<br />
DISTRIBUTION.—Eastern Atlantic, from the Hebrides<br />
southward to Mauritania (?), including the<br />
Azores and the Canary Islands; Mediterranean;<br />
?South Africa. Sublittoral, from a few meters to<br />
about 110 m. All of these records require verification.<br />
* Inachus grallator, new species<br />
FIGURES 73, 74<br />
PInachus dorsettensis.—A. Milne Edwards and Bouvier, 1900:<br />
143 [part?: Cape Verde Islands, 75-90 and 318 m].—<br />
Doflein, 1904:72 [Bane de la Seine; Spanish Sahara].—<br />
Odhner, 1923:19 [Angola, 72-108 m].—Monod, 1956:526<br />
[? part].—Longhurst, 1958:89 [Sierra Leone, 72-118<br />
m].—Guinot and Ribeiro, 1962:77 [Angola, 150-220 m].<br />
Inachus dorsettensis.—Capart, 1951:70, fig. 21, pi. 1: fig. 6, pi.<br />
2: fig. 11 [part, not specimens from Spanish Sahara?].—
288<br />
Crosnier, 1970:1218. [Not Inachus dorsettensis (Pennant,<br />
1777).]<br />
MATERIAL EXAMINED.—Pillsbury Material: Nigeria:<br />
Sta 255, 264-269 m, 36 (includes holotype), 1$ (L, W).<br />
Undaunted Material: Angola: Sta 96, 162 m, 1? (L).<br />
Other Material: Congo: 05°03'S, 11°23'E, 247-250 m,<br />
sandy mud, 23 Jan 1968, A. Crosnier, 3$ ov (W).<br />
DESCRIPTION.—Carapace (Figure 73a) distinctly<br />
longer than broad, length 1.06 to 1.12<br />
times width. Gastric region of carapace with 4<br />
tubercles in transverse row anterior to erect gastric<br />
spine. Cardiac region of carapace with erect<br />
dorsal spine. Each branchial region with shorter<br />
erect spine, not markedly recurved anteriorly,<br />
and with dorsal tubercle anterior to each spine.<br />
Hepatic lobe with 2 distinct tubercles, and<br />
smaller, less conspicuous tubercles scattered on<br />
surface. Rostrum (Figures 73b, 744) short, spines<br />
broad, lateral margins convergent anteriorly.<br />
Eyes large, but not extending laterally beyond<br />
strong postorbital spine. Interantennular spine<br />
well developed, clearly visible in dorsal view (Figure<br />
73a,b) overreaching rostral spines in some<br />
specimens. Branchial margin of carapace tuberculate,<br />
anteriorly continuous with free epimeral<br />
margin. Basal antennal segment (Figure 73b)<br />
with row of tubercles, posteriormost largest, anterolateral<br />
angle lacking prominent spine.<br />
Chelipeds subequal to or slightly longer than<br />
carapace, slender, not markedly inflated (slightly<br />
inflated in some males), equal and similar in both<br />
FIGURE 73.—Inachus grallator, new species: a, dorsal view; b,<br />
ventral view of front; c, male pleopod. (All from Capart,<br />
1951, fig. 21, pi. 1: fig. 6, pi. 2: fig. 11.)<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
sexes; ischium with inner row of 4 or 5 erect<br />
tubercles; merus rounded, with low, blunt spines<br />
and tubercles, primarily on ventral surface; carpus<br />
with scattered small tubercles and spinules;<br />
palm covered with small, blunt spines; dactylus<br />
much longer than palm, cutting edge lacking<br />
enlarged proximal tooth or teeth.<br />
Second pereiopod (Figure 74c) about 4 (3.8 to<br />
4.5, mean 4.1) times as long as carapace, very<br />
slender, merus, propodus, and dactylus each<br />
longer than carapace; dactylus unarmed, subequal<br />
to or slightly shorter than propodus. Third<br />
pereiopod not extending to end of propodus of<br />
second, overreaching carpus of second by half of<br />
propodus and all of dactylus; latter shorter than<br />
propodus, lacking subdistal spines ventrally.<br />
Fourth pereiopod extending to or slightly overreaching<br />
propodus of third; dactylus slightly<br />
curved, with 2 low, subdistal tubercles, subequal<br />
to propodus. Fifth pereiopod (Figure 74d,e)<br />
scarcely overreaching propodus of fourth by tip<br />
of dactylus; latter similar to those of third and<br />
fourth pereiopods, subequal to propodus, with 2<br />
subapical tubercles apically. Male pleopod shown<br />
in Figure 73c.<br />
MEASUREMENTS.—Carapace lengths of males<br />
14.2 to 15.5 mm, of non-ovigerous females 12.7 to<br />
13.0 mm, of ovigerous females 14.5 to 14.6 mm.<br />
REMARKS.—Inachus grallator is the southern<br />
counterpart of Inachus dorsettensis, which it closely<br />
resembles. It may be a smaller species than /.<br />
dorsettensis, Atlantic populations of which may<br />
attain a carapace length of 30 mm (Christiansen,<br />
1969), almost twice as large as adult specimens of<br />
/. grallator available for study. It also appears to<br />
live in deeper water, occurring off <strong>West</strong> Africa in<br />
depths of 162-269 m. Zariquiey Alvarez (1968)<br />
noted that /. dorsettensis is usually found at depths<br />
of 30-40 m in Spain, and Christiansen (1969)<br />
recorded it from 6-10 m to about 110 m, and,<br />
because Bouvier (1940) noted that it occurred<br />
from the shore to 550 m, it seems possible that he<br />
was dealing with more than one species. Material<br />
of /. dorsettensis from Spain in the collections of<br />
the <strong>Smithsonian</strong> <strong>Institution</strong> comprises more than<br />
30 lots from the Ria de Arosa, where it is common
NUMBER 306 289<br />
FIGURE 74.—Inachus grallator, new species, male paratype, cl 14.2 mm, Pillsbury Sta 255: a,<br />
carapace, lateral view; b, front, dorsal view; c, second pereiopod; d, fifth pereiopod; e, dactylus<br />
of fifth pereiopod. Inachus dorsettensis (Pennant), male, cl 15.1 mm, Spain:/, front, dorsal view;<br />
g, second pereiopod; h, fifth pereiopod; i, dactylus of fifth pereiopod.<br />
in depths of less than 40 m, although 8 lots were<br />
taken in depths between 45 and 70 m.<br />
The best way to distinguish /. grallator from /.<br />
dorsettensis is by the slenderness of the pereiopods<br />
and the shape of the dactyli of the fifth pereio-<br />
pods. In /. grallator, the pereiopods are slenderer,<br />
appearing longer, and the dactyli of the fifth<br />
pereiopods are much less curved, being subsimilar<br />
to those of the fourth pereiopods (compare Figure<br />
74d,e with 746,i), as well as comparatively longer.
290<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
In adult females of /. grallator the dactylus of the Griffin was misled in using the accounts of Monod<br />
fifth pereiopod is longer than the carapace, (1956) and Barnard (1950) to help identify his<br />
whereas in /. dorsettensis it is shorter. In females of material with /. dorsettensis. Monod was not deal-<br />
the new species the pereiopods are slightly stouter ing with /. dorsettensis but with an undescribed<br />
than in males, but they are noticeably slenderer species, /. nanus, new species (p. 291). Although<br />
than the same pereiopods of /. dorsettensis. we have seen no material from South Africa,<br />
Although Forest (1965a:392-393), in differen- Barnard's (1950) account of/, dorsettensis suggests<br />
tiating the Mediterranean /. communissimus Rizza that he was dealing with /. complectens, a much<br />
from /. dorsettensis, stated that in the latter species spinier species with sharper rostral teeth, in which<br />
the carapace is longer than broad (mean length/ the four dorsal spines of the carapace all attain<br />
width ratio of 1.1 to 1), that the second pereiopod<br />
was three times as long as the carapace, and that<br />
the merus of the second leg was equal to the<br />
carapace, our material from Spain suggests that<br />
/. dorsettensis may be somewhat variable in these<br />
features. Our specimens have the carapace<br />
length/width ratio varying from 0.97 in ovigerous<br />
females to 1.04 in males; the carapace of adult<br />
females is noticeably broader than long. In our<br />
specimens the second pereiopods are 3.29 to 4.09<br />
(mean 3.65) times as long as the carapace, being<br />
comparatively longer in males, and, in all of the<br />
specimens measured, the merus is perceptibly<br />
longer than the carapace.<br />
As Forest (I965a:391) noted, Atlantic specimens<br />
of /. dorsettensis appear to be much larger<br />
than do Mediterranean specimens; the former<br />
may attain a carapace length of 30 mm, whereas<br />
the latter rarely exceed 20 mm in length. Our<br />
material suggests that in the Mediterranean population<br />
the dorsal spines of the carapace are much<br />
slenderer and higher than they are in material<br />
the same height (in /. dorsettensis, I. grallator, and<br />
/. nanus, the median spines are higher than the<br />
branchials). A redescription of /. complectens<br />
(Rathbun) is in preparation.<br />
Specimens identified with /. dorsettensis from<br />
<strong>West</strong> <strong>African</strong> localities by A. Milne Edwards and<br />
Bouvier (1900), Odhner (1923), Longhurst<br />
(1958), Guinot and Ribeiro (1962), and, possibly,<br />
part of the material referred to /. dorsettensis by<br />
Monod (1956) may be referable to /. grallator; all<br />
of these records have been questioned in the<br />
synonymy, above. Both Doflein and Odhner commented<br />
on the small size of specimens they identified<br />
with /. dorsettensis from deep water off <strong>West</strong><br />
Africa. All of their material, as well as all of the<br />
material listed above under /. dorsettensis from<br />
NW <strong>African</strong> localities, should be restudied.<br />
Some of the deeper records for /. dorsettensis,<br />
assigned to /. nanus, new species (p. 291), may<br />
well be based on /. grallator. The specimen from<br />
Senegal illustrated by Monod (1956, fig. 720;<br />
from Atlantic localities except in very large fe- Figure 75*/ herein) with long dorsal spines resemmales<br />
in which the median spines usually are bles /. grallator in that feature, but the interanten-<br />
worn down. If the Mediterranean population nular spine appears to be small as in typical /.<br />
proves to be distinct, the name Inachus parvirostris nanus.<br />
(Risso, 1816), which Risso originally proposed as ETYMOLOGY.—The specific epithet is derived<br />
Macropus parvirostris, is available.<br />
from the Latin and means "one who goes on<br />
Although Griffin (1974) synonymized Pseudo- stilts," alluding to the long, slender legs of this<br />
collodes complectens Rathbun, 1911, with /. dorsetten- species.<br />
sis (Pennant, 1777), reexamination of Griffin's TYPE-LOCALITY.—Gulf of Guinea off Nigeria,<br />
material from the western Indian Ocean, as well 03°49'N, 07°38'E to 03°48'N, 07°42'E, in 264as<br />
the type of P. complectens, leads us to believe 269 m (Pillsbury Sta 255).<br />
that the Indian Ocean form is a distinct species. DISPOSITION OF TYPES.—The holotype (Crust.<br />
As noted above Griffin was correct in synonymiz- D. 27154) a male from Pillsbury Sta 255, is in the<br />
ing Pseudocollodes with Inachus. We suspect that Rijksmuseum van Natuurlijke Historie, Leiden;
NUMBER 306 291<br />
paratypes also are in that collection and in the<br />
National Museum of Natural History, <strong>Smithsonian</strong><br />
<strong>Institution</strong>, Washington, D.C.<br />
BIOLOGY.—Inachus grallator is a deep water species,<br />
known to occur in depths between 100 m<br />
and 250-300 m on soft bottom; Capart (1951)<br />
reported specimens from green mud, brown sandy<br />
mud, muddy sand and green muddy sand. The<br />
specimens recorded herein from the Congo were<br />
taken on sandy mud in 247-250 m.<br />
Ovigerous females have been recorded in January<br />
and November (Capart, 1951; present paper).<br />
DISTRIBUTION.—<strong>West</strong> Africa, from a few localities<br />
between Nigeria and Angola. Records in the<br />
literature include the following:<br />
Gabon: 45 mi [72.5 km] NxE of Port-Gentil, 00°S, 08°-<br />
58'E, 250-300 m (Capart, 1951).<br />
Angola: 25 mi [40 km] WNW of Ambriz, 07°39'S,<br />
12°47'40"E, 100 m; 8 mi [12.8 km] NxE of Baia dos Elefantes,<br />
13°05'S, 12°46'E, 100-110 m; 18 mi [29 km] WSW<br />
of Baia dos Tigres, 16°36'S, 11°27'E, 110 m (all Capart,<br />
1951). 16°37'S, 11°22'E, 126 m (Crosnier, 1970).<br />
Inachus guentheri (Miers, 1879)<br />
Inachus guentheri.—Monod, 1956:529, figs 723-730 [Senegal,<br />
Gabon, Angola].<br />
SYNONYM.—Inachus antarcticus Doflein, 1904.<br />
DISTRIBUTION.—Off <strong>West</strong> Africa and South<br />
Africa, sublittoral, from about 18 to more than<br />
180 m.<br />
Inachus leptochirus Leach, 1817<br />
Inachus leptochirus Leach, 1817, in 1815-1875, pi. 22B.—A.<br />
Milne Edwards and Bouvier, 1894:7 [Azores]; 1899:45<br />
[key]; 1900:145 [part; Spain; Azores; Mauritania].—Doflein,<br />
1904:73 [Seine Seamount].—Bouvier, 1922:79, pi. 2:<br />
fig. 5 [color] [Josephine Seamount; Princesse Alice<br />
Bank; S of Almadena].—Balss, 1922:72 [listed].—Monod,<br />
1933b:503 [p. 48 on separate] [listed].—Chapman and<br />
Santler, 1955:376 [Azores].—Maurin, 1968b:484 [Spanish<br />
Sahara].—Zariquiey Alvarez, 1968:472, fig. 157a,b<br />
[Spain; references].—Christiansen, 1969:104, fig. 43, map<br />
36 [North Atlantic ].—Tiirkay, 1976a:26 [listed], 40, fig.<br />
35 [Morocco].<br />
SYNONYM.—Inachus affinis Rizza, 1839.<br />
DISTRIBUTION.—Eastern Atlantic, from the He-<br />
brides southward to Mauritania, including the<br />
Azores; Mediterranean. Sublittoral, from about<br />
30 m to more than 500 m.<br />
*Inachus nanus, new species<br />
FIGURE 75<br />
Inachus dorsettensis.—Monod, 1956:526, 632, figs. 715-722<br />
[?part].—Rossignol, 1957:116 [key]. [Not Inachus dorsettensis<br />
(Pennant, 1777).]<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 68, 70 m, broken shell, 36, 29 ov (L). Sta 69, 29 m, coral<br />
or rock, 1(5 (W).<br />
Ivory Coast: Sta 42,62-75 m, mud with brown, branched<br />
Foraminifera, 1
292<br />
a<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 75.—Inachus nanus, new species: a, dorsal view; b, front, ventral view; c, carapace, lateral<br />
view; d, carapace of different specimen, lateral view (this could be based on /. grallator, new<br />
species); e, male pleopod. (All from Monod, 1956, figs. 715-717, 720, 722.)<br />
half the carpus beyond front. Chelipeds in females<br />
slenderer than in males, about 1.1 times as long<br />
as carapace, carpus not extending to front. Chelipeds<br />
in both sexes with lower margin of merus<br />
and inner border of carpus very spiny. Palm<br />
tuberculate in both sexes, length about 1.7 times<br />
height in females, 1.3-1.4 times greatest height in<br />
males. Dactylus of chela with 2 rounded teeth<br />
proximally on cutting edge in males.<br />
Second pereiopod about 4 times carapace
NUMBER 306 293<br />
length in males, about 3.5 times carapace length<br />
in females (most second pereiopods detached);<br />
dactylus with minute tubercles ventrally, shorter<br />
than propodus, latter slightly shorter than merus.<br />
Dactylus of third pereiopod not overreaching propodus<br />
of second. Second to fifth pereiopods decreasing<br />
in size posteriorly, fifth about twice as<br />
long as carapace in males, about 1.5 times as long<br />
as carapace in females. Dactylus of fifth pereiopod<br />
curved but not strongly falcate, with 1 or 2<br />
large, subdistal spinules flanked proximally by<br />
line of smaller spinules and tubercles.<br />
Terminal somite of male abdomen rounded<br />
apically. Male pleopod as shown (Figure I5e).<br />
MEASUREMENTS.—Carapace lengths of males<br />
4.9 to 8.9 mm, of ovigerous females 4.8 to<br />
8.3 mm.<br />
REMARKS.—Inachus nanus resembles the Mediterranean<br />
/. communissimus Rizza and differs from<br />
the other Atlantic species of Inachus, which have<br />
a transverse row of four tubercles on the gastric<br />
region. It differs from /. dorsettensis (Pennant) and<br />
/. grallator, new species, in having a very small<br />
interantennular spine, which is not visible between<br />
the teeth of the rostrum in dorsal view.<br />
Inachus nanus, which has a maximum carapace<br />
length of less than 10 mm, is a much smaller<br />
species than /. communissimus; in the latter species<br />
Forest (1965a) recorded specimens with carapace<br />
lengths up to 26 mm. The carapace in /. nanus is<br />
longer than broad or about as broad as long; in<br />
/. communissimus it is much broader than long. In<br />
/. nanus the posterolateral margin of the carapace<br />
is continuous with the free epimeral margin<br />
whereas in /. communissimus it is not continuous,<br />
being interrupted on the anterior part of the<br />
branchial region.<br />
Some of the <strong>West</strong> <strong>African</strong> records of/, dorsettensis,<br />
referred above to /. grallator, new species, may<br />
well be based on /. nanus; this can only be determined<br />
by examining the material.<br />
TYPE-LOCALITY.—Gulf of Guinea off the Ivory<br />
Coast, 05°02.5'N, 03°49.5'W to 05°05'N, 03°-<br />
52*W, in 62-75 m {Pillsbury Sta 42).<br />
DISPOSITION OF TYPES.—The holotype (Crust.<br />
D. 27155), a male, cl 8.3 mm, cb 7.8 mm, is in<br />
the Rijksmuseum van Natuurlijke Historie, Leiden;<br />
paratypes also are deposited there and in the<br />
National Museum of Natural History, <strong>Smithsonian</strong><br />
<strong>Institution</strong>.<br />
ETYMOLOGY.—The specific epithet is from the<br />
Latin and refers to the diminutive size of this<br />
species.<br />
BIOLOGY.—Inachus nanus is a sublittoral species,<br />
occurring in depths between 29 and 118 m. Only<br />
three of the 21 depth records available are from<br />
100 m or more and only one is from less than 40<br />
m. Although three of the lots taken by the Pillsbury<br />
were from rough bottom, coral or rock, most of<br />
the specimens were taken on mud, either with<br />
Jullienella or bryozoans. Some of the deeper records<br />
in Monod (1956) may be based on specimens<br />
of/, grallator.<br />
Ovigerous females have been taken in February,<br />
May, June, July, September, and October,<br />
suggesting that the species spawns all year<br />
(Monod, 1956; Pillsbury).<br />
DISTRIBUTION.—Inachus nanus is a sublittoral<br />
<strong>West</strong> <strong>African</strong> species, known from a few localities<br />
between Senegal and Gabon. The following records<br />
are all from Monod (1956):<br />
Senegal: S of Cap Vert, 97-98 m; Fosse de Kayar, 100 m;<br />
S of Fosse de Kayar, 50 m; SE of lie de la Madeleine, Dakar,<br />
48 m; off Goree, 95, 96, and 132 m.<br />
Sierra Leone: No specific locality, 118 m.<br />
Gabon: Libreville, 60 m.<br />
It has not previously been recorded from Liberia,<br />
Ivory Coast, Ghana, Nigeria, or Cameroon,<br />
but all of these lie within its known range.<br />
Inachus phalangium (Fabricius, 1775)<br />
Inachus phalangium.—Monod, 1956:531 [references].—Zariquiey<br />
Alvarez, 1968:472, fig. 159c [Spain; references].—<br />
Christiansen, 1969:102, fig. 42, map 35 [North Atlantic].<br />
Inachus dorhynchus.—Rossignol, 1957:115 [key].—Forest and<br />
Games, 1960:357 [Morocco].<br />
SYNONYMS.—ICancer tribulus Linnaeus, 1767<br />
[name suppressed by ICZN]; Inachus dorynchus<br />
Leach, 1814; Macropus aracnides Risso, 1816.<br />
DISTRIBUTION.—Eastern Atlantic, from about<br />
60°N latitude southward to the Cape Verde Is-
294<br />
lands; Mediterranean. Sublittoral, from about 10<br />
to at least 150 m.<br />
Inachus thoracicus Roux, 1830<br />
lnachus thoracicus.—Capart, 1951:68 [Congo].—Monod, 1956:<br />
532, figs. 731-735 [Senegal, Gambia].—Rossignol, 1957:<br />
115 [key].—Longhurst, 1958:89 [Sierra Leone].—Maurin,<br />
1968a:48, 59; 1968b:480, 484, 486, fig. 5 [both Spanish<br />
Sahara, Mauritania].—Zariquiey Alvarez, 1968:473, figs.<br />
157c,d, 158a,b [Spain; references].<br />
SYNONYM.—Inachus cocco Rizza, 1839.<br />
DISTRIBUTION.—Eastern Atlantic, from southern<br />
Europe to the Congo, including the Canary<br />
Islands; Mediterranean. Sublittoral, to at least<br />
100 m.<br />
Genus Macropodia Leach, 1814<br />
Macropodia Leach, 1814:395 [type-species: Cancer longirostris<br />
Fabricius, 1775, by monotypy; gender: feminine; name<br />
1699 on Official List].<br />
Peridromus Gistel, 1848:ix [replacement name for Macropodia<br />
Leach, 1814 (as Macropus); type species: Cancer longirostris<br />
Fabricius, 1775; gender: masculine].<br />
REMARKS.—Specific identification of the eastern<br />
Atlantic species of Macropodia has remained<br />
particularly difficult until recently when Forest<br />
and Zariquiey Alvarez (1964) reviewed the Mediterranean<br />
species and their nomenclature (Forest,<br />
1965b). They pointed out that in the past<br />
identification often was based on keys provided<br />
by A. Milne Edwards and Bouvier (1899:48) or<br />
Bouvier (1940:362) and that results obtained<br />
through using these keys generally were unsatisfactory.<br />
Forest and Zariquiey Alvarez (1964) recognized<br />
five species in the Mediterranean: M. rostrata<br />
(Linnaeus, 1761); M. longipes (A. Milne Edwards<br />
and Bouvier, 1899); M. longirostris (Fabricius,<br />
1775); M. czernjawskii (Brandt, 1880); and a<br />
new species they named M. linaresi.<br />
In discussing the literature relating to these<br />
species, Forest (1965b:349) pointed out that the<br />
epithet longirostris in the past had been applied to<br />
three distinct taxa: M. longirostris proper, M. longipes,<br />
and a third species from areas in the north-<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
eastern Atlantic north of the Mediterranean, M.<br />
tenuirostris Leach, 1814. Thus many earlier records<br />
of M. longirostris from the Mediterranean were<br />
based on that species or M. longipes.<br />
Monod (1956:559) included M. longirostris as a<br />
member of the <strong>West</strong> <strong>African</strong> fauna, based on<br />
earlier records by A. Milne Edwards and Bouvier<br />
(1900:156, 157) who reported two males and a<br />
female from the Bane d'Arguin, one male from<br />
Senegal, and one female from the Cape Verde<br />
Islands, as well as material from Morocco. Forest<br />
and Zariquiey Alvarez (1964:228) considered M.<br />
longirostris to be a Mediterranean endemic and<br />
identified many of A. Milne Edwards and Bouvier's<br />
records of M. longirostris with M. longipes,<br />
noting (1964:226, footnote) that material reported<br />
by these authors from three localities,<br />
including Morocco and Senegal, no longer was in<br />
the collection at Paris. Apparently there are no<br />
substantiated records for either M. longipes or M.<br />
longirostris from tropical <strong>African</strong> waters, and we<br />
have omitted these species from our accounts<br />
below.<br />
Material from the third locality, the Strait of<br />
Bonifaccio (between Corsica and Sardinia), is in<br />
the collection of the National Museum of Natural<br />
History, <strong>Smithsonian</strong> <strong>Institution</strong>, under catalog<br />
number USNM 22978. It consists of a single<br />
male, cl 15.9 mm, clearly identifiable with M<br />
longirostris. Outside of the Mediterranean, M. longipes<br />
is known with certainty from the Bay of<br />
Biscay (Forest and Zariquiey Alvarez, 1964:226).<br />
The eastern Atlantic species of Macropodia still<br />
require a great deal of work, especially the status<br />
of the species occurring on the NW <strong>African</strong> coast,<br />
including M. longirostris auctorum (including<br />
Peres, 1964:20, Morocco), M. longipes auctorum<br />
(Forest and Gantes, 1960:357, Morocco; Bas, Arias,<br />
and Guerra, 1976, table 3, Spanish Sahara;<br />
Tiirkay, 1976a:40, fig. 33, Portugal and Morocco),<br />
M. egyptia sensu A. Milne Edwards and<br />
Bouvier, 1900, and M. intermedia Bouvier, 1940<br />
(see p. 300). We believe that the status of Mediterranean<br />
material identified with M. rostrata<br />
(Linnaeus, 1761) also requires investigation.
NUMBER 306 295<br />
Key to Tropical <strong>West</strong> <strong>African</strong> Species of Macropodia<br />
1. Dactylus of fifth leg slightly curved 2<br />
Dactylus of fifth leg strongly arcuate 4<br />
2. Rostrum longer than antennal peduncle, more than 2/3 postrostral length<br />
of carapace. [Nuchal spine present. Basal article of antenna with strong<br />
spines ventrally] M. longipes<br />
Rostrum shorter than antennal peduncle, less than half postrostral length<br />
of carapace 3<br />
3. Rostrum extending beyond midlength of distal article of antennal peduncle.<br />
Nuchal spine present. Orbital margin with dorsal spines or tubercles.<br />
Basal article of antenna with strong spines ventrally M. gilsoni<br />
Rostrum falling short of midlength of distal article of antennal peduncle.<br />
Nuchal spine absent. Orbital margin smooth. Basal article of antenna<br />
smooth, unarmed M. spinulosa<br />
4. Epistome with longitudinal ridge on each side extending from pore of<br />
antennnal gland to basal article of antenna M. macrocheles<br />
Epistome lacking longitudinal ridge on each side 5<br />
5. Rostrum longer than antennal peduncle. Basal segment of antenna<br />
unarmed 6<br />
Rostrum not reaching end of antennal peduncle. Basal segment of antenna<br />
armed with spinules and tubercles 7<br />
6. Rostrum more than 2/3 postrostral length of carapace. Basal article of<br />
antenna smooth M. longicornis<br />
Rostrum less than half postrostral length of carapace. Basal article of<br />
antenna irregularly tuberculate M. straeleni<br />
7. Rostrum extending beyond midlength of distal segment of antennal peduncle.<br />
Gastric region with sharp, erect spine ... M. doracis, new species<br />
Rostrum not extending to midlength of distal segment of antennal peduncle.<br />
Gastric region with low, conical prominence<br />
Af. hesperiae, new species<br />
Macropodia doracis, new species Rostrum (Figure 76a) moderately long, extending<br />
almost to but falling short of end of distal<br />
FIGURE 76 segment of antennal peduncle, ornamented with<br />
„ , . , „ , , , „ . curved hairs. Apices of rostral teeth appressed.<br />
Stenorhynchus aeeyptius.—A. Milne Edwards and Bouvier, r \ r .<br />
1900:155 [part] [not Sunorynchus tgyptius H. Milne Ed- Rostrum upturned dorsally. Nuchal region with<br />
wards, 1834 = Cancer longirostns Fabricius, 1775]. lateral spinule, visible in dorsal view (Figure 76a).<br />
MATERIAL EXAMINED.—PilUbury Material: None.<br />
Other Material: Cape Verde Islands: between Ilheu Branco<br />
and Ilheu Raso, 110-180 m, sand, rocks, Talisman No. 105,<br />
Protogastric region smooth, lacking dorsal tubercies<br />
medially. Hepatic regions each with low,<br />
conical prominence dorsally. Hepatic lobes each<br />
- i i i i J J L I n r><br />
,, ¥ , " „ „ nk, , , s<br />
27Jul 1883, l9ov(W, holotype).<br />
with bluntly rounded tubercle apically. Gastric<br />
r 7<br />
. . J .<br />
region with high, erect spine dorsally. Cardiac<br />
DESCRIPTION.—Size small, carapace length of region with broad, obtuse prominence dorsally,<br />
adult less than 10 mm (7.8 mm). lower than gastric spine. Branchial regions each
296<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 76.—Macropodia doracis, new species, holotype, ovigerous female, cl 7.8 mm, Cape Verde<br />
Islands: a, carapace, dorsal view; b, carapace, lateral view; c, chela; d, third pereiopod; e, fourth<br />
pereiopod;/, fifth pereiopod. (All pereiopods slightly damaged.)<br />
with 1 low dorsal epibranchial tubercle, 1 broadly<br />
rounded anterolateral epibranchial tubercle, 1<br />
dorsal mesobranchial tubercle, and 1 metabranchial<br />
tubercle situated almost on posterior margin.<br />
Anterolateral margins with rounded tubercle<br />
posterior to hepatic lobe.<br />
Basal antennal segment (Figure 76b) with 2<br />
tubercles on each side, larger, almost spiniform,<br />
anteriormost. Distal article of antennal peduncle<br />
about twice as long as penultimate. Epistome<br />
with rounded lobule near opening of antennal<br />
gland and with tubercle at base of antenna.<br />
Ocular peduncle lacking distinct anterior lobe.<br />
Cornea with distal dorsal tubercle.<br />
Cheliped (Figure 76V) of adult female slender,<br />
fingers, measured dorsally, longer than palm, latter<br />
with row of spinules dorsally. Merus with 2<br />
prominent dorsal tubercles.<br />
Dactylus of second pereiopod slender, slightly<br />
curved distally, apex broken; merus shorter than<br />
carapace. Merus of each walking leg with blunt,<br />
unarmed dorsal projection (Figure 76d). Dactyli<br />
of fourth and fifth pereiopods (Figure 76eJ)<br />
strongly arcuate, ventral margin toothed along<br />
entire margin.<br />
MEASUREMENTS.—Carapace length of ovigerous<br />
female 7.8 mm.<br />
REMARKS.—Macropodia doracis approaches M.<br />
hesperiae, (p. 298), in general facies, but differs in<br />
having a longer rostrum, which extends almost to
NUMBER 306 297<br />
the end of the antennal peduncle, in having a<br />
shorter distal segment on the antennal peduncle,<br />
and in having a sharp, erect gastric spine. The<br />
present species may be closest to Macropodia aegyptia<br />
sensu Bouvier (1940:364, fig. 220), from the<br />
Canary Islands, but in that species the epistome<br />
has more spines and the basal segment of the<br />
antennal peduncle is armed with three large<br />
sharp spines rather than a spinule and a tubercle.<br />
TYPE-LOCALITY.—Cape Verde Islands, between<br />
Ilheu Branco (16°39'N, 24°41'W) and IIheu<br />
Raso (16°37'N, 24°36 / W) in 110-180 m.<br />
DISPOSITION OF TYPES.—The unique holotype<br />
(USNM 22980), an ovigerous female, cl 7.8 mm,<br />
is in the National Museum of Natural History,<br />
<strong>Smithsonian</strong> <strong>Institution</strong>. Apparently it was sent<br />
to the <strong>Smithsonian</strong> on exchange early in the<br />
1900's.<br />
ETYMOLOGY.—The specific epithet is derived<br />
from a Latin name for the Cape Verde Islands,<br />
Doraces insulae.<br />
DISTRIBUTION.—Known only from the typelocality<br />
in the Cape Verde Islands.<br />
Nigeria: Sta 237, 101 m, 16\ 29 (L). Sta 239, 73 m, 86, 79<br />
(L). Sta 241, 59-63 m, mud and shell, 116\ 89 (5 ov) (W).<br />
Cameroon: Sta 259, 59 m, mud and broken shell, 36, 39<br />
(W). Sta 260, 46 m, Id (L).<br />
Geronimo Material: Gabon: Sta 235, 100 m, 16* (W).<br />
Undaunted Material: Angola: Sta 95, 126 m, 26, 19 ov (L).<br />
Other Material: Cameroon: 03°54'N, 08°53'E, 64 m, Aug<br />
1963, A. Crosnier, 56, 29 (1 ov) (W). 03°55.5'N, 08°52.5'E,<br />
60 m, Ombango, A. Crosnier, 5 Jan 1963, 19 ov (W).<br />
* Macropodia gilsoni (Capart, 1951)<br />
DESCRIPTION.—Capart, 1951:65.<br />
Figures: Capart, 1951, fig. 20, pi. 1: figs. 4, 10,<br />
pi. 2: fig. 3; Monod, 1956, figs. 811-822.<br />
Male Pleopod: Capart, 1951, pi. 2: fig. 3 (Cabinda);<br />
Monod, 1956, fig. 821 (Senegal).<br />
MEASUREMENTS.—Carapace lengths of males<br />
6.5 to 19 mm, of non-ovigerous females 7.5 to<br />
14 mm, of ovigerous females 11 to 16.7 mm.<br />
REMARKS.—Macropodia gilsoni, M. hesperiae, new<br />
species, M. intermedia, and M. longipes are the only<br />
<strong>West</strong> <strong>African</strong> species of the genus with strong,<br />
well-developed nuchal spines; these spines are<br />
also present in the northern species, M. tenuirostris<br />
(Leach) (Christiansen, 1969, fig. 47). Further, M.<br />
gilsoni and M. intermedia are the only species in the<br />
genus with spines on the supraorbital margin.<br />
Differences between these two species are pointed<br />
Achaeopsis gilsoni Capart, 1951:65, fig. 20, pi. 1: figs. 4, 10, pi. out (p. 300) under the account of M. intermedia.<br />
2: fig. 3.—Rossignol, 1957:115 [key].<br />
Macropodia gilsoni appears to be quite variable<br />
Macropodia gilsoni.—Monod, 1956:555, figs. 811-822.— in some features, especially in the extent of spi-<br />
Longhurst, 1958:89.—Rossignol, 1962:123.—Crosnier, nation: the supraorbital spines may be reduced<br />
1964:34.<br />
to tubercles; the walking legs often have but one<br />
Macropodia intermedia.—Guinot and Ribeiro, 1962:78.—Forest<br />
and Guinot, 1966:115.—Crosnier, 1970:1215 [listed], spine or even up to five spines on the merus of<br />
1218. [Not M. intermedia Bouvier, 1940.]<br />
the fourth leg rather than three; the hairiness of<br />
Macropodia.—Voss, 1966:22.<br />
the legs is variable; and in some specimens the<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 68, 70 m, broken shell, 4
298<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
bottoms, including bottoms with shells and bry- dorsal view. Protogastric region smooth, lacking<br />
ozoans or foliate Foraminifera.<br />
dorsal tubercles medially. Hepatic regions each<br />
Ovigerous females have been found in all with low, conical prominence dorsally. Hepatic<br />
months but July, indicating that the species lobes each with apical spinule, flanked mesially<br />
spawns all year.<br />
with spine on anterior margin in some specimens.<br />
DISTRIBUTION.—<strong>West</strong> Africa, from localities be- Gastric and cardiac regions each with low, conical<br />
tween Senegal to Angola (16°37'S); sublittoral, prominence. Branchial regions with poorly<br />
in depths between 37 and about 200 m. Capart marked prominences: 1 low, rounded epibran-<br />
(1951) studied numerous specimens from localichial, 1 broad dorsal mesobranchial, 1 low metaties<br />
between Gabon and Angola, and Monod branchial near posterior margin, and sharp tu-<br />
(1956) had material from Senegal and Guinea. bercles laterally. Anterolateral margins with<br />
Records since 1956 include the following: sharp, anteroventrally directed spine posterior to<br />
Senegal: 12°55.5'N, 17°33'W, 65-75 m (Forest and<br />
Guinot, 1966).<br />
Guinea-Bissau: 10°19'N, 16°34'W, 60-73 m (Forest and<br />
Guinot, 1966).<br />
hepatic lobe.<br />
Basal antennal segment (Figure lib) with 1-3<br />
spinules or sharp tubercles, anteriormost largest.<br />
Distal article of antennal peduncle about 3 times<br />
Sierra Leone: No specific locality, 54-200 m (Longhurst,<br />
1958).<br />
Ivory Coast: 04°35'N, 06°40'W to 04°35'N, 06°41'W, 64<br />
m (Voss, 1966).<br />
Cameroon: No specific locality, deeper than 50 m (Crosnier,<br />
1964).<br />
as long as penultimate. Epistome with spinule<br />
near opening of antennal gland, occasionally with<br />
spinule anteriorly near base of antenna.<br />
Ocular peduncles lacking distinct anterior lobe.<br />
Cornea with distinct dorsal tubercle.<br />
Gabon: W of Barre des Portugais; W of Mayumba, 110<br />
m (Rossignol, 1962).<br />
Angola: Luanda, 102-122 m (Guinot and Ribeiro, 1962).<br />
16°37'S, 11°22'E, 126 m (Crosnier, 1970).<br />
Chelipeds of adult males larger and more robust<br />
than in females. Fingers as long as palm,<br />
latter with row of spinules dorsally. Merus with<br />
3 erect dorsal spines or lobes.<br />
* Macropodia hesperiae, new species<br />
Dactylus of second pereiopod (Figure lie) slender,<br />
slightly curved distally, 7/9 as long as merus,<br />
FIGURE 77<br />
latter longer than carapace and antennal peduncles.<br />
Meri of second through fifth pereiopods each<br />
Macropodia linaresi.—Forest and Guinot, 1966:117 [not Macrowith<br />
single large distal dorsal spine, latter lower<br />
podia linaresi Forest and Zariquiey Alvarez, 1964].<br />
Macropodia.—Voss, 1966:31.<br />
on merus of fifth pereiopod. Dactyli of fourth and<br />
fifth (Figure lid) pereiopods strongly arched,<br />
MATERIAL EXAMINED.—Pillsbury Material: Ivory<br />
Coast: Sta 60, 79-82 m, coral or rock, 16\ 1$ (L). Sta 65, 46ventral<br />
margin toothed along entire margin.<br />
MEASUREMENTS.—Carapace lengths of males<br />
49 m, 16* (holotype), 1? (L).<br />
Ghana: Sta 22, 51 m, rough bottom, 16* (W).<br />
Nigeria: Sta 230, 82-97 m, hard ground, with gorgonians,<br />
coral, rock, 1(5 (W).<br />
5.2 to 9.4 mm, of females 4.2 and 7.3 mm.<br />
REMARKS.—Macropodia hesperiae resembles M.<br />
linaresi Forest and Zariquiey Alvarez, 1964, but<br />
differs as follows: The rostrum is longer, with the<br />
DESCRIPTION.—Size small, carapace lengths of apices of the spines divergent rather than ap-<br />
adults less than 10 mm. Rostrum (Figure 77a,*) pressed; the dorsal prominences of the carapace<br />
short, overreaching base but falling short of mid- are much less prominent; the carapace is<br />
dle of fifth article of antennal peduncle, orna- smoother and less hairy; and the legs are longer,<br />
mented with curved hairs. Apices of rostral teeth<br />
divergent in some specimens. Rostrum slightly<br />
upturned dorsally (Figure Tib). Nuchal region<br />
usually with lateral spinule or tubercle, visible in<br />
the merus of the second pereiopod being longer<br />
than the carapace and antennal peduncles combined.<br />
If our interpretation of the original account of
NUMBER 306 299<br />
a<br />
FIGURE 77.—Macropodia hesperiae, new species. Male paratype, cl 6.0 mm, Pillsbwy Sta 60: a,<br />
carapace; b, front, lateral view; c, second pereiopod; d, fifth pereiopod. Female paratype, cl 4.2<br />
mm, Pillsbury Sta 60: e, carapace.<br />
M. linaresi is correct, the basal antennal segment<br />
in that species lacks distinct spines; it was described<br />
as "avec des granules ou des courtes spinules"<br />
(Forest and Zariquiey Alvarez, 1964:241).<br />
Also, in that species, the merus of the second<br />
pereiopod was described as being slightly longer<br />
than the carapace. We have examined a male cl<br />
7 mm, and a female, cl 7 mm (RMNH Crust. D.<br />
20133), from Cadaques, Spain, and found that<br />
both specimens had three spinules ventrally on<br />
the basal segment of the antenna, the anteriormost<br />
largest, and, in the female, the merus of the<br />
second pereiopod is longer than the carapace; all<br />
the pereiopods of the male were detached.<br />
We have no hesitation in identifying material<br />
identified with M. linaresi from Senegal by Forest<br />
and Guinot (1966:117) with M. hesperiae as they<br />
noted that "elle ne differe guere des exemplaires<br />
typiques .. . que par un rostre legerement plus<br />
grele et une carapace plus lisse et plus glabre."<br />
Macropodia hesperiae is a much smoother species<br />
than M. linaresi.<br />
TYPE-LOCALITY.—Off the Ivory Coast,<br />
04°15'N, 07°32 / W to 04°12'N, 07°35.5'W, 46-49<br />
m, Pillsbury Sta 65.<br />
DISPOSITION OF TYPES.—The holotype (Crust.<br />
D. 23891), a male from Pillsbury Sta 65, is in the<br />
Rijksmuseum van Natuurlijke Historie, Leiden;<br />
paratypes also are at Leiden and in the National<br />
Museum of Natural History, <strong>Smithsonian</strong> <strong>Institution</strong>,<br />
Washington, D.C.<br />
ETYMOLOGY.—The specific name is derived<br />
from the Latin name for the Gulf of Guinea,<br />
Hesperium mare.<br />
BIOLOGY.—Macropodia hesperiae is a sublittoral<br />
species known from depths of 46-49 m, 51 m, 79-<br />
82 m, and 82-97 m (Pillsbury) and 65-75 m<br />
(Calypso; Forest and Guinot, 1966). Our specimens
300 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
were taken on rough bottom at three or four<br />
stations. The Calypso specimens were taken on<br />
mud, sand, and compact sand (sable construit).<br />
An ovigerous female was taken in May by the<br />
Calypso (Forest and Guinot, 1966).<br />
DISTRIBUTION.—Gulf of Guinea, from off Senegal,<br />
the Ivory Coast, Ghana, and Nigeria, in<br />
depths between 46-49 and 82-97 m. Records in<br />
the literature include the following:<br />
Senegal: 12°55.5'N, 17°33'W, 65-75 m (Forest and<br />
Guinot, 1966).<br />
Nigeria: 06° 1 l'N, 03°36'E to 06° 10'N, 03°38'E, 82-97 m<br />
(Voss, 1966).<br />
Macropodia intermedia Bouvier, 1940<br />
Macropodia longirostris var. intermedia Bouvier, 1940:366.—Capart,<br />
1951:76.<br />
?Macropodia gilsoni.—Maurin, 1968a:48, 62; 1968b:484, 486,<br />
489, fig. 5 [not Macropodia gilsoni (Capart, 1951)].<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Morocco: No specific locality, holotype of<br />
Macropodia longirostris var. intermedia, 16* (MP).<br />
Remarks.—Although several authors have synonymized<br />
Macropodia gilsoni Capart, 1951, with<br />
Macropodia intermedia Bouvier, 1940 (p. 297), we<br />
believe that two taxa may be involved, M. intermedia<br />
from areas to the north of the tropical<br />
region and M. gilsoni from the Gulf of Guinea<br />
proper, and, until more material from the two<br />
regions can be studied together, we prefer to<br />
recognize two species.<br />
The holotype of M. intermedia, a male, cl 19<br />
mm, cb 13.5 mm, was examined by Manning at<br />
the Museum national d'Histoire naturelle in Paris<br />
in 1971. Several differences between it and our<br />
material of M. gilsoni were observed: (1) The<br />
rostrum is longer than the antennal peduncle; it<br />
is shorter in M. gilsoni (Monod, 1956, fig. 814).<br />
(2) The posterior margins of the antennular fossae<br />
as well as those portions that overlap the basal<br />
segment of the antenna are smooth; they usually<br />
are spinulose in M. gilsoni (Monod, 1956, figs.<br />
814, 815). (3) The ventral spine on the ischium of<br />
the fifth pereiopod appears to be much less prom-<br />
inent and appears to be directed laterally rather<br />
than posteriorly as in M. gilsoni.<br />
Although these differences are relatively minor,<br />
they suggest to us that M. intermedia should be<br />
considered distinct from M. gilsoni, at least for the<br />
time being.<br />
DISTRIBUTION.—Eastern Atlantic, from the NW<br />
coast of Africa, sublittoral. All but the first of the<br />
following records require verification:<br />
Morocco: Atlantic coast (Bouvier, 1940).<br />
Spanish Sahara: Pulpito Bay, 20-30 m (Capart, 1951).<br />
Between Cabo Corbeiro and Cabo Blanco, 60-80 m<br />
(Maurin, 1968a). Between Cabo Barbas and Cabo Blanco,<br />
50-90 m (Maurin, 1968b).<br />
Mauritania: S Bane d'Arguin, 90 150 m (Maurin,<br />
1968a). Bane d'Arguin, 40-60, 60-70, 90-100 m (Maurin,<br />
1968b).<br />
Macropodia longicornis (A. Milne Edwards<br />
and Bouvier, 1899)<br />
Macropodia longicornis.—Monod, 1956:562 [references].<br />
DISTRIBUTION.—Cape Verde Islands; sublittoral,<br />
to at least 275 m.<br />
Macropodia longipes (A. Milne Edwards and<br />
Bouvier, 1899)<br />
Stenorhynchus longipes A. Milne Edwards and Bouvier, 1899:<br />
48.<br />
IMocropodia longirostris.—Monod, 1956:559 [references].—<br />
Rossignol, 1957:115 [key].—Peres, 1964:20 [Morocco].<br />
[Not Cancer longirostris Fabricius, 1775.]<br />
Macropodia longipes.—Forest and Games, 1960:357 [Morocco].—Forest<br />
and Zariquiey Alvarez, 1964:226, 241<br />
[key], figs. 2, 6, 13 [review of Mediterranean species].—<br />
Forest, 1965b:349, 350 [discussion].—Bas, Arias, and<br />
Guerra, 1976, table 3 [Spanish Sahara].—Tiirkay, 1976a:<br />
25 [listed], 40, fig. 33 [Portugal, Morocco].<br />
Macropodia tenuirostris longipes.—Forest, 1978:337, figs. 9, 16.<br />
REMARKS.—Forest (1978:337) considers M. longipes<br />
to be the southern subspecies of M. tenuirostris<br />
(Leach, 1814).<br />
DISTRIBUTION.—Eastern Atlantic, from the<br />
Gulf of Gascogne to Mauritania, Cape Verde<br />
Islands, and western Mediterranean; sublittoral,<br />
in 50-445 m (Forest, 1978).
NUMBER 306 301<br />
*Macropodia macrocheles (A. Milne Edwards<br />
and Bouvier, 1898)<br />
Macropodia macrocheles.—Capart, 1951:77, fig. 23.—Monod,<br />
1956:560, 632, figs. 823-827.—Rossignol, 1957:115<br />
[key].—Longhurst, 1958:89.—Guinot and Ribeiro, 1962:<br />
78.—Forest and Guinot, 1966:116.<br />
Macropodia.—Voss, 1966:27.<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 82, 146-150 m, 1(5, 19 ov (L).<br />
Nigeria: Sta 255, 264-269 m, 36, 2$ (1 ov) (L, W).<br />
Geronimo Material: Gabon: Sta 197, 200 m, 1$ (W).<br />
Other Material: Mauritania: Off Cabo Blanco, 240 m,<br />
Talisman, 13 Jul 1883, syntype, 1$ ov (W, USNM 22979).<br />
DESCRIPTION.—Capart, 1951:77.<br />
Figures: Capart, 1951, fig. 23; Monod, 1956,<br />
figs. 823-827.<br />
Male Pleopod: Monod, 1956, fig. 827 (Senegal).<br />
MEASUREMENTS.—Carapace lengths of males<br />
12.3 to 21 mm, of non-ovigerous females 11.8 to<br />
15.2 mm, of ovigerous females 8.0 to 13.8 mm.<br />
BIOLOGY.—Macropodia macrocheles is a deep water<br />
species, occurring between 96 and 300 m on<br />
soft bottoms (mud and sand, muddy sand), generally<br />
in depths greater than 150 m.<br />
Ovigerous females have been taken in January,<br />
February, May, June, and December (Capart,<br />
1951; Monod, 1956; Pillsbury).<br />
DISTRIBUTION.—<strong>West</strong> Africa, from localities between<br />
Mauritania and Angola, in depths between<br />
96 and 300 m. Monod (1956) reported the species<br />
from Senegal and Sierra Leone; since 1956 it has<br />
been taken from the following localities:<br />
Guinea-Bissau: 10°32'N, 16°53.5'W, 174 m (Forest and<br />
Guinot, 1966).<br />
Sierra Leone: 08°45'N, 14°38'W, 220-240 m (Longhurst,<br />
1958).<br />
Liberia: 04°57'N, 09°30'W to 04°58'N, 09°32'W, 146-<br />
150 m (Voss, 1966).<br />
Angola: Baia dos Tigres, 150-200 m (Guinot and Ribeiro,<br />
1962).<br />
* Macropodia spinulosa (Miers, 1881)<br />
FIGURE 78a<br />
Stenorhynchus rostratus var. spinulosus Miers, 1881a:206.<br />
Stenorhynchusphalangium.—Studer, 1882:335; 1883:7 [not Cancer<br />
phalangium Fabricius, 1775].<br />
Macropodia rostrata spinulosa.—Rathbun, 1900a:293 [listed].<br />
Macropodia rostrata.—Rathbun, 1900a:293 [listed].—Balss,<br />
1922:71.— Odhner, 1923:18.—Monod, 1933b:501<br />
[listed].—Capart, 1951:74, pi. 1: fig. 3.—Sourie, 1954b:<br />
147.—Monod, 1956:562, figs. 828-836.—Rossignol, 1957:<br />
115 [key].—Longhurst, 1958:89.—Gauld, 1960:72.—<br />
Guinot and Ribeiro, 1962:78, pi. 4: figs. 2, 3.—Forest and<br />
Guinot, 1966:116.—Le Loeuff and Intes, 1968:46, table<br />
1.—Crosnier, 1970:1218. [Not Macropodia rostrata (Linnaeus,<br />
1761).]<br />
Stenorhynchus rostratus.—Doflein, 1904:69.—Lenz and<br />
Strunck, 1914:272 [part, not South <strong>African</strong> specimens?].<br />
[Not Macropodia rostrata (Linnaeus, 1761).]<br />
?Stenorhynchus phalangium.—Stimpson, 1907:22 [Madeira]<br />
[not Cancer phalangium Fabricius, 1775].<br />
MATERIAL EXAMINED.—Pillsbury Material: Ivory<br />
Coast: Sta 46, 38-42 m, mud with dense Jullienella, 56, 10$<br />
(7 ov) (W). Sta 47, 37 m, bottom with Jullienella, 16 (L). Sta<br />
48, 22 m, Id, 3$ ov (L). Sta 65, 46-49 m, 16* (L).<br />
Ghana: Sta 16, 46 m, mud with Foraminifera, shells, ld\<br />
1$ (W). Sta 17, 48 m, fine sand and green mud, 16 (L). Sta<br />
23, 42 m, foliate brown to orange bryozoans, 26, 2$ ov (L).<br />
Sta 24, 35-37 m, dark red bryozoans, 26, 19 ov (W). Sta 28,<br />
49-53 m, 10
302<br />
a<br />
FIGURE 78.—a, Macropodia spinulosa (Miers) (from Monod,<br />
1956, fig. 828); b, Stenorhynchus lanceolatus (Brulle) (from<br />
Monod, 1956, fig. 838).<br />
rostrata (a character mentioned by Forest and<br />
Zariquiey Alvarez, 1964:241, key); only the proximal<br />
third or fourth of the dactylus of the fifth<br />
pereiopod is armed with spinules or low tubercles<br />
whereas in M. rostrata the proximal two-thirds to<br />
three-fourths is armed (most of the dactylus of<br />
that leg is naked in M. spinulosa, lacking even<br />
setae); and the dorsal spines of the carapace are<br />
much longer (Monod, 1956, fig. 830). In addition<br />
the pereiopods are longer, with the third leg<br />
almost five times rather than three times as long<br />
as the carapace (Christiansen, 1969:112) and, as<br />
noted by Miers (188la:206) the chelipeds are very<br />
spinulose. Some of these differences already had<br />
been noted by Guinot and Ribeiro (1962:79) and<br />
by Crosnier (1970:1218).<br />
We have compared our material of M. spinulosa<br />
with M. rostrata from the North Sea, as well as<br />
from the Mediterranean, and we found some<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
differences in material of M. rostrata from those<br />
two areas. The Mediterranean form is more slender<br />
(compare carapace shape in Forest and Zariquiey<br />
Alvarez, 1964, fig. 1, and Christiansen,<br />
1969, fig. 46), a feature reflected in the shape of<br />
the epistome, which in Mediterranean specimens<br />
is as long as wide but in northern specimens is<br />
much broader than long. Also, the carapace in<br />
Mediterranean specimens is much smoother and<br />
less hairy. Finally, in Mediterranean specimens<br />
the dactylus of the fifth leg is longer, more slender,<br />
less curved, less setose, and has low denticles only<br />
on the proximal third; in northern specimens the<br />
dactylus is short, more strongly curved, much<br />
more setose, and has erect spinules on the proximal<br />
two-thirds. These differences strongly suggest<br />
that two species may be involved. If they prove<br />
to be constant and the Mediterranean form is a<br />
distinct species, the name Macropodia inermis (Heller,<br />
1856) is available. It was named originally as<br />
Stenorhynchus inermis from the Adriatic Sea (Heller,<br />
1856:3).<br />
Material from South Africa identified with M.<br />
rostrata by Lenz and Strunck (1914) and Barnard<br />
(1950, 1954) resembles M. spinulosa rather than<br />
M. rostrata in having a short rostrum, with the<br />
margins of the antennular fossae visible at the<br />
base of the rostrum. Barnard's material, at least,<br />
differs from both of these more northern species<br />
in having distal spinules on the basal segment of<br />
the antenna, an apical seta on the eye, and in<br />
having spinules along the entire margin of the<br />
dactylus of the fifth pereiopod.<br />
BIOLOGY.—Macropodia spinulosa is a sublittoral<br />
species, having been taken in depths between ca<br />
1 and 126 m; of 55 depth records available, only<br />
3 (5%) are from depths of more than 100 m (108,<br />
110, and 126 m), 11 (20%) are from between 50<br />
and 100 m, 11 (20%) are from depths of 20 m or<br />
less, and the remainder (55%) are from depths<br />
between 20 and 50 m. The deeper material identified<br />
here with this species probably should be<br />
restudied.<br />
The species generally occurs on soft bottoms,<br />
mud or muddy sand, but it was taken by the<br />
Calypso (Forest and Guinot, 1966) on mud, rocks,
NUMBER 306 303<br />
calcareous algae, sand, and Foraminifera; mud,<br />
sand, and compacted sand (sable construit); mud;<br />
mud with Area; and on calcareous algae. Our<br />
specimens were taken on mud, mud with Foraminifera,<br />
fine sand and green mud, and on bottom<br />
with bryozoans. Sourie (1954b) found it on<br />
coarse, shelly sand, bottom with Area and Pyura,<br />
in 10-12 m in the Baie de Dakar. Longhurst<br />
(1958) found it on shelly mud in 60 m, and Le<br />
Loeuff and Intes (1968) considered it a eurythermic<br />
species, living on muddy sand in 25-40<br />
m.<br />
Ovigerous females have been collected from<br />
January through June, and August (Capart,<br />
1951; Monod, 1956; Guinot and Ribeiro, 1962;<br />
Forest and Guinot, 1966; Crosnier, 1970; herein<br />
p. 301).<br />
DISTRIBUTION.—<strong>West</strong> Africa, from the Cape<br />
Verde Islands and at least Senegal on the <strong>African</strong><br />
mainland S to Angola; sublittoral, in depths between<br />
1 and 126 m, generally in less than 50 m.<br />
Records of M. rostrata from areas N of Senegal,<br />
such as those by Capart (1951) from Spanish<br />
Sahara and Stimpson (1907) from Madeira, especially<br />
the latter, may be based on either this<br />
species or M. rostrata proper. They require verification.<br />
Records in the literature include the following:<br />
Cape Verde Islands: 15°40'N, 23°06'W) 38 fm [70 m]<br />
(Studer, 1882, 1883).<br />
Senegal: Off Dakar, 14°40'-41'N, 17°18'30"-20'30"W,<br />
20-25 m; 14°38'-41'N, 17°2O'-23'W, 22-34 m; 14°38'-<br />
40'N, ^H'-iaW, 22-27 m; off Dakar; Anse Bernard, 8-<br />
10 m; Baie de Hann; between Goree and Thiaroye-sur-Mer<br />
(as Tiaroye), 15 m; and between Dakar and Goree, 16 m (all<br />
Monod, 1956). Baie de Dakar, 10-12 m (Sourie, 1954b).<br />
Goree (the type-locality) (Miers, 1881a; Lenz and Strunck,<br />
1914; Monod, 1956); in 20 m (Balss, 1922). Around Goree,<br />
ca. 1, 5-11, 7-8, 13, 15-20, 19, 20, 25, 40-41, and 46-50 m;<br />
between Thiaroye-sur-Mer and Bel-Air, 9 m; Mbao; SE lie<br />
de la Madeleine, 35 and 48 m; and off Joal, 15-17 m<br />
(Monod, 1956). 13°01'N, 17°24'W, 51-55 m, and 12°55.5'N,<br />
17°33'W, 65-75 m (Forest and Guinot, 1966).<br />
Guinea: Off lie Kassa, lies de Los (Capart, 1951). 09°-<br />
36'N, 13°57'W, 18-30 m (Forest and Guinot, 1966).<br />
Sierra Leone: No specific locality, 60 m (Longhurst, 1958).<br />
Ivory Coast: Off Grand Bassam, 25-40 m (Le Loeuff and<br />
Intes, 1968).<br />
Ghana: Off Accra, 26 and 44 m (Monod, 1956; Gauld,<br />
1960).<br />
Nigeria: Off the mouths of the Niger River, 04°03'N,<br />
06°12'E, 32 m (Forest and Guinot, 1966).<br />
Principe: 01°43'10"N, O7°28'2O"E, 73 m, and 01°43'N,<br />
07°28'55'E, 37m (Forest and Guinot, 1966).<br />
Annobon: 2 mi [3.2 km] off Annobon, 18-20 m (Capart,<br />
1951).<br />
Gabon: Libreville, 60 m (Monod, 1956).<br />
Congo: 11 mi [17.7 km] WSW Pointe-Noire, 04°52'S,<br />
ll°39'30"E, 58-60 m (Capart, 1951).<br />
Cabinda: Off the lighthouse (Capart, 1951).<br />
Zaire: Off the mouth of the Congo River, 44 m (Doflein,<br />
1904). 28 mi [32 km] WNW of Banana, 05°54'S, 11°58'-<br />
30"E, 50 m, and 05°56'S, 12°E, 50-60 m (Capart, 1951).<br />
Angola: Benguela, 13-26 m; Baia da Caota, 13, 16, and<br />
18 m; and Baia Farta, 5 m (Guinot and Ribeiro, 1962).<br />
Porto Alexandre, 72 and 108 m (Odhner, 1923). 18 mi<br />
[29 km] WSW Baia dos Tigres, 16°36'S, 11°27'E, 110 m<br />
(Capart, 1951). 16°37'S, 11°22'E, 126 m (Crosnier, 1970).<br />
* Macropodia straeleni Capart, 1951<br />
Macropodia straeleni Capart, 1951:79, fig. 24, pi. 1: fig. 5.—<br />
Monod, 1956:566, fig. 837.—Rossignol, 1957:115<br />
[key].—Forest and Guinot, 1966:117.<br />
MATERIAL EXAMINED.—Pilbbury Material: Liberia:<br />
Sta 68, 70 m, broken shell, 2c? (L).<br />
Ivory Coast: Sta 42, 62-75 m, mud with brown, branched<br />
Foraminifera, 26* (L). Sta 60, 79-82 m, coral or rock, 26, 29<br />
ov (W). Sta 65, 46-49 m, 1$ ov (L).<br />
Nigeria: Sta 239, 73 m, 1$ ov (W).<br />
Other Material: Congo: 05°20'S, 11°37'E, 200 m, 10 Dec<br />
1963, A. Crosnier, 36, 1? ov (W).<br />
DESCRIPTION.—Capart, 1951:79.<br />
Figures: Capart, 1951, fig. 24, pi. 1: fig. 5.<br />
MEASUREMENTS.—Carapace lengths of males<br />
5.1 to 8.7 mm, of ovigerous females 5.2 to 6.5<br />
mm.<br />
BIOLOGY.—Macropodia straeleni occurs sublittorally<br />
in moderately deep water. Capart (1951)<br />
took it in 73 to 140-150 m on sandy mud and it<br />
was taken by the Calypso in 65-75 m on mud,<br />
sand, and compacted sand (sable construit) (Forest<br />
and Guinot, 1966). The Pillsbury specimens,<br />
for which data are available, were taken on mud
304<br />
with brown, branched Foraminifera in 62-75 m,<br />
broken shell in 70 m, and on coral or rock in 79-<br />
82 m. The deepest recorded occurrence of the<br />
species is in 200 m off the Congo and the Ivory<br />
Coast (Forest and Guinot, 1966).<br />
Ovigerous females have been taken in May,<br />
June, October, November, and December (Capart,<br />
1951; Forest and Guinot, 1966; present paper).<br />
DISTRIBUTION.—<strong>West</strong> Africa, from localities between<br />
Senegal (12°55.5'N) and Angola (07°39'S),<br />
in depths between 46-49 m and 200 m. Capart<br />
(1951) based his account on material from off the<br />
Congo, Cabinda, and Angola; Monod (1956)<br />
added no material. Other records in the literature<br />
include the following:<br />
Senegal: 12°55.5'N, 17°33'W, 65-75 m (Forest and<br />
Guinot, 1966).<br />
Ivory Coast: No specific locality, 200 m (Forest and<br />
Guinot, 1966).<br />
Genus Stenorhynchus Lamarck, 1818<br />
Pactolus Leach, 1815b: 19 [type-species: Pactolus boscii Leach,<br />
1815, a subjective junior synonym of Cancer seticomis<br />
Herbst, 1788, by monotypy; gender: masculine; suppressed<br />
by the International Commission on Zoological<br />
Nomenclature under its plenary powers (Opinion 763,<br />
1966); name 1778 on Official Index].<br />
Stenotynckus Lamarck, 1818:236 [invalid original spelling of<br />
Stenorhynchus; name 1779 on Official Index].<br />
Stenorhynchus Lamarck, 1818:236 [emendation for Stenorynchus<br />
by the International Commission of Zoological Nomenclature<br />
under its plenary powers (Opinion 763, 1966); typespecies:<br />
Cancer seticomis Herbst, 1788, by designation by<br />
the International Commission on Zoological Nomenclature<br />
under its plenary powers (Opinion 763, 1966:19);<br />
gender: masculine; name 1700 on Official List].<br />
* Stenorhynchus lanceolatus (Brulle, 1837)<br />
FIGURE 784<br />
Leptopodia lanceolata Brulle, 1837, unnumbered pi.: fig. la,b.<br />
Leptopodia Canariensis Brulle, 1839:15.<br />
Leptopodia sagittaria.—Bru\le, 1839:15.—White, 1847a: 1<br />
[part].—Stimpson, 1858d:219.—Kingsley, 1880b:383<br />
[part].—Miers, 1886:3, 4.—Osorio, 1887:221; 1888:187,<br />
191.—Koelbel, 1892:114.—Osorio, 1898:187, 192.—A.<br />
Milne Edwards and Bouvier, 1900:153—Stimpson, 1907:<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
23.—Balss, 1922:72. [Not Cancer Sagittarius Fabricius, 1793<br />
= Cancer seticomis Herbst, 1788.]<br />
"une nouvelle espece de ce genre" [Leptopodia].—Guerin-<br />
Meneville, 1844:10.<br />
Leptopodia Sagittarius.—Herklots, 1861:136 [part].<br />
Leptopodia vittata (Guerin MS).—Kingsley, 1880b:384.<br />
Leptopodia sagitaria.—Osorio, 1889:130; 1898:185.<br />
Stenorynchus Sagittarius.—Rathbun, 1900a:293.—Stimpson,<br />
1907:23.<br />
Stenorhynchus Sagittarius.—Odhner, 1923:19.<br />
Stenorynchus seticomis.—Rathbun, 1925:13 [part, not pi. 2, 3].<br />
[Not Stenorhynchus seticomis (Herbst, 1788).]<br />
Stenorhynchus seticomis.—Monod, 1933b:503.—Capart, 1951:<br />
81, fig. 25.—Sourie, 1954b: 147.—Monod, 1956:567, figs.<br />
838, 839.—Rossignol, 1957:78, 115 [key].—Longhurst,<br />
1958:89.—Gauld, 1960:72.—Guinot and Ribeiro, 1962:<br />
79.—Rossignol, 1962:123.—Ribeiro, 1964:21.—Crosnier,<br />
1964:38, fig. on pi. A.—Forest and Guinot, 1966:117.—<br />
Maurin, 1968b:484, 486.—Le Loeuff and Intes, 1968,<br />
table 1; 1969:65.—Uschakov, 1970:455 [listed].<br />
Stenorynchus.—Voss, 1966:17, 19.<br />
Stenorhynchus lanceolatus.—Yang, 1967:220.—Barr, 1975:47<br />
[discussion].<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 70, 33 m, branched Foraminifera, 1$ (L).<br />
Ivory Coast: Sta 42, 62-75 m, mud with brown, branched<br />
Foraminifera, 1
NUMBER 306 305<br />
Stenorhynchus (S. seticornis, S. lanceolatus, and Stenorhynchus<br />
species A), gave the differences between<br />
the latter two species in the following tabular<br />
form:<br />
Stenorhynchus species A<br />
Propodite of female chela<br />
smooth<br />
Hiatus of female chela<br />
narrow<br />
Meropodite of male chela<br />
strongly granulated<br />
S. lanceolatus<br />
Propodite of female chela<br />
covered by fine spines<br />
Hiatus of female chela<br />
round; similar to S.<br />
seticomis<br />
Meropodite of male chela<br />
smooth<br />
The differences between S. seticornis (Herbst)<br />
and S. lanceolatus were discussed by Yang (1967:<br />
221) as follows:<br />
S. seticomis is also different from S. lanceolatus in having<br />
three spines at the distal margin of the meropodite (rather<br />
than two). Also S. seticomis is distinguished from 5. lanceolatus<br />
by having smooth propodite of the female chelae, and<br />
strongly granulated meropodite of the male chelae. The tip<br />
of the male first pleopod of S. lanceolatus is round without a<br />
tooth-like projection and resembles that of S. species A<br />
(based on one large specimen). However, the hiatus of the<br />
female chela and the spination of pereiopod carpodite of<br />
Stenorhynchus lanceolatus is very similar to that of 5. seticomis.<br />
The future larval study of S. lanceolatus will show the affinity<br />
of this species to Stenorhynchus sp. A or 5. seticomis.<br />
Figures: Capart, 1951, fig. 25; Monod, 1956,<br />
figs. 838, 839.<br />
Color: Koelbel (1892:115) mentioned "tief<br />
purpurbraunen, manchmal auch lauchgriinen<br />
Riickenstreifen" and noted in a footnote<br />
"Vermoge eines ausgepragt griinlichen<br />
Schimmers entspricht die Farbung derartiger<br />
Streifen vollstandig jener der Fliigeldecken der<br />
Mannchen von Dyticus marginalise He described<br />
the background of the stripes as "Elfenbeinweiss."<br />
A. Milne Edwards and Bouvier (1900:<br />
154) described the color of the species as follows:<br />
Les couleurs de ce Crabe sont disposees d'une facon tres<br />
remarquable. Une serie de bandes brunes, jaunatres ou<br />
presque blanches, s'etendent longitudinalement; elles se reunissent<br />
en avant sur la ligne mediane, puis se dirigent<br />
en arriere en suivant les contours de la carapace, les plus<br />
longues etant en dehors, les plus courtes au centre et la<br />
region cardiaque pouvant etre consideree comme le centre<br />
de ces lignes concentriques. Les pattes sont brunes, relevees<br />
de taches et de bandes longitudinales d'un jaune vif. Les<br />
doigts des pinces sont d'un bleu violace.<br />
Capart (1951:82) described the color as follows:<br />
"La carapace de couleur orange, avec bande<br />
longitudinale violette, les pattes et pinces marquees<br />
de bandes de memes couleurs."<br />
Male Pleopod: Monod, 1956, fig. 839 (Sierra<br />
Leone).<br />
MEASUREMENTS.—The carapace lengths of the<br />
present male specimens ranged from 18 to 24<br />
mm, in the females from 19 to 28 mm (the same<br />
range was shown by the ovigerous females). In<br />
the literature males have been reported with cl<br />
18.2-54.5 mm (Guinot and Ribeiro, 1962), 62.5<br />
mm (Miers, 1886), 71 and 75 mm (Rossignol,<br />
1957) and 72 mm (Capart, 1951); A. Milne Edwards<br />
and Bouvier (1900) gave the span of their<br />
males as 200 to 250 mm. Non-ovigerous females<br />
with cl 14.4-49.5 mm and ovigerous females with<br />
cl 36.9, 40.2, and 44.2 mm were listed by Guinot<br />
and Ribeiro (1962). Monod (1956) gave the diameter<br />
of the eggs as 0.6 to 0.7 mm.<br />
REMARKS.—Brulle (1837) gave a figure of the<br />
present species and on the plate used the name<br />
Leptopodia lanceolata for it. Later the same author<br />
(Brulle, 1839), now using the name Leptopodia<br />
canariensis sank it as a synonym of Leptopodia sagittaria<br />
(Fabricius, 1793) [= Stenorhynchus seticomis<br />
(Herbst, 1788)]. Practically all later authors accepted<br />
this synonymy. Only recently Yang (1967)<br />
showed that not one but three distinct species of<br />
the present genus have to be distinguished in the<br />
Atlantic region, one <strong>West</strong> <strong>African</strong> and two western<br />
Atlantic. Yang showed that the correct name<br />
for the <strong>West</strong> <strong>African</strong> species is Stenorhynchus lanceolatus<br />
(Brulle, 1837) and he indicated the differences<br />
between the three species. He based the<br />
<strong>West</strong> <strong>African</strong> form largely on part of the Pillsbury<br />
material listed above. Dr. Won Tack Yang intends<br />
to publish a more extensive paper on this<br />
problem, for which reason we only deal briefly<br />
with it here.<br />
BIOLOGY.—The species has been collected at<br />
depths between 6 and 96 m (extremes 2-6 m, 5-<br />
7 m, 80 m, 75-90 m, and 96 m); more than 85%<br />
of the catches were from between 20 and 80 m.
306<br />
The bottom on which the species was found was<br />
variously reported: sand and rock; sand and shells<br />
(A. Milne Edwards and Bouvier, 1900). Mud;<br />
green mud; brown and green mud; sand, mud,<br />
and rock (Capart, 1951). Coarse shelly sand, bottom<br />
with Area and Pyura (Sourie, 1954b). Mud<br />
and sand; shell and sand (Longhurst, 1958).<br />
Stones; rock (Guinot and Ribeiro, 1962). Mud;<br />
mud and sand; mud and shells; sand and rock;<br />
sand, rock, coral, and calcareous algae; calcareous<br />
algae; calcareous and other algae; sand, calcareous<br />
and other algae; rock (Forest and Guinot,<br />
1966).<br />
Ovigerous females have been reported from all<br />
months except February (Capart, 1951; Monod,<br />
1956; Guinot and Ribeiro, 1962; Forest and<br />
Guinot, 1966; present paper).<br />
Monod (1956) reported upon a specimen carrying<br />
a branchial isopod parasite.<br />
DISTRIBUTION.—Stenorhynchus lanceolatus is an<br />
eastern Atlantic species, known from Madeira,<br />
the Canary Islands, the Cape Verde Islands, and<br />
from numerous localities on the <strong>African</strong> mainland<br />
between Spanish Sahara and Angola. The following<br />
records are in the literature:<br />
Madeira: No specific locality (White, 1847a; Miens, 1886).<br />
Off the S side of Madeira 15 fm [27 m] (Stimpson, 1858d,<br />
1907).<br />
Canary Islands: E coast of Isla de Gran Canaria and near<br />
the harbor of Puerto de Cabras, Isla de Fuertaventura<br />
(Koelbel, 1892). Near Isla de Tenerife, 28°28'N, 18°32'W<br />
[of Paris, 16°12'W of Greenwich], 80 m, and Estrecho de la<br />
Bocaina, 28°49'N, 16°13'W [of Paris, 13°53'W of Greenwich],<br />
30 m (A. Milne Edwards and Bouvier, 1900).<br />
Cape Verde Islands: No specific locality (Osorio, 1898).<br />
Sao Vicente (Miers, 1886; Osorio, 1888). Channel between<br />
Santo Antao and Sao Vicente, 75-90 m (A. Milne Edwards<br />
and Bouvier, 1900). Faja di Agua, Brava, 8 and 50 m<br />
(Guinot and Ribeiro, 1962; Ribeiro, 1964).<br />
Spanish Sahara: 21°05'N, 17°14'W, 43-45 m (Forest and<br />
Guinot, 1966). Between Cabo Barbas and Cabo Blanco and<br />
between Cabo Corbeiro and Cabo Blanco, 200 m (Maurin,<br />
1968b).<br />
Mauritania: Bane d'Arguin, 50 m (Maurin, 1968b).<br />
Senegal: No specific locality (Herklots, 1861; Kingsley,<br />
1880b). S of He de la Madeleine, 40 m; near Dakar,<br />
14°41'30"-41"N, 17°18'30"-20'30"W,20-25m; 14°39'30"-<br />
4C30"N, 17°16'-18°30'W, 18-19 m; 14°38'-39'N, 17° 16.5'n.S'W,<br />
26 m; between Goree and Dakar, 16 m; area around<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Goree, 33-35 and 41 m; Bane du Seminole, Baie de Goree,<br />
ca. 38 m; between Cap Manuel and S point of Goree, 24-27<br />
m; and off Ngaparou (all Monod, 1956). Baie de Dakar, 10-<br />
12 m (Sourie, 1954b). Rufisque, 14°30'N, 17°25'W, 24 m<br />
(Capart, 1951). 12°55.5'N, 17°33'W, 65-75 m (Forest and<br />
Guinot, 1966).<br />
Guinea-Bissau: 10°19'N, 16°34'W, 60-73 m (Forest and<br />
Guinot, 1966).<br />
Guinea: No specific locality (Uschakov, 1970). 09°44'N,<br />
13°56'W, 10 m; 09°40'N, 14°21'W, 25 m; 09°16'N,<br />
13°42'W, 20 m; 09°16'N, 13°34'W, 10 m; 09°07'N,<br />
13°41'W, 25 m; and 09°N, 13°50'W, 30 m (Monod, 1956).<br />
09°36'N, 13°57'W, 18-30 m (Forest and Guinot, 1966). Off<br />
Conakry, 10 m; Matakong, ca. 15 m (Monod, 1956). Between<br />
lie Tamara and lie Roume, lies de Los, 10-12 m<br />
(Capart, 1951).<br />
Sierra Leone: No specific locality, in 25 m (Longhurst,<br />
1958). 08°38'-08°42'N, 8-12 m (Monod, 1956).<br />
Ivory Coast: 05°05'N, 04°59.5'W, 22 m (Voss, 1966;<br />
Yang, 1967). Lagoon at Abidjan, 05°16'N, 04°01'20"W<br />
(Forest and Guinot, 1966). Off Port-Bouet (Monod, 1956).<br />
Off Grand-Bassam, 15-60 m (Le Loeuff and Intes, 1968).<br />
Over all of the continental shelf (Le Loeuff and Intes, 1969).<br />
Ghana: 04°46'N, 02°30'W to 04°45'N, 02°33'W, 61-<br />
64 m (Voss, 1966; Yang, 1967). Off Takoradi (Gauld, 1960).<br />
Off Accra, 32-51 m (Monod, 1956; Gauld, 1960).<br />
Nigeria: No specific locality (Monod, 1956). Off the<br />
mouths of the Niger River, 04°03'N, 06°12'E, 32 m (Forest<br />
and Guinot, 1966). Off the Niger delta, 04°03'N, 05°41'E to<br />
04°07'N, 05°40'E, 33 m, and 04°04'N, 06°18'E to 04°05'N,<br />
06°22'E, 30 m (Yang, 1967).<br />
Cameroon: No specific locality, in 30-50 m (Crosnier,<br />
1964).<br />
Principe: 01°43'10"N, 07°28'20"E, 73 m; 01°43'N,<br />
07°28'55"E, 37 m; 01°38'35"N, 07°21'35"E, 35 m;<br />
01°35'N, 07°28'E, 45 m (Forest and Guinot, 1966).<br />
Sao Tome: No specific locality (Osorio, 1887, 1889,<br />
1898; Balss, 1922). Praia da Fernao Dias (Osorio, 1888).<br />
00°25'40"N, 06°40'10"E, 50 m; OO°25'15"N, 06°43'05"E,<br />
8-30 m; Morro Peixe, 2-6 m (Forest and Guinot, 1966).<br />
Annobon: No specific locality, 18-20 m (Capart, 1951).<br />
N of San Antonio, 23 m (Forest and Guinot, 1966).<br />
Congo: Off Pointe-Noire (Monod, 1956; Rossignol, 1957).<br />
W of Pointe-Noire, 40 m (Rossignol, 1962). 32 mi [51.5 km]<br />
W of Pointe-Noire, 04°48'S, 1 l°30'E, 50 m; 11 mi [17.7 km]<br />
WSW of Pointe-Noire, 04°52'S, 1 l°39'30"E, 58-60 m; 8 mi<br />
[12.9 km] WSW of Pointe-Noire, 04°53'S, 11°53'E, 50-70 m<br />
(Capart, 1951).<br />
Cabinda: 05°03'S, 11°24'E, 36-48 m (Capart, 1951).<br />
Angola: No specific locality (Miers, 1886; Guinot and<br />
Ribeiro, 1962). 11 mi [17.7 km] W of Cabo Ledo, Luanda,<br />
09°40'S, 13°02'E, 80 m (Capart, 1951). Benguela, 13-26 m;<br />
Baia Farta, 30 m; Baia da Caota, 13.5 and 30 m (Guinot
NUMBER 306 307<br />
and Ribeiro, 1962). Porto Alexandre, Mocamedes (Odhner,<br />
1923).<br />
Subfamily MAJINAE Samouelle, 1819<br />
Genus Maja Lamarck, 1801<br />
Maja Lamarck, 1801:154 [type-species: Cancer squinado<br />
Herbst, 1788, by subsequent designation under the plenary<br />
powers of the International Commission on Zoological<br />
Nomenclature in Opinion 511; gender: feminine; name<br />
1260 on Official List].<br />
Meria Griffith and Pidgeon, 1833:165 [erroneous spelling of<br />
Maja].<br />
Paramya de Haan, 1837, pi. 24 [type-species: Pisa {Paramya)<br />
spinigera de Haan, 1837, by monotypy; gender: feminine].<br />
Mamaia Stebbing, 1905:23 [substitute name for Maja; typespecies:<br />
Cancer squinado Herbst, 1788; gender: feminine].<br />
REMARKS.—Guinot and Ribeiro (1962:76, footnote),<br />
in their account of M. squinado, remarked<br />
on the status of <strong>West</strong> <strong>African</strong> species of Maja as<br />
follows: "Nous identifions provisoirement ces<br />
specimens a Maja squinado, mais la systematique<br />
des Maja ouest-africaines n'est pas entierement<br />
satisfaisante et fera l'objet d'une mise au point<br />
ulterieure." To our knowledge these authors have<br />
not pursued this problem, and, inasmuch as we<br />
lack material of Maja from tropical localities, we<br />
have not been able to investigate it ourselves.<br />
Those working with <strong>West</strong> <strong>African</strong> Maja in the<br />
future should take this into account.<br />
Maja crispata Risso, 1827<br />
Maia crispata Risso, 1827:23.<br />
Maja verrucosa.—Monod, 1956:477 [references].—Zariquiey<br />
Alvarez, 1968:447, figs. 149b, 150a-f, 156a [Spain; references].<br />
SYNONYMS.—Maia verrucosa H. Milne Edwards,<br />
1834 (see Holthuis, 1977b:72); Cancer majodes<br />
Nardo, 1847.<br />
DISTRIBUTION.—Eastern Atlantic, from Portugal<br />
to Cabo Blanco and the Cape Verde Islands,<br />
Mediterranean; shallow water, littoral and sublittoral.<br />
Maja goltziana d'Oliveira, 1888<br />
Maja goltziana.—Capart, 1951:100, fig. 33 [Gabon].—<br />
Monod, 1956:478, 632, figs. 644, 645 [Guinea, Sierra<br />
Leone, Nigeria, Principe].—Rossignol, 1957:116 [key];<br />
1962:121 [Congo].—Crosnier, 1967:339, figs. 31, 32<br />
[Congo, Annobon].—Longhurst, 1958:88 [Sierra Leone].<br />
—Zariquiey Alvarez, 1968:447 [Portugal; references].—<br />
Massuti, 1970:127 [Rio Muni].<br />
Maia goltziana.—Le Loeuff and Intes, 1968, table 1 [Ivory<br />
Coast].<br />
Maia goltziana.—Maurin, 1968a: 30, 48 [Morocco, Spanish<br />
Sahara]; 1968b:484 [Spanish Sahara].<br />
DISTRIBUTION.—Eastern Atlantic, from the<br />
Mediterranean and Portugal S to the Congo,<br />
including Annobon and Principe islands in the<br />
Gulf of Guinea; sublittoral, in 15-200 m.<br />
Maja squinado (Herbst, 1788)<br />
Maja squinado.—Capart, 1951:98, fig. 32 [Spanish Sahara,<br />
Mauritania].—Monod, 1956:474, figs. 638-643 [Spanish<br />
Sahara, Mauritania, Senegal, Guinea].—Figueira, 1960:<br />
11 [Azores].—Guinot and Ribeiro, 1962:75 [Cape Verde<br />
Islands, Angola].—Ribeiro, 1964:18 [Cape Verde Islands].—Forest<br />
and Guinot, 1966:95 [Guinea-Bissau].—<br />
Crosnier, 1967:339 [Gabon; Congo or Gabon].—Monod,<br />
1967:182, pi. 16: fig. 2 [no localities].—Zariquiey Alvarez,<br />
1968:446, figs. 149a, 150g,h [Spain; references].—Christiansen,<br />
1969:131, fig. 54, map 47 [North Atlantic].—<br />
Kensley, 1970:180 [South-<strong>West</strong> Africa].—Ribeiro, 1973:6<br />
[Cape Verde Islands].<br />
Maia squinado.—Sourie, 1954b: 147 [Senegal].<br />
Maia squinado.—Chapman and Santler, 1955:375<br />
[Azores].—Forest and Gantes, 1960:356 [Morocco].—<br />
Gauld, 1960:72 [Ghana].—Maurin, 1968a:48; 1968b:484<br />
[both Spanish Sahara].—Bas, Arias, and Guerra, 1976:<br />
169 [Spanish Sahara].<br />
Maja squinada.—Massuti, 1970:127 [Rio Muni].<br />
SYNONYM.—Maia squinado var. brachydactyla<br />
Balss, 1922.<br />
DISTRIBUTION.—Eastern Atlantic, from the<br />
North Sea southward to South-<strong>West</strong> Africa, including<br />
the Mediterranean; sublittoral to about<br />
75 m.<br />
Subfamily PISINAE Dana, 1851<br />
Genus Apiomithrax Rathbun, 1897<br />
Phycodes A. Milne Edwards, 1869:374 [invalid junior<br />
homonym of Phycodes Guenee, 1852 (Lepidoptera); type-
308<br />
species: Phycodes antennarius A. Milne Edwards, 1869, a<br />
subjective junior synonym of Micropisa violacea A. Milne<br />
Edwards, 1867, by monotypy; gender: masculine].<br />
Apiomithrax Rathbun, 1897b: 164 [substitute name for Phycodes<br />
A. Milne Edwards, 1869; type-species: Phycodes antennarius<br />
A. Milne Edwards, 1869, by monotypy; gender:<br />
masculine].<br />
Apiomithrax bocagei (Osorio, 1887)<br />
Micropisa violacea.—A. Milne Edwards and Bouvier, 1900:<br />
130 [part].—Rathbun, 1925:303, pi. 241: figs. 5-8<br />
[part].—Capart, 1951:93, fig. 30, pi. 2: figs. 12, 13 [part,<br />
all but one specimen from Dakar (Monod, 1956:509)].<br />
[Not Micropisa violacea A. Milne Edwards, 1867.]<br />
Apiomithrax bocagei.—Monod, 1956:508, figs. 692-702.—<br />
Longhurst, 1958:89.—Gauld, 1960:72.—Guinot and Ribeiro,<br />
1962:76.—Forest and Guinot, 1966:105.—LeLoeuff<br />
and hues, 1968:46, 70, table 1, figs. 52, 61; 1969:63, 64,<br />
65.<br />
Micropisa bocagei.—Rossignol, 1957:78, 116 [key], pi. 1: figs.<br />
6-9, pi. 3: fig. 2.<br />
Micropisa (Apiomithras) bocagei.—Rossignol, 1962:122.<br />
Micropisa spinosa Forest and Guinot, 1966:105 [nomen nudum;<br />
discussion of manuscript name].<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: <strong>West</strong> Africa: 1906, F. P. Vermeulen, 1
NUMBER 306 309<br />
recorded many specimens from localities between<br />
Senegal and the Congo. Since 1956 it has been<br />
reported from the following:<br />
Sierra Leone: No specific locality, 8-25 m (Longhurst,<br />
1958).<br />
Ivory Coast: No specific locality (Le Loeuff and Intes,<br />
1969). Off Sassandra, off Fresco, off Grand-Lahou, off Jacqueville,<br />
off Grand-Bassam, 8-40 m (Le Loeuff and Intes,<br />
1968).<br />
Ghana: No specific locality, shore to 20 m (Gauld, 1960).<br />
Principe: Between Ponta da Mina and Ponta Novo Destino,<br />
6 m (Forest and Guinot, 1966).<br />
Congo: Pointe-Noire (Rossignol, 1957). Baie de Pointe-<br />
Noire (Rossignol, 1962).<br />
Angola: No specific locality; Benguela (Forest and<br />
Guinot, 1966). Luanda, beach seine; Porto Amboim, 40 m;<br />
Baia de Benguela, shore (Guinot and Ribeiro, 1962).<br />
* Apiomkhrax violaceus<br />
(A. Milne Edwards, 1867)<br />
Micropisa eryophora De Rochebrune, 1883:167.—Rathbun,<br />
1900a:294 [listed].—Balss, 1922:73 [listed].—Monod,<br />
1956:513.<br />
Herbstia eryophora.—Miers, 1886:49 [listed].—Rathbun,<br />
1893a:93 [listed].<br />
Micropisa violacea.—Capart, 1951:93 [part, not illustrated<br />
specimen].—Sourie, 1954b: 147, 152.—Rossignol, 1957:78,<br />
116 [key], fig. 2, pi. 1: figs. 2-5, pi. 3: fig. 1.—Buchanan,<br />
1958:20, 24.—Crosnier, 1964:31, 32.<br />
Apiomithrax violaceus.—Monod, 1956:502, figs. 682-691.—<br />
Longhurst, 1958:89.—Gauld, 1960:72.—Monod, 1963,<br />
fig. 41 [no material].—Forest and Guinot, 1966:104.—Le<br />
Loeuff and Intes, 1969:64 [discussion].<br />
Micropisa (Apiomithras) violacea.—Rossignol, 1962:122 [erroneous<br />
spelling].<br />
SYNONYMS.—Phycodes antennarius A. Milne Edwards,<br />
1869; Micropisa eryophora De Rochebrune,<br />
1883.<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 70, 33 m, branched Foraminifera, Ic5 (L).<br />
Other Material: Dahomey: Harbor of Cotonou, 26 Mar<br />
1964, H. Hoestlandt, 1 juv 9 (L).<br />
Congo: Off Pointe-Noire, 04°46'15"S, 11°48'15"E, 18 m,<br />
25 Jan 1966, Ombango, A. Crosnier, \6 (W).<br />
DESCRIPTION.—Monod, 1956:500-501 (comparison<br />
with A. violaceus).<br />
Figures: Monod, 1956, figs. 682-691.<br />
Male Pleopod: Monod, 1956, figs. 687-689 (Senegal);<br />
Rossignol, 1957, pi. 3: fig. 1 (Congo).<br />
MEASUREMENTS.—Our specimens are all quite<br />
small, having carapace lengths of 10.5 mm or less.<br />
Monod (1956) recorded a specimen 50 mm long.<br />
REMARKS.—The identity of the species described<br />
by De Rochebrune (1883) as Micropisa<br />
eryophora, from the mouths of the Casamance and<br />
Gambia rivers, has remained unsolved until now.<br />
Miers (1886) placed it in Herbstia and Balss (1922)<br />
retained it in Micropisa but suggested it could be<br />
identified with A. violaceus. Monod (1956) listed it<br />
as ''''Micropisa eryophora" and discussed it after his<br />
account of Apiomithrax, but did not identify it<br />
with any <strong>West</strong> <strong>African</strong> species then known.<br />
De Rochebrune's longer French account (1883:<br />
167, 168), based on a small specimen, 18 mm<br />
long, 15 mm wide, is as follows:<br />
Carapace plus longue que large, couverte ainsi que les<br />
pattes de longs poils grisatres enchevetres; les parties [p. 168]<br />
inferieures garnies de poils courts, rigides; front bifide, a<br />
comes courtes et obtuses; bord externe de 1'orbite arme d'une<br />
dent obtuse, dirigee en dehors; bords lateraux garnis d'epines<br />
assez longues, molles; trois tubercules saillants disposes sur<br />
une ligne transversale au niveau de la region gastrique;<br />
quatre autres epines au centre de la carapace; regions branchiales<br />
et hepatiques supportant des epines assez fortes,<br />
terminees par une etranglement surmonte d'un bouton de<br />
consistance molle; pinces faibles, a doigts ecartes, se touchant<br />
seulement a leur extremite; pattes ambulatoires assez fortes,<br />
laineuses.<br />
Couleur generale brun rougeatre, visibles a travers le<br />
feutrage des poils; pointe des epines rose, pattes rosees.<br />
The body proportions of M. eryophora, length<br />
6/5 width, suggest that it is based upon an Apiomithrax<br />
rather than a Herbstia or Micropisa; small<br />
specimens of Herbstia are relatively slender<br />
(Monod, 1956, figs. 649, 650) and Micropisa ovata<br />
(Monod, 1956, fig. 669) lacks dorsal spines on the<br />
midline of the carapace. That the branchial and<br />
hepatic spines were described as terminating in<br />
spongy buttons suggests that the specimen was<br />
covered with a sponge, not that the spines were<br />
enlarged distally. The mention of long setae on<br />
the carapace and legs suggests that De Rochebrune<br />
was dealing with the relatively hairy A.<br />
violaceus rather than A. bocagei. Finally, the de-
310<br />
scription of three prominent tubercles arranged<br />
in a transverse row across the level of the gastric<br />
region plus four other spines in the center of the<br />
carapace (not described by De Rochebrune as<br />
being in a longitudinal row) is precisely the condition<br />
figured by Monod (1956, fig. 682) for A.<br />
violaceus. We have no hesitation in identifying<br />
Micropisa eryophora De Rochebrune, 1883, with<br />
Micropisa violacea A. Milne Edwards, 1867.<br />
Although Monod (1956:502, synonymy) indicated<br />
that the type-locality of Micropisa violacea<br />
was "St. Vincent du Cap Vert," A. Milne Edwards<br />
(1867a: 33-35) actually had material from<br />
two localities, Sao Vicente and "la cote<br />
d'Angola," and specimens from these two localities<br />
are syntypes. Monod (1933b:506 and 1956:<br />
508) examined the specimen from Angola and,<br />
like A. Milne Edwards, considered it to be conspecific<br />
with the material from the Cape Verde<br />
Islands. However, the type-locality of M. bocagei<br />
also is Angola. In order to insure that the name<br />
Micropisa violacea will continue to be used in the<br />
sense originally employed by A. Milne Edwards<br />
(1867a), we select the specimen figured by him<br />
on the plate (pi. 21: fig. 1) that accompanied his<br />
account of the species in volume 1 of Les Fonds de<br />
la Mer. Apparently the plates that accompanied<br />
this volume were not issued with it but were<br />
issued after livraison 24 of volume 2, published in<br />
1872, i.e., with one of the livraisons of that volume<br />
issued between 1873 and 1876 (see Rehder, 1946:<br />
74, for an account of the dates of publication of<br />
the individual livraisons and some of the figures).<br />
BIOLOGY.—Like A. bocagei, this species apparently<br />
prefers soft bottoms in relatively shallow<br />
water. Although it has been recorded from a<br />
depth of 110-180 m from the Talisman collections,<br />
it usually occurs in depths of 35 m or less. The<br />
single specimen taken by the Pillsbury was collected<br />
from branched Foraminifera in 33 m.<br />
Sourie (1954b) in a study of sand habitats, found<br />
it in Senegal in 10-12 m on bottom with Area and<br />
Pyura, in coarse shelly sand and on fine shelly<br />
sand with mud and with Molgula hannensis (Peres),<br />
2-7 m. In a study of the rocky shore fauna of<br />
Senegal, Sourie (1954a) included it with the "hy-<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
pobiotes lapidicoles mobiles," the "hypobioses detricicoles"<br />
where it was associated with Sidonops<br />
senegalensis (Topsent) and Eurythoe complanata (Pallas),<br />
and as a representative of the "horizons<br />
moyens et inferieurs" on rocky shores. Buchanan<br />
(1958) reported that it was abundant in the sandy<br />
silt community in 8-20 fin (15-36 m) and also in<br />
the inshore fine sand community in 3-8 fm (6-15<br />
m) off Accra, Ghana. Monod (1956) recorded a<br />
few notes on habitat: bottom with Palythoa and<br />
Molgula on sand in 15-20 m, muddy sand in 10<br />
m, and specimens collected at the shore, beach,<br />
or reef. Longhurst (1958) found the species off<br />
Sierra Leone in sandy mud in estuaries or on<br />
shelly sand offshore, in depths between 6 and<br />
25 m. Crosnier (1964) found it off Cameroon in<br />
0-32 m, in sand with Foraminifera on rocky<br />
bottom with gorgonians, and characterized it as<br />
a warm water crustacean. Gauld (1960) noted<br />
that it was common on rocky shores throughout<br />
Ghana and was occasionally taken sublittorally.<br />
Forest and Guinot (1966) recorded the following<br />
habitats: from mud with Area in 32 m off the<br />
Niger delta; from rocks, coral, sand, and calcareous<br />
algae in 3-10 to 35 m off Principe; from<br />
rocks on shore and from calcareous algae, rocks,<br />
coral, sand, on an anchor and in a sponge, in 2 to<br />
10 m off Sao Tome; and from similar bottoms in<br />
7 to 60 m off Annobon.<br />
Ovigerous females have been taken in January<br />
through June and July through September<br />
(Monod, 1956; Forest and Guinot, 1966).<br />
DISTRIBUTION.—This species, like A. bocagei, occurs<br />
off <strong>West</strong> Africa from numerous localities<br />
between Cabo Blanco and Angola; in depths to<br />
110-180 m, generally in less than 35 m; unlike A.<br />
bocagei, it also occurs off Brazil (Rathbun, 1925).<br />
Monod (1956) reported on many specimens from<br />
localities between Mauritania and Gabon; in addition<br />
to Monod's records, A. violaceus has been<br />
recorded from the following:<br />
Senegal: Dakar (Sourie, 1954b). Mouth of the Casamance<br />
River (De Rochebrune, 1883).<br />
Gambia: Mouth of the Gambia River (De Rochebrune,<br />
1883).
NUMBER 306 311<br />
Sierra Leone: No specific locality, in 6-25 m (Longhurst,<br />
1958).<br />
Ghana: Off Accra, 6-36 m (Buchanan, 1958). No specific<br />
locality (rocky shores throughout Ghana) (Gauld, 1960).<br />
Nigeria: Off the mouths of the Niger River, 04°03'N,<br />
06°12'E, 32 m (Forest and Guinot, 1966).<br />
Cameroon: No specific locality, 0-32 m (Crosnier, 1964).<br />
Principe: Baia das Agulhas, 4-8 m; 01°38'35"N, O7°2l'-<br />
35"E, 35 m; Ilheus dos Mosteiros, 3-10 m (Forest and<br />
Guinot, 1966).<br />
Sao Tome: In front of Sao Tome, 8 m; 00°20'N, 06°46'E,<br />
10 m; Ponta Diogo Vaz; Praia Santa Catarina; Ilheu Macaco;<br />
off Ponta Diogo Nunes, shore and 4-5 m; in front of<br />
Ponta Oquedelrei; Morro Peixe; and Ilheu das Cabras, shore<br />
(all Forest and Guinot, 1966).<br />
Annobon: 01°24'04"S, 05°36'45"E, 7-10 m; 01°26'15"S,<br />
05°35'40"E, 60 m; NW coast, Isla Tortuga, 15-40 m (all<br />
Forest and Guinot, 1966).<br />
Congo: Pointe-Noire (Rossignol, 1957). Baie de Pointe-<br />
Noire (Rossignol, 1962).<br />
REMARKS.—Apparently this species is rare off<br />
<strong>West</strong> Africa. Monod (1956) recorded six specimens<br />
and only two were taken by the Pillsbury.<br />
Barnard (1950) has recorded this species from<br />
South Africa and suggested that E. longimana<br />
Stimpson, 1858, should be considered a synonym<br />
of/s. aspera. Capart (1951) considered E. longimana<br />
to be a distinct species, but Griffin (1964, 1974)<br />
left it in synonymy. Crosnier (1970) agreed with<br />
Barnard and Griffin and pointed out additional<br />
similarities between material available to him<br />
and that reported by Barnard. If those considering<br />
E. longimana to be a synonym of E. aspera are<br />
correct, the latter species occurs from Norway<br />
southward to South Africa.<br />
BIOLOGY.—Little information is available for<br />
<strong>West</strong> <strong>African</strong> specimens of E. aspera, but apparently<br />
it prefers a firm substrate with relatively<br />
Genus Eurynome Leach, 1814<br />
large particles in it. Chapman and Santler (1955)<br />
Eurynome Leach, 1814:431 [type-species: Cancer asper Pennant, reported it from rocks in a harbor. Longhurst<br />
1777, by monotypy; gender: feminine; name 1623 on<br />
Official List].<br />
(1958) took one specimen on shelly sand in 100 m<br />
and Forest and Guinot (1966) also reported one<br />
specimen from algae and calcareous algae in<br />
60 m. Our specimens were taken on broken shell<br />
* Eurynome aspera (Pennant, 1777)<br />
Eurynome aspera.—Capart, 1951:86, pi. 2: fig. 4.—Chapmanin<br />
70 m. The overall depth range is wide, from<br />
and Santler, 1955:375.—Monod, 1956:480, figs. 646- 10 m or less to 550 m (Monod, 1956).<br />
648.—Longhurst, 1958:89.—Forest and Guinot, 1966: DISTRIBUTION.—Eurynome aspera is widely dis-<br />
95.—Zariquiey Alvarez, 1968:462, figs. 14g, 153a, 154f<br />
tributed in the eastern Atlantic, from Norway to<br />
[Spain; references].—Christiansen, 1969:126, fig. 52, map<br />
45 [North Atlantic].—Crosnier, 1970:1217, pi. 1: figs. 1-4 Angola, including the Mediterranean, and, pos-<br />
[Mediterranean in part].—Tiirkay, 1976a:25 [listed], 39 sibly, from South Africa, in depths between 10<br />
[Portugal in part].<br />
and 550 m. Monod (1956) reported on material<br />
from Senegal and Principe Island; in addition to<br />
SYNONYMS.—Eurynome scutellata Risso, 1827; Eu-<br />
Monod's records the species has been recorded<br />
rynome boletifera Costa, 1838; tEurynome longimana<br />
from the following:<br />
Stimpson, 1858; Eurynome aspera var. acuta A.<br />
Milne Edwards and Bouvier, 1900.<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Azores: Horta, Faial (Chapman and Santler, 1955).<br />
Sta 68, 70 m, broken shell, 16", 1 juv (L).<br />
Morocco: 33°27.7'N, 08°50.8'W, 161-168 m; and<br />
Undaunted Material: Angola: Sta 96, 162 m, 1
312<br />
*Eurynome parvirostris Forest and Guinot,<br />
1966<br />
Eurynome parvirostris Forest and Guinot, 1966:95, fig. 9.<br />
MATERIAL EXAMINED.—Pillsbury Material: Ghana:<br />
Sta 16, 46 m, mud with Foraminifera, shells, 1$ (W). Sta 23,<br />
42 m, foliate brown to orange bryozoans, 1$ ov (L).<br />
Other Material: Dahomey: Off Grand-Popo, 30 m, Petersen<br />
grab, 23 Feb 1964, Guinean Trawling Survey, Tr 34, Sta 2,<br />
1(5 (L).<br />
DESCRIPTION.—Forest and Guinot, 1966:95.<br />
Figure: Forest and Guinot, 1966, fig. 9.<br />
MEASUREMENTS.—Our specimens have carapace<br />
lengths of 6 mm. The two type-specimens<br />
had carapace lengths of 6.3 mm each.<br />
BIOLOGY.—This species may have different<br />
habitat requirements than E. aspera, for most<br />
specimens have been taken on soft bottoms: mud<br />
with Area in 32 m (Forest and Guinot, 1966), and<br />
mud with Foraminifera and shells in 46 m, or<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
mud with foliate bryozoans in 42 m. The known<br />
depth range is relatively narrow, from 30 to 46<br />
m.<br />
DISTRIBUTION.—Off tropical <strong>West</strong> Africa,<br />
where it has been recorded from off Dahomey,<br />
Ghana, and off the mouths of the Niger River,<br />
04°03'N, 06°12'E, 32 m (Forest and Guinot,<br />
1966).<br />
Genus Herbstia H. Milne Edwards, 1834<br />
Key to Species of Adult Herbstia from <strong>West</strong> Africa<br />
Herbstia H. Milne Edwards, 1834:301 [type-species: Cancer<br />
condyliatus Fabricius, 1787, by monotypy; gender: feminine;<br />
name 1625 on Official List].<br />
Rhodia Bell, 1835:169 [type-species: Rhodia pyriformis Bell,<br />
1835, by monotypy; gender: feminine].<br />
Herbstiella Stimpson, 1871b:93 [type-species: Herbstia depressa<br />
Stimpson, 1860, by original designation; gender: feminine].<br />
Fisheria Lockington, 1877:72 [type-species: Fisheria depressa<br />
Lockington, 1877, a subjective junior synonym of Herbstiella<br />
camptacantha Stimpson, 1871, by monotypy; gender:<br />
feminine].<br />
1. Posterior median margin of carapace with trilobed projection. [Opposable<br />
margins of dactyli of walking legs with fixed, triangular teeth]<br />
H. condyliata<br />
Posterior median margin of carapace with single projection 2<br />
2. Carapace, smooth, regions poorly defined. Upper surface of palm in males<br />
smooth. Opposable margins of dactyli of walking legs with fixed triangular<br />
teeth H. nitida, new species<br />
Carapace rugose, regions well defined. Upper surface of palm in males with<br />
row of tubercles. Opposable margins of dactyli of walking legs lacking<br />
fixed triangular teeth H. rubra<br />
* Herbstia condyliata (Fabricius, 1787)<br />
FIGURE 79<br />
Cancer condyliatus Fabricius, 1787:324.<br />
Herbstia condyliata.—Monod, 1933a:212.—Forest and Games,<br />
1960:356.—Zariquiey Alvarez, 1968:455, fig. 153c,d<br />
[Spain; references].<br />
Herbstia rubra.—Sourie, 1954a: 113, 254, 256, 294; 1954b:<br />
147.—Chapman and Sander, 1955:375.—Monod, 1956:<br />
482, figs. 650-653 [part, not fig. 649]. [Not Herbstia rubra<br />
A. Milne Edwards, 1869.]<br />
SYNONYMS.—Mithrax herbsti Risso, 1827; Mithrax<br />
scaber Costa, 1840.<br />
MATERIAL EXAMINED.—Pillsbury Material: Ghana:<br />
Sta 22, 51 m, rough bottom, Ic5 (L).<br />
Other Material: Madeira: SE coast, near Canical, 32°44'N,<br />
16°44'W, 0-22 m, shore collecting, snorkeling, and diving,<br />
11 Mar 1976, Orwersaagd Sta 48, \S (L). S coast Ponta da<br />
Garajau, 32°38'N, lG^l'W, 5-26 m, diving, 17 Mar 1976,<br />
Orwersaagd Sta 111, 1$ (L).<br />
DESCRIPTION.—Carapace (Figure 79a,c,d) pyriform,<br />
irregular dorsally, ornamented with numerous<br />
spines, regions well marked; surface covered<br />
with thin coat of low, dark setae. Gastric, cardiac,<br />
and intestinal regions each elevated in midline.
NUMBER 306 313<br />
a<br />
FIGURE 79.—Herbstia condyliata (Fabricius). Juvenile, cl 4.2 mm, Sicily: a, carapace; b, dactylus<br />
of fifth pereiopod. Male, cl 10.8 mm, Spain: c, carapace. Male, cl 29.6 mm, Pillsbwy Sta 22: d,<br />
carapace; e, dactylus of fifth pereiopod; /, abdomen; g, apex of gonopod.<br />
Gastric region with 3 raised protogastric tubercles<br />
in transverse line anteriorly (Figure 79
314<br />
podus, ventral margin with slender spines and<br />
usually with 1 or 2 triangular teeth (Figure 19b,e),<br />
large in juvenile, smaller in adult.<br />
Male gonopod and abdomen as shown (Figure<br />
79/*).<br />
MEASUREMENTS.—Carapace lengths of males<br />
7.8 to 29.6 mm, of female 18 mm.<br />
REMARKS.—Herbstia condyliata can readily be<br />
distinguished from other <strong>West</strong> <strong>African</strong> species of<br />
the genus, H. rubra and H. nitida, new species, by<br />
its larger size, much more spinulose carapace, and<br />
presence of a trilobed projection on the posterior<br />
margin of the carapace (as shown in Monod,<br />
1956, fig. 650, and herein, Figure *19d), where it<br />
is almost quadrilobed. The chelipeds in adult<br />
males of H. condyliata are greatly enlarged, being<br />
much longer than the carapace; in the two<br />
smaller <strong>West</strong> <strong>African</strong> species the chelipeds in<br />
adult males are subsimilar to those of females.<br />
Herbstia condyliata agrees with H. nitida, new species,<br />
and differs from H. rubra in having fixed,<br />
triangular teeth on the opposable margin of the<br />
dactyli of the walking legs.<br />
Because smaller specimens of this species have<br />
been confused with the adults of the two smaller<br />
<strong>West</strong> <strong>African</strong> species, we have sketched the carapace<br />
shape in a juvenile, cl. 4.2 mm (W), from<br />
Sicily (Figure 79a) and a larger specimen from<br />
Spain, cl. 10.8 mm (L) (Figure 79c), as well as an<br />
outline of our adult from Ghana (Figure 79d). In<br />
the smallest specimen, the carapace is very slender,<br />
its dorsal armature is reduced, and there is<br />
but one posterior spine. In the specimen from<br />
Spain the characteristic three posterior spines of<br />
the adult are developed, although the spinulation<br />
of the carapace is still reduced. Monod (1956, fig.<br />
650) figured a female 15.5 mm long, in which the<br />
carapace is still more slender than in the adult<br />
but in which the posterior spines of the carapace<br />
are well developed. In Monod's figure, the transverse<br />
line of three protogastric tubercles is not<br />
shown.<br />
A. Milne Edwards and Bouvier (1900:128) suggested<br />
that H. rubra might only be a race of H.<br />
condyliata and Monod (1956:485) more or less<br />
agreed with this:<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
II n'est nullement impossible que l'etude de materiaux<br />
plus nombreux, et provenant en particulier des regions marocaine<br />
ou saharienne, n'oblige a tenir un jour rubra pour<br />
synonyme de condyliata, ou tout au moins comme une forme<br />
subordonnee, moins tuberculee, et aussi plus petite.<br />
Monod was working with juveniles of H. condyliata<br />
as well as adults of both other species, and,<br />
in view of the similarities between specimens of<br />
similar size of all three species, it is not surprising<br />
that they were not separated by him.<br />
Although it is impossible to be certain in view<br />
of the relatively small number ofHerbstia recorded<br />
from <strong>West</strong> Africa and the relatively unreliable<br />
literature records, it seems likely that H. rubra is<br />
restricted to the Cape Verde Islands, H. nitida,<br />
new species, occurs only on the offshore islands of<br />
the Gulf of Guinea, and H. condyliata occurs on<br />
the continental shelf.<br />
The male pleopod of H. condyliata, the typespecies<br />
of the genus, is shown in Figure 19g. It<br />
resembles that shown by Garth (1958, pi. S: figs.<br />
7, 8) for the eastern Pacific species H. pubescens<br />
Stimpson and H. pyriformis (Bell) in that there is<br />
a subapical rounded, spinulose knob flanked by<br />
a distinct notch; this knob is present on the<br />
gonopods of all three species reported here. As<br />
noted by Garth (1958:300) in his key to <strong>West</strong><br />
American Herbstia, the other western American<br />
species lack the subapical notch on the male<br />
pleopod. Further study may well indicate that<br />
Herbstia, as currently recognized, comprises representatives<br />
of more than one genus.<br />
BIOLOGY.—Sourie (1954b) found this species<br />
on coarse, shelly sand, with mud, bottom with<br />
Area and Pyura, in 2-7 m, at Anse de Hann, Baie<br />
de Dakar, and subsequently included the species<br />
as a component of the rocky shore fauna of<br />
Senegal. Monod (1933a:212) reported two specimens<br />
associated with sponges, and, in 1956, reported<br />
a specimen collected during a spring low<br />
tide. The single specimen taken by the Pillsbury<br />
was taken on rough bottom at a depth of 51 m.<br />
DISTRIBUTION.—Mediterranean Sea and adjacent<br />
Atlantic south to Ghana. Since 1956 it has<br />
been recorded from Morocco: Temara, Skhirat,<br />
and David (Forest and Games, 1960).
NUMBER 306 315<br />
Barrois (1888) recorded the species from the low tubercles and several larger spines arranged<br />
Azores, and also indicated, without citing a ref- in dorsal and ventral rows, spination variable,<br />
erence, that the species occurred in the Canary absent in some specimens; carpus with numerous<br />
Islands. The specimens reported from Horta and tubercles and 1 larger spine on outer margin.<br />
Pasteleiro, Faial, Azores, by Chapman and San- Palm not markedly inflated in males, with 1 or 2<br />
der (1955) as H. rubra may be referable to H. (usually 1) posteriorly directed spines on dorsal<br />
condyliata; H. rubra is known with certainty onlyarticular<br />
knob, remainder of surface smooth but<br />
from the Cape Verde Islands.<br />
setose. Fingers shorter than palm, cutting edges<br />
crenulate, not strongly toothed or gaping.<br />
*Herbstia nitida, new species<br />
Walking legs slender, long, fifth leg longer than<br />
carapace, ornamented with corneous bristles and<br />
FIGURE 80<br />
longer simple setae. Merus with row of corneous<br />
bristles dorsally, occasionally with dorsal and pos-<br />
?Herbstia rubra.—Monod, 1956:482 [part].—Forest and<br />
Guinot, 1966:97. [Not Herbstia rubra A. Milne-Edwards,<br />
1869.]<br />
terior spines or with irregular tubercles posteriorly,<br />
not heavily armed. Dactylus (Figure 80/)<br />
shorter than propodus, ventral margin with 1-4<br />
MATERIAL EXAMINED.—Pillsbury Material: Annobon: prominent, triangular teeth and slender spines.<br />
Sta 275, 9-69 m, rubble of coralline algae, 3c5, 3$ (2 ov), 1 Male abdomen and gonopods as shown (Figure<br />
juv (W). Sta 282, 18-37 m, nodular coralline algae, 3
316<br />
a<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 80.—Herbstia nitida, new species, paratype, male, cl 9.0 mm, Pillsbury Sta 282: a, dorsal<br />
view; b, carapace; c, front, dorsal view; d, front, ventral view; e, chela; /, dactylus of fifth<br />
pereiopod; g, abdomen; h, apex of gonopod.<br />
DISPOSITION OF TYPES.—The holotype (Crust.<br />
D.31780), a male with a carapace length of 8.9<br />
mm from Pillsbury Sta 282, is in the Rijksmuseum<br />
van Natuurlijke Historie, Leiden. Paratypes have<br />
been deposited in the <strong>Smithsonian</strong> <strong>Institution</strong>,<br />
Washington, D.C., and in Leiden.<br />
BIOLOGY.—All of our material came from Annobon<br />
Island, in depths between 9 and 55 m, in<br />
the beds of calcareous algae described by Forest<br />
(1959) and Voss (1966). If our assumption is<br />
correct that the material from the offshore islands<br />
of the Gulf of Guinea previously reported as H.<br />
rubra actually is identifiable with H. nitida, the<br />
latter species lives on rough bottom in moderate<br />
depths on rock, coral, algae, and calcareous algae.<br />
Forest and Guinot (1966) recorded the following<br />
habitats and depths (all under H. rubra): Principe,<br />
calcareous algae in 37-73 m; Sao Tome (6 localities),<br />
in rocks and coral in 0-6 and at 30 m, in<br />
calcareous algae in 6 and 10 m, in a sponge from<br />
rocks and calcareous algae in 2-6 m; and Annobon<br />
(5 localities), in rocks and coral, in calcareous<br />
algae, sand, and coral in 7-10 m, and in calcareous<br />
algae in 23-60 m.<br />
Ovigerous females have been recorded in May<br />
and June (Forest and Guinot, 1966; Pillsbury).<br />
ETYMOLOGY.—The specific epithet is from the<br />
Latin nitidus, shining, neat, referring to the<br />
smooth carapace of this species.<br />
DISTRIBUTION.—Gulf of Guinea, where it is<br />
known to occur off Annobon in 15-55 m. We<br />
9
NUMBER 306 317<br />
suspect that all of the records of H. rubra given<br />
below are based on H. nitida:<br />
Principe: No specific locality, 15 m (Monod, 1956). 01°-<br />
43'10"N, 07°28'20"E, 73 m, and 01°43'N, 07°28'55'E, 37 m<br />
(Forest and Guinot, 1966).<br />
Sao Tome: Off Sao Tome, 8 m; 00°20'N, 06°46'E, 10 m;<br />
Ponta Diogo Vaz, 0-6 m and 30 m; in front of Ponta<br />
Oquedelrei, 6 m; and Morro Peixe, 2-6 m (all Forest and<br />
Guinot, 1966).<br />
Annobon: No specific locality, 12 m (Monod, 1956).<br />
01°24'04"S, 05°36'45"E, 7-10 m; O1°27.5'S, 05°36.5'E,<br />
35 m; N of San Antonio, 23 m; 01°26'15"S, 05°35'40"E, 60<br />
m; and Isla Tortuga, NW coast (all Forest and Guinot,<br />
1966).<br />
Herbstia rubra A. Milne Edwards, 1869<br />
FIGURE 81<br />
Herbstia rubra A. Milne Edwards, 1869:354.—Miers, 1886:49,<br />
pi. 7: fig. 1,1a.—Rathbun, 1893a:93 [listed].—A. Milne<br />
Edwards and Bouvier, 1900:128, pi. 19: fig. 17 [fig. 16 in<br />
text].—Balss, 1922:73 [listed].—Monod, 1956:482, fig. 649<br />
[part, not fig. 650].—Guinot and Ribeiro, 1962:73.—Ribeiro,<br />
1964:19.<br />
MATERIAL EXAMINED.—Pillsbury Material: None.<br />
Other Material: Cape Verde Islands: Sao Vicente, 20 m, 26<br />
Jul 1883, Talisman, \6, 1$ (L, W). Sao Vicente, Challenger,<br />
BMNH 84.31, 1
318 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
which the chelipeds of the male are not greatly<br />
enlarged and there is but one posterior projection<br />
on the carapace. This species differs from H.<br />
nitida, new species, in having the regions of the<br />
carapace better developed, more dorsal tubercles<br />
and spines on the carapace, a narrower posterior<br />
projection on the carapace, more spines on stouter<br />
walking legs, a row of spines on the upper surface<br />
of the palm in males, and no triangular teeth on<br />
the opposable margin of the dactylus of the walking<br />
legs.<br />
An outline of the carapace and a sketch of the<br />
dactylus of a walking leg are shown in Figure 81.<br />
BIOLOGY.—There is little information available<br />
on the ecology of this species. It was reported<br />
from coralline algae in 75 m (A. Milne Edwards<br />
and Bouvier, 1900), and near shore and in 6 to<br />
20 m (Guinot and Ribeiro, 1962; Ribeiro, 1964).<br />
DISTRIBUTION.—<strong>West</strong> Africa, from the Cape<br />
Verde Islands (the type-locality) and possibly the<br />
Azores; most records in the literature require<br />
verification. They include the following:<br />
Cape Verde Islands: No specific locality (A. Milne Edwards,<br />
1869). Sao Vicente (Miers, 1886). Praia Matiota, Sao<br />
Vicente, shore, and Porta da Furna, Brava, 6-20 m (Guinot<br />
and Ribeiro, 1962; Ribeiro, 1964). Between Ilheu Branco<br />
and Ilheu Raso, 75 m (A. Milne Edwards and Bouvier,<br />
1900). Ponta do Sol, Ilha de Santo Antao (Monod, 1956).<br />
Genus Micropisa Stimpson, 1858<br />
Micropisa Stimpson, 1858a:218 [type-species: Micropisa ovala<br />
Stimpson, 1858, by monotypy; gender: feminine].<br />
Micropisa ovata Stimpson, 1858<br />
Micropisa ovata.—Monod, 1956:495, figs. 669-681 [Cape<br />
Verde Islands, Senegal, Gabon(?)].—Guinot and Ribeiro,<br />
1962:74 [Cape Verde Islands].—Ribeiro, 1964:19 [Cape<br />
Verde Islands].<br />
DISTRIBUTION.—Off tropical <strong>West</strong> Africa, from<br />
the Cape Verde Islands and Senegal, with one<br />
doubtful record from Gabon; sublittoral, from<br />
less than 10 to more than 110 m.<br />
Genus Pisa Leach, 1814<br />
Arctopsis Lamarck, 1801:155 [type-species: Arctopsis lanata<br />
Lamarck, 1801, by monotypy; gender: feminine; sup-<br />
pressed for purposes of the Law of Priority but not for<br />
those of the Law of Homonymy by the International<br />
Commission on Zoological Nomenclature in opinion 708<br />
(1964); name 1701 on Official Index].<br />
Pisa Leach, 1814:431 [type-species: Cancer biaculeatus Montagu,<br />
1813, a subjective junior synonym of Maja armata<br />
Latreille, 1803, by monotypy; gender: feminine; name<br />
1597 on Official List}.<br />
Blastus Leach, 1814:431 [type-species: Cancer tetraodon Pennant,<br />
1777, by monotypy; gender: masculine].<br />
*Pisa armata (Latreille, 1803)<br />
Maja armata Latreille, 1803c:98.<br />
Pisa Gibbsii.—Studer, 1882:335.<br />
Pisagibbsi.—Capart, 1951:90, fig. 28, pi. 2: fig. 23.—Monod,<br />
1956:486, fig. 654.—Rossignol, 1957:116 [key].—Figueira,<br />
1960:12.—Crosnier, 1964:34.<br />
Pisa armata.—Forest and Gantes, 1960:356.—Forest and<br />
Guinot, 1966:98.—Zariquiey Alvarez, 1968:454, figs.<br />
151d, 152e, 154d [Spain; references].—Christiansen, 1969:<br />
124, fig. 51, map 44 [North Atlantic].—Crosnier, 1970:<br />
1215 [listed], 1218.—Turkay, 1976a:25 [listed], 39 [Portugal].<br />
Pisa.—Voss, 1966:31.<br />
Pisa gibbi.—Maurin, 1968b:486 [erroneous spelling].<br />
SYNONYMS.—Cancer biaculeata Montagu, 1813;<br />
Pisa gibbsii Leach, 1815.<br />
MATERIAL EXAMINED.—Pillsbury Material: Nigeria:<br />
Sta 230, 82-97 m, hard ground with gorgonians, coral, and<br />
rock, 1
NUMBER 306 319<br />
habits moderate depths on rough ground on the<br />
shelf. The Pillsbury specimen was taken on hard<br />
ground with gorgonians, coral, and rock in 82-<br />
97 m. Capart (1951) reported material collected<br />
at depths from 20-30 to 110 m, on sandy mud,<br />
that from 20-30 m being the shallowest record<br />
off <strong>West</strong> Africa. Crosnier (1964) characterized it<br />
as a cold-water species, occasionally occurring<br />
deeper than 50 m off Cameroon. Forest and<br />
Guinot (1966) recorded it from mud, sand, and<br />
"sable construit" in 65-75 m off Senegal and on<br />
mud, shell, and Cidaris in 60-73 m off Guinea-<br />
Bissau. Maurin (1968b) found it off Morocco in<br />
40-60 m on bottom with sponge, Suberites, and<br />
asteroid, Allopatiria.<br />
Ovigerous females have been recorded from<br />
<strong>West</strong> <strong>African</strong> localities in March, May, August,<br />
October, November (Capart, 1951; Monod,<br />
1956), and summer (Figueira, 1960).<br />
DISTRIBUTION.—Eastern Atlantic, from the<br />
southern North Sea and England to Angola,<br />
Mediterranean; sublittoral, to at least 162 m.<br />
Monod (1956) reported material from Senegal,<br />
Guinea, and Angola; other records include the<br />
following localities.<br />
Azores: Horta (Figueira, 1960).<br />
Morocco: Temara (Forest and Games, 1960).<br />
Mauritania: Bane d'Arguin, 40-60 and 90-100 m<br />
(Maurin, 1968b).<br />
Cape Verde Islands: 15°40'N, 23°06'W, 38 fm [70 m]<br />
(Studer, 1882).<br />
Senegal: 12°55.5'N, 17°33'W, 65-75 m (Forest and<br />
Guinot, 1966).<br />
Guinea-Bissau: 10°19'N, 16°34'W, 60-73 m (Forest and<br />
Guinot, 1966).<br />
Nigeria: 06° 1 l'N, 03°36'E to 06° 10'N, 03°38'E, 82-97 m<br />
(Voss, 1966).<br />
Cameroon: No specific locality, more than 50 m (Crosnier,<br />
1964).<br />
Angola: 16°41'S, 11°21'E, 162 m (Crosnier, 1970).<br />
The latter record appears to be the southernmost<br />
as well as the deepest for the species.<br />
*Pisa calva Forest and Guinot, 1966<br />
Une espece nouvelle de Pisa.—Forest, 1959, pi. 3: fig. 1. Pisa<br />
(n. sp.).—Forest, 1959:19.<br />
Pisa calva Forest and Guinot, 1966:99, figs. 10, lla-f, 13.<br />
MATERIAL EXAMINED.—Pillsbury Material: Annobon:<br />
Sta 275, 9-69 m, rubble of coralline algae, 66, 4$ (3 ov) (L).<br />
Sta 282, 18-37 m, nodular coralline algae, 16*, 7$ (5 ov), 2<br />
juv (L, W). Sta 283, 51-55 m, nodular coralline algae,<br />
116\ 89 (5ov), 3juv(W).<br />
Other Material: Annobon: 01°24'S, 05°37'30"E, 7-8 m, 11<br />
Dec 1965, Ombango, A. Crosnier, 29 (W).<br />
DESCRIPTION.—Forest and Guinot, 1966:100.<br />
Figures: Forest and Guinot, 1966, figs. 10,<br />
lla-f, 13.<br />
Male Pleopod: Forest and Guinot, 1966, fig. 13.<br />
MEASUREMENTS.—Carapace lengths of males 5<br />
to 11 mm, of females 5 to 8 mm, of ovigerous<br />
females 5 to 10 mm, of juveniles 3 to 5 mm. The<br />
largest specimens recorded by Forest and Guinot<br />
(1966) were a male 14 mm long and a female 10.5<br />
mm long.<br />
REMARKS.—As Chace (1966:653, footnote)<br />
pointed out, this species is very close to Pisa<br />
sanctaehelenae Chace (1966:651, fig. 14) from Saint<br />
Helena. However, P. calva has less divergent rostral<br />
spines and much more slender pereiopods<br />
(Figure 82), with 10 or 11 rather than 5-7 fixed,<br />
triangular teeth on the flexor margin of the dactylus.<br />
In our specimens, the propodus of the sec-<br />
FIGURE 82<br />
FIGURE 82.—Pisa calva Forest and Guinot, male, cl 11.0 mm,<br />
Pisa carinimana.—Monod, 1956:488 [part] [not Pisa carinimana Pillsbury Sta 283: a, second pereiopod; b, dactylus of second<br />
Miers, 1879].<br />
pereiopod, enlarged.
320<br />
ond pereiopod in the males is not markedly swollen<br />
distally as in P. sanctaehelenae.<br />
Apparently Monod's (1956) record of P. carinimana<br />
from Annobon was based on this species<br />
(Forest and Guinot, 1966:104) and his record of<br />
P. carinimana from Principe may also be based on<br />
P. calva. However, Forest and Guinot, who reexamined<br />
Monod's material from Annobon, could<br />
not locate the specimen from Principe and they<br />
noted (1966:104) that both species were taken at<br />
the same station off Principe by the Calypso.<br />
BIOLOGY.—The Pillsbury representatives of this<br />
species all were taken off Annobon Island in beds<br />
of coralline algae in 9-69 m; the Ombango specimens<br />
were taken at Annobon in 7-8 m. Forest<br />
and Guinot (1966) recorded the following habitats<br />
for their material: Principe (5 stations): on<br />
mud, calcareous algae, shell, rocks, and coral in<br />
31-73 m; Sao Tome (12 stations): from a beach<br />
seine in 0-4 m and in rocks, coral, calcareous<br />
algae, "epave," algae, sand, and mud in 5-50 m;<br />
Annobon (6 stations): calcareous algae, sand,<br />
coral, algae, and rocks in 7-60 m.<br />
Ovigerous females have been collected in May,<br />
June, and July.<br />
DISTRIBUTION.—<strong>West</strong> Africa, from the offshore<br />
islands of the Gulf of Guinea; sublittoral, from 0-<br />
4 to 73 m. Records in the literature include the<br />
following:<br />
Principe: 01°38'25"N, 07 o 22 / 05 w E, 31 m; 01°43'10"N,<br />
07°28'20"E, 73 m; 01°43'N, O7°28'55"E, 37 m; in front of<br />
Baia de Santo Antonio, 50 m; Tinhosa Grande Island, 12 mi<br />
[19.3 km] SSW of Principe, 01°20'45"N, 07°17'37"E, 25-40<br />
m (all Forest and Guinot, 1966).<br />
Sao Tome: In front of Sao Tome, 8 m; 00°20'N, 06°46'E,<br />
10 m; N of Ilheu das Cabras, 32 m; Ponta Diogo Vaz, 30 m;<br />
Praia Santa Catarina, 15-18 m; 00°25'40"N, 06°40'10"E,<br />
50 m; 00°25'15"N, 06°43'05"E, 8-30 m; Baia de Ana de<br />
Chaves, 5 m; in front of Ponta Oquedelrei, 6 m; Morro<br />
Peixe, 0-4 m; in front of Ponta Sao Sebastiao, 11 m (all<br />
Forest and Guinot, 1966).<br />
Annobon: No specific locality, 12 m (Monod, 1956).—<br />
NW coast, Isla Tortuga, 15-40 m; N of Santo Antonio,<br />
23 m; 01°27.5'S, 05°36.5'E, 35 m; 01°26'15 w S, O5°35'4O"E,<br />
60 m; 01°25'10"S, 05°36'10"E, 20-25 m; 01°24'04"S,<br />
05°36'45"E, 7-10 m (all Forest and Guinot, 1966).<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
*Pisa carinimana Miers, 1879<br />
Pisa carinimana.—Capart, 1951:87, fig. 27, pi. 2: fig. 8.—<br />
Sourie, 1954b: 147.—Monod, 1956:488, figs. 655-668<br />
[part].—Rossignol, 1957:116 [key].—Longhurst, 1958:<br />
89.—Forest, 1959:19 [discussion].—Gauld, 1960:72.—<br />
Rossignol, 1962:122.—Guinot and Ribeiro, 1962:74.—<br />
Forest and Guinot, 1966:99, figs, llg, 12.—Zariquiey<br />
Alvarez, 1968:452, fig. 155a [Spain; references].—Uschakov,<br />
1970:439, 455 [listed].—Tiirkay, 1957a: 71<br />
[listed], 74.<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 69, 29 m, coral or rock, 1$ ov (L). Sta 70, 33 m, branched<br />
Foraminifera, 86, 8$ (7 ov) (W).<br />
Ivory Coast: Sta 42, 62-75 m, mud with brown, branched<br />
Foraminifera, 16 (W). Sta 46, 38-42 m, mud with dense<br />
Jullienella, 4
NUMBER 306 321<br />
100 m off Ghana. The Pillsbury collected this<br />
species in depths between 29 and 82 m, usually<br />
on relatively soft bottom, sand and mud, or mud<br />
with bryozoans or Foraminifera; two stations<br />
were made on rock and coral. Monod (1956)<br />
recorded material collected on bottom with Palythoa<br />
and Molgula and sand in 15-20 m, on sand in<br />
10-45 m, muddy sand in 10-15 m, and on pebbles<br />
in 14.5 m. Guinot and Ribeiro (1962) reported<br />
material from Angola in depths between 5 and<br />
90 m on sand and muddy sand. Forest and Guinot<br />
(1966) noted the following habitats for the species:<br />
mud, shell, gorgonians, and ascidians in 43-<br />
45 m; mud, sand, and compact sand [sable construit]<br />
in 65-75 m; mud and shells in 18-30 m;<br />
muddy sand and Foraminifera in 21-27 m;<br />
gravel, shells, and Foraminifera in 50 m; mud<br />
with Area in 32 m, and mud, calcareous algae,<br />
and shells in 31 m. This species was taken with P.<br />
calva by the Calypso at this latter station off Principe<br />
Island. Uschakov (1970) found the species in<br />
clear water on hard sand in depths greater than<br />
10 m off Guinea. Apparently this species prefers<br />
habitats on softer bottoms than does P. calva,<br />
which the Pillsbury took only in coralline algae.<br />
Ovigerous females have been recorded in all<br />
months but July (Monod, 1956; Forest and<br />
Guinot, 1966).<br />
DISTRIBUTION.—Pisa carinimana is an eastern Atlantic<br />
species, known from localities between the<br />
Canary Islands (the type-locality) and Spanish<br />
Sahara southward to Angola (12°36'S), including<br />
Melilla in the Mediterranean (Zariquiey Alvarez,<br />
1968); littoral and sublittoral to a depth of about<br />
100 m. Monod (1956) recorded it from numerous<br />
localities between Mauritania and Gabon; other<br />
records include the following:<br />
Spanish Sahara: 21°05'N, 17°14'W, 43-45 m (Forest and<br />
Guinot, 1966).<br />
Mauritania: Off Cap Blanc, 20°37.3'N, 17°24.4'W, 57 m<br />
(Tiirkay, 1975a)<br />
Senegal: Baie de Dakar, 10-12 m (Sourie, 1954b).<br />
12°55.5'N, 17°33'W, 65-75 m (Forest and Guinot, 1966).<br />
Guinea: No specific locality, depth greater than 20 m<br />
(Uschakov, 1970). 09°40'N, 14°05'W, 18 m, and 09°36'N,<br />
13°57VJ, 18-30 m (Forest and Guinot, 1966).<br />
Sierra Leone: No specific locality, 12-34 m (Longhurst,<br />
1958).<br />
Ivory Coast: 05°02.5'N, 05°25'W, 21-27 m (Forest and<br />
Guinot, 1966)<br />
Ghana: Off Accra, 14-100 m (Gauld, 1960). 04°36.5'N,<br />
0l°31'W, 50 m (Forest and Guinot, 1966).<br />
Nigeria: Off the mouths of the Niger River, 04°03'N,<br />
06°12'E, 32 m (Forest and Guinot, 1966).<br />
Principe: No specific locality (Forest, 1959). 01°38'25"N,<br />
07°22'05"E, 31 m (Forest and Guinot, 1966).<br />
Gabon: W of Pointe Claire (as Pointe Clara) and mouth<br />
of Gabon River, 20-40 m (Rossignol, 1962).<br />
Congo: Baie de Pointe-Noire (Rossignol, 1962).<br />
Cabinda: No specific locality, in 38-39 m (Guinot and<br />
Ribeiro, 1962).<br />
Angola: Baia da Caota, Benguela, 11-12 m, 13 m, 16 m,<br />
18 m, and 30 m; Baia Farta, 22 m, 22-28 m, and 90 m;<br />
Sombreiro, 5 m; between Sombreiro and Ponta da Caruita,<br />
23 m (all Guinot and Ribeiro, 1962).<br />
Pisa nodipes (Leach, 1815)<br />
Pisa nodipes.—Monod, 1956:486 [references].—Forest and<br />
Games, 1960:356 [Morocco].—Zariquiey Alvarez, 1968:<br />
454, figs. 151e, 152f, 154e, [Spain; references].—Maurin,<br />
1968a:59, 107 [Mauritania, Mediterranean]; 1968b:486,<br />
489 [Mauritania].<br />
SYNONYM.—Inachus musivus Otto, 1821.<br />
DISTRIBUTION.—Eastern Atlantic, from the Canary<br />
Islands, the Cape Verde Islands, Morocco,<br />
Mauritania, and the Mediterranean; sublittoral,<br />
to about 75 m.<br />
Pisa tetraodon (Pennant, 1777)<br />
Pisa Mraodon.—Caparx, 1951:92, fig. 29, pi. 2: fig. 16 [Mauritania].—Monod,<br />
1956:485 [references].—Forest and<br />
Games, 1960:356 [Morocco].—Zariquiey Alvarez, 1968:<br />
452, figs. 6d, 151a, 152a, 154a [Spain; references].<br />
SYNONYMS.—Cancer hircus Fabricius, 1781 (see<br />
Rathbun, 1925:195); Cancer praedo Herbst, 1796;<br />
Pisa convexa Brandt, 1880.<br />
DISTRIBUTION.—Eastern Atlantic, from England<br />
to Cabo Blanco, Mauritania, Mediterranean;<br />
sublittoral, shallow water to about 50 m<br />
(100 m in Monod, 1956).
322<br />
Family MIMILAMBRIDAE Williams, 1979<br />
MIMILAMBRIDAE Williams, 1979:399.<br />
This family, comprising one genus and species,<br />
does not occur within the study area.<br />
Family PARTHENOPIDAE MacLeay, 1838<br />
PARTHENOPINA MacLeay, 1838:55, 58 [corrected to Parthenopidae<br />
by Bell, 1844:45; name 362 on Official List].<br />
AETHRINAE Dana, 1851b: 127 [originally cited as Oethrinae].<br />
CRYPTOPODIINAE Stimpson, 1871a: 137.<br />
HEPATINAE Stimpson, 1871a: 154 [see p. 325].<br />
EUMEDONINAE Neumann, 1878:17.<br />
LAMBRINAE Neumann, 1878:17.<br />
EASTERN ATLANTIC GENERA.—Five, Daldorjia,<br />
Heterocrypta, Parthenope, Sakaila, new genus, and<br />
Solenolambrus, each represented by species occurring<br />
off tropical <strong>West</strong> Africa.<br />
EASTERN ATLANTIC SPECIES.—Eleven, of which<br />
eight occur off <strong>West</strong> Africa. Several name changes<br />
have been made since this family was studied by<br />
Monod (1956), as follows:<br />
Name in Monod<br />
Osachila stimpsoni<br />
(Calappidae)<br />
Lambrus massena (including<br />
various forms)<br />
Lambrus miersi<br />
Lambrus macrochelos<br />
Lambrus expansus<br />
Heterocrypta maltzani<br />
Solenolambrus noordendei<br />
Parthenope bouvieri<br />
Current Name<br />
Sakaila africana,<br />
new genus, new species*<br />
(Parthenopidae)<br />
Parthenope massena*<br />
Parthenope miersii*<br />
Parthenope notialis,<br />
new species*<br />
Parthenope expansa*<br />
Heterocrypta maltzami*<br />
Solenolambrus noordendei*<br />
Daldorjia bouvieri<br />
Three nominal species occur to the north of the<br />
tropical region:<br />
Heterocrypta marionis A. Milne Edwards, 1881.<br />
In deep water off the Azores, Toulon, France, and<br />
the Bay of Biscay (Bouvier, 1940).<br />
Parthenope angulifrons Latreille, 1828. Mediterranean<br />
(Zariquiey Alvarez, 1968), with one questionable<br />
record from Senegal (Monod, 1956:587).<br />
Parthenope macrochelos (Herbst, 1790). Mediter-<br />
ranean and adjacent Atlantic (Zariquiey Alvarez,<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
1968). See under P. notialis, new species (p. 335),<br />
for possible records along the NW <strong>African</strong> coast.<br />
The status of Lambrus spinosissimus Osorio, 1923<br />
(see Nobre, 1936:85) remains to be determined.<br />
One specimen originated from the latitude of<br />
Lisbon, the other from Morocco. Osorio's account<br />
is not available to us. According to Nobre's quote<br />
of the original, L. spinosissimus (cl 33 mm, cb 37<br />
mm) is a larger species than the <strong>West</strong> <strong>African</strong> P.<br />
notialis (cl 5-21 mm, cb 11-23 m) (see p. 334).<br />
Osorio described the upper surface of the palm of<br />
the cheliped as follows (our translation): "The<br />
upper surface of the hand also shows spines,<br />
rather widely spaced one from the other, and the<br />
lower surface is covered with granules, which do<br />
not stop at the base of the fingers, but, on the<br />
contrary, are widely distributed on them." This<br />
description applies well to P. macrochelos; in P.<br />
notialis the upper surface of the palm of the<br />
cheliped is almost smooth. We believe that L.<br />
spinosissimus Osorio, 1923 should be identified<br />
with the northern Parthenope macrochelos (Herbst,<br />
1790) rather than the smaller tropical species, P.<br />
notialis.<br />
Subfamily AETHRINAE Dana, 1851<br />
Genus Heterocrypta Stimpson, 1871<br />
Heterocrypta Stimpson, 1871b: 102 [type-species: Cryptopodia<br />
granulata Gibbes, 1850, by original designation; gender:<br />
feminine; name 1626 on Official List].<br />
* Heterocrypta maltzami Miers, 1881<br />
Heterocrypta Maltzami Miers, 1881a:209, pi. 13: fig.l.<br />
Heterocrypta Maltzani.—Miers, 1881a:364, 374.—A. Milne Edwards<br />
and Bouvier, 1900:121, pi. 19: fig.6 [part ?].—Balss,<br />
1921:54.—Bouvier, 1940:315 [part ?].<br />
Heterocrypta maltzani.—Ortmann, 1893b:417.—Rathbun,<br />
1900a:296.—Sourie, 1954b: 150.—Monod, 1956:589, figs.<br />
862-867.—Longhurst, 1958:89.—Gauld, 1960:72.—<br />
Guinot and Ribeiro, 1962:80.—Rossignol, 1962:123.—Ribeiro,<br />
1964:21.—Forest and Guinot, 1966:120.—Zariquiey<br />
Alvarez, 1968:442 [Spain; references].—Maurin,<br />
1968b:486.—Tiirkay, 1975a:71 [listed], 74, fig. 6.<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 68, 70 m, broken shell, 1
NUMBER 306 323<br />
Ivory Coast: Sta 46, 38-42 m, mud with denseJullienella,<br />
29 (1 ov) (L).<br />
Ghana: Sta 23, 42 m, foliate brown to orange bryozoans,<br />
16* (L). Sta 24, 35-37 m, dark red bryozoans, 3
324<br />
tion to consider that "le specimen de H. maltzani<br />
Miers signale par T. Odhner (1923 p. 20) de Port<br />
Alexandre (Angola), appartient vraisemblablement<br />
a cette espece nouvelle [= Solenolambrus noordendei]"<br />
Monod (1956:590, 595) seemed to accept<br />
Capart's conclusions. Guinot and Ribeiro (1962:<br />
80, 81) showed that H. maltzami does occur rather<br />
commonly in Angola and concluded that "on<br />
peut done supposer que 1 'echantillon d'Odhner<br />
appartient bien a H. maltzani." This conclusion<br />
seems to be accepted by Grosnier (1967:340, 1970:<br />
1219). As long as Odhner's specimen has not been<br />
reexamined, speculation on its identity is futile:<br />
The size of the specimen may fit either species,<br />
the locality of it lies within the known range of<br />
H. maltzami, but outside that of Solenolambrus noordendei,<br />
while the depth at which Odhner's specimen<br />
was collected lies well within the depth range<br />
of S. noordendei, but is much greater than at present<br />
known for tropical <strong>West</strong> <strong>African</strong> H. maltzami.<br />
BIOLOGY.—Monod (1956:593) gave the depth<br />
range of the species as "sublittoral, jusqu'a 400<br />
metres." Our specimens were obtained at depths<br />
of 33 to 70 m. All previous <strong>West</strong> <strong>African</strong> records<br />
also are from far less than 100 m depth: 18-28 m<br />
(Miers, 1881a: type-locality), 10 m, 20-22 m<br />
(Balss, 1921), 10, 14-15,15, 15, 15-20, 15-20, 16-<br />
18, 19, 19, 20, 20, 20, 22, 23-24, 25, 25, 27-28,<br />
30, 30, 31, 32, 32, 35, 36, 37, 38, 38, 40-41, 41,<br />
46-50, 50 m (Monod, 1956), 0-10, 3-17 m<br />
(Sourie, 1954b), 18-60 m (Longhurst, 1958), 19-<br />
37 m (Gauld, 1960), 5, 8, 11-12, 13, 16, 22 m<br />
(Guinot and Ribeiro, 1962), 6-8 m (Rossignol,<br />
1962), 8 m (Ribeiro, 1964), 18 m (Forest and<br />
Guinot, 1966), 30-40 m (Maurin, 1968b), 40,<br />
57 m (Tiirkay, 1975a). On the other hand all the<br />
records of the species from the Mediterranean,<br />
the Bay of Biscay, the Azores, and north of the<br />
Gape Verde Islands are much deeper: 100 to<br />
550 m. The <strong>West</strong> <strong>African</strong> specimens were found<br />
on the following types of bottom: shelly and<br />
muddy (Miers, 1881a), sand with Palythoa and<br />
Molgula (Monod, 1956), coarse shelly sand with<br />
Area and Pyura (Sourie, 1954b), muddy sand,<br />
shelly sand, and shelly mud (Longhurst, 1958),<br />
sand (Guinot and Ribeiro, 1962), mud and shells<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
(Forest and Guinot, 1966), and sand or shelly<br />
sand (Maurin, 1968b).<br />
DISTRIBUTION.—Heterocrypta maltzami has been<br />
reported from the Bay of Biscay, the Mediterranean<br />
(as far as the Adriatic Sea), the Azores, N of<br />
the Cape Verde Islands (16°55'-16°51'N, 27°27'-<br />
27°29'W of Paris = 25°07'-25°09'W of Greenwich),<br />
and from tropical <strong>West</strong> Africa. Monod<br />
(1956) summarized earlier records and reported<br />
material from Senegal, Ghana, and Gabon; records<br />
not in Monod include the following:<br />
Spanish Sahara: OffCabo Blanco, 20°43.5'N, 17° 10.8'W,<br />
40 m, and 2O°37.3'N, 17°24.4'W, 57 m (Tiirkay, 1975a).<br />
Mauritania: Bane d'Arguin, 30-40 m (Maurin, 1968b).<br />
Cape Verde Islands: Baia de Porto Grande, Sao Vicente<br />
(Guinot and Ribeiro, 1962; Ribeiro, 1964).<br />
Senegal: Anse de Hann and Anse Bernard near Dakar<br />
(Sourie, 1954b).<br />
Guinea: 09°40'N, 14°05'W, 18 m (Forest and Guinot,<br />
1966).<br />
Sierra Leone: No specific locality, 18-60 m (Longhurst,<br />
1958).<br />
Ghana: Off Accra, 18-37 m (Gauld, 1960).<br />
Congo: Baie de Pointe-Noire, 6-8 m (Rossignol, 1962).<br />
Angola: Baia da Caota, 11-12, 13, and 16m; Sombreiro,<br />
5 m; and Baia Farta, 22 m (all Benguela) (Guinot and<br />
Ribeiro, 1962).<br />
Genus Sakaila, new genus<br />
TYPE-SPECIES.—Sakaila africana, new species.<br />
ETYMOLOGY.—It is with great pleasure that we<br />
dedicate this genus to Professor Tune Sakai, the<br />
eminent Japanese carcinologist, who pointed out<br />
the differences between the species assigned here<br />
to the new genus and the species of Osachila sensu<br />
stricto, here restricted to the American and central<br />
Atlantic species (Rathbun, 1937:248). The<br />
gender of the generic name is feminine.<br />
DEFINITION.—Carapace broad, suboval in<br />
shape, narrowing anteriorly, regularly arcuate<br />
laterally, margins thin, scalloped or lobed. Front<br />
narrow, produced anteriorly, bilobed. Orbits<br />
small. Eyes scarcely or not at all visible in dorsal<br />
view. Antennules oblique. Antennae at inner angle<br />
of orbit. Buccal cavity broad anteriorly. Efferent<br />
branchial orifices separated, not meeting at<br />
midline. Merus of third maxilliped shorter than
NUMBER 306 325<br />
ischium, anterior margin of merus divided into 2<br />
lobes. Chelipeds symmetrical. Walking legs compressed,<br />
dorsally toothed or with large, irregular<br />
tubercles.<br />
REMARKS.—Sakaila can readily be distinguished<br />
from Osachila Stimpson, 1871, by the<br />
position of the orbits, the distinctly separated<br />
efferent branchial channels, by the ischium of the<br />
third maxillipeds being shorter than the merus<br />
and being notched anteriorly, and by the spined<br />
or tuberculate walking legs.<br />
Guinot (1966:754) noted "pour des raisons<br />
d'ordre taxonomique, la separation d'O. stimpsoni,<br />
0. japonica, et 0. imperialis dans un genre ou sousgenre<br />
nouveau pourrait se justifier car plusieurs<br />
caracteres concomitants les distinguent des autres<br />
Osachila." Guinot (1968b) referred to 0. stimpsoni<br />
[sensu Monod, 1956] as a primitive Osachila. As a<br />
result of her study of the affinities of Osachila<br />
sensu lato, she assigned it, Aethra, Hepatus, Hepatella,<br />
and Actaeomorpha to a group that she called<br />
"parthenoxystomienne," and tentatively assigned<br />
this group to the Parthenopinae (Guinot, 1967b:<br />
841). This group was retained in the parthenopids<br />
by Sakai (1976). Pending further studies, we retain<br />
Sakaila in the Parthenopidae, subfamily<br />
Aethrinae, following Sakai (1976:288).<br />
In addition to the type-species, Sakaila africana,<br />
new species, we assign two other species to this<br />
genus, both from Japanese waters: Osachila imperialis<br />
Sakai, 1963, and Osachila japonica Sakai, 1963<br />
(Sakai, 1963, 1965). Osachila expansa Takeda<br />
(1977), from the Ogasawara Islands, in which the<br />
maxillipeds are strongly narrowed anteriorly,<br />
may be retained in Osachila sensu stricto.<br />
* Sakaila africana, new species<br />
FIGURE 83<br />
Osachila stimpsoni.—Monod, 1956:100, 624, figs. 874-876.—<br />
Forest, 1959:15.—Forest and Guinot, 1966:51.—Guinot,<br />
1966:747, 750, 752-755, figs. 2, 6, 14; 1967b:828-830,<br />
832-838, figs. 26, 29, 32, 33; 1968b: 165, fig. 15 [discussion].<br />
[Not Osachila stimpsonii Studer, 1883.]<br />
MATERIAL EXAMINED.—Pillsbury Material: Annobon:<br />
Sta 284, 73 m, black basaltic rocks, 1
326<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
FIGURE 83.—Sakaila africana, new genus, new species, holotype, male cl 23 mm, Pillsbury Sta 284:<br />
a, carapace, dorsal view; b, carapace, ventral view; c, chela; d, fifth pereiopod; e, abdomen.<br />
buccal area shows correctly that the infraorbital<br />
tooth, which lies against the outer margin of the<br />
antennal peduncle, continues posteriorly and medially<br />
as a flattened lobe, which partly overlaps<br />
the area along the anterolateral margin of the<br />
mouthfield. In our specimen this lobe ends in the<br />
anterior and largest of the three deep pits in the<br />
subhepatic region. These pits are not shown in<br />
Guinot's figure.<br />
The outer surface of the merus of the cheliped<br />
(Figure 83r) is reticulate by the presence of shorter<br />
and longer ridges and tubercles, rather than being<br />
"granuleuse" as described by Monod. The upper<br />
surface of the merus, not described by Monod, is
NUMBER 306 327<br />
very narrow, widening somewhat anteriorly. The<br />
inner margin shows three high teeth, which are<br />
anteroposteriorly compressed; the outer margin<br />
shows a sharp high rim-like crest, the anterior<br />
part of which is highest and is fused with the<br />
anterior inner tooth to a high screen-like structure<br />
on the anterodorsal margin of the merus. The<br />
inner surface of the carpus has an anterodorsal<br />
and an anteroventral tooth connected with a<br />
ridge-like row of smaller teeth. The ridge on the<br />
inner surface of the palm bears a row of tubercles,<br />
one near the middle being high and sharp. The<br />
fingers have some of the tubercles rather strong,<br />
especially near the upper margin (in the dactylus),<br />
the lower margin (in the fixed finger), and<br />
near the base. The fingers of both chelae are<br />
inflected downward (Figure 83c), making a blunt<br />
angle with the longitudinal axis of the palm. In<br />
Monod's (1956) figure 875 the fingers are shown<br />
directed straight forward.<br />
Through the kindness of H.-E. Gruner, Zoologisches<br />
Museum, Berlin, we were able to compare<br />
our specimens with syntypes of Osachila stimpsonii<br />
Studer, 1883, from Ascension Island. Studer's<br />
species is correctly placed in Osachila. A redescription<br />
of 0. stimpsonii is in preparation by Manning<br />
in collaboration with Fenner A. Chace, Jr., in a<br />
report on the marine decapods of Ascension Island.<br />
TYPE-LOCALITY.—Annobon Island, 01°30'S,<br />
05°36'E, in 73 m.<br />
DISPOSITION OF TYPES.—The holotype (Crust.<br />
D. 31541), a male taken at Pillsbury Sta 284, is in<br />
the Rijksmuseum van Natuurlijke Historie, Leiden.<br />
The male paratype (USNM 139766) taken<br />
at Geronimo Sta 235 is in the National Museum of<br />
Natural History, <strong>Smithsonian</strong> <strong>Institution</strong>, Washington,<br />
D.C.<br />
BIOLOGY.—Sakaila africana has been collected in<br />
depths between 65-75 and 132 m. The type of<br />
bottom on which it was found has been indicated<br />
as mud and sand, 65 to 75 m (Forest and Guinot,<br />
1966) and black basaltic rocks, 73 m {Pillsbury).<br />
Ovigerous females were collected in July<br />
(Monod, 1956).<br />
ETYMOLOGY.—The specific epithet is derived<br />
from Latin and refers to the occurrence of the<br />
species off the <strong>West</strong> <strong>African</strong> coast.<br />
DISTRIBUTION.—Off tropical <strong>West</strong> Africa. It has<br />
not previously been recorded from either Gabon<br />
or Annobon. Records in the literature include the<br />
following:<br />
Senegal: S of Goree, 96 m; off Goree, 132 m (Monod,<br />
1956). 12°55.5'N, 17°33'W, 65-75 m (Forest, 1959; Forest<br />
and Guinot, 1966; Guinot, 1966, 1967b, 1968b).<br />
Subfamily PARTHENOPINAE MacLeay, 1838<br />
Genus Daldorfia Rathbun, 1904<br />
Parlhenope Fabricius, 1798:315, 352 [type-species: Cancer horridus<br />
Linnaeus, 1758, by selection by H. Milne Edwards,<br />
1838, in 1836-1844, pi. 26: fig. 2; a junior homonym of<br />
Parthenope Weber, 1795; gender: feminine; name 1679 on<br />
Official Index].<br />
Daldorfia Rathbun, 1904:171 [type-species: Cancer horridus<br />
Linnaeus, 1758, by monotypy; gender: feminine; name<br />
1582 on Official List].<br />
Daldorfia bouvieri (A. Milne Edwards, 1869)<br />
Parthenope bouvieri A. Milne Edwards, 1869:350.—Capart,<br />
1951:106, fig. 36 [Cape Verde Islands].—Monod, 1956:<br />
595, figs. 871, 872 [Cape Verde Islands; references].—<br />
Forest and Guinot, 1966:121 [Principe, Sao Tome].—<br />
Crosnier, 1967:340 [Congo]; 1969:535 [Congo].<br />
DISTRIBUTION.—<strong>West</strong> Africa, from the Cape<br />
Verde Islands and the Gulf of Guinea at the<br />
localities listed above, in 4-5 to 91 m.<br />
Genus Parthenope Weber, 1795<br />
Parthenope Weber, 1795:92 [type-species: Cancer longimanus<br />
Linnaeus, 1758, by subsequent designation by Rathbun,<br />
1904:171; gender: feminine; name 1581 on Official List].<br />
Lambrus Leach, 1815a:308 [type-species: Cancer longimanus<br />
Linnaeus, 1758, by monotypy; gender: masculine; name<br />
1678 on Official Index).<br />
Platylambnts Stimpson, 1871a: 129 [type-species: Lambrus crenulatus<br />
Saussure, 1858, a subjective junior synonym of<br />
Lambrus serratus H. Milne Edwards, 1834, by subsequent<br />
designation by Rathbun, 1925:516; gender: masculine].<br />
Aulacolambrus Paulson, 1875:9 [type-species: Lambrus pisoides<br />
Adams and White, 1848, by monotypy; gender: masculine].<br />
Pseudolambrus Paulson, 1875:9 [type-species: Parthenope calap-
328<br />
poides Adams and White, 1848, by monotypy; gender:<br />
masculine].<br />
Enoplolambrus A. Milne Edwards, 1878, in 1873-1881:147<br />
[type-species: Lambrus carenatus H. Milne Edwards, 1834,<br />
by monotypy; gender: masculine].<br />
Parthenolambrus A. Milne Edwards, 1878, in 1873-1881:148<br />
[type-species: Parthenope tarpeius Adams and White, 1849,<br />
by subsequent designation by Rathbun, 1925:528; gender:<br />
masculine].<br />
Rhinolambrus A. Milne Edwards, 1878, in 1873-1881:148<br />
[type-species: Cancer contrarius Herbst, 1804, by original<br />
designation; gender: masculine].<br />
Parthenopoides Miers, 1879a:672 [type-species: Lambrus massena<br />
Roux, 1830, by monotypy; gender: masculine].<br />
Oncodolambrus De Man, 1906:400 [type-species: Lambrus (Oncodolambrus)<br />
praedator De Man, 1906, by monotypy; gender:<br />
masculine].<br />
* Parthenope expansa (Miers, 1879)<br />
FIGURE 84<br />
Lambrus (Parthenopoides) expansus Miers, 1879b:25, pi. 5:<br />
fig. 9.<br />
Parthenolambrus expansus.—Adensamer, 1898:611 [Mediterranean].<br />
Lambrus expansus.—Monod, 1956:588 [references].—Holthuis<br />
and Gottlieb, 1958:119 [listed].—Forest and Guinot, 1966:<br />
120.<br />
Parthenope expansus.—Pastore, 1975:145, 147, figs. 1-3 [Mediterranean].<br />
MATERIAL EXAMINED.—Pillsbury Material: Annobon:<br />
Sta 283, 51-55 m, nodular coralline algae, 2$ (L, W).<br />
DESCRIPTION.—Carapace triangular in outline.<br />
Front bluntly rounded, its sides forming a straight<br />
line with sides of carapace. Orbits visible above<br />
as small cavities in lateral margin of carapace,<br />
but eyes, when retracted, exactly fill cavity and<br />
lateral margin appears unbroken. In posterior<br />
half, lateral margin slightly widened and showing<br />
3 shallow teeth just before posterolateral angle.<br />
Posterior margin slightly and evenly convex,<br />
showing few broad, inconspicuous lobes in median<br />
part. Dorsal surface of carapace uneven, but<br />
showing hardly any tubercles. Front somewhat<br />
concave, sunk in middle. Transverse broad ridge<br />
or elevated area present in metagastric region<br />
behind sunken area. Cervical groove indicated by<br />
wide depression. Oblique ridge present over each<br />
branchial region ending in posterolateral angle<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
and running parallel to lateral margin of carapace.<br />
Posterior part of carapace, between branchial<br />
ridges, bearing median and 2 submedian<br />
elevations, median strongest. Surface of carapace<br />
rather uniformly and finely pitted, with few tubercles<br />
visible on elevated portions, but not very<br />
distinct. General shape of carapace in many respects<br />
resembling more that of Heterocrypta than<br />
that of Parthenope.<br />
Eyes small, completely retractable in their orbits,<br />
visible in dorsal view.<br />
Antennules with basal segment very wide,<br />
reaching beyond antennal peduncle and forming<br />
greater part of the lower inner margin of orbit.<br />
Antennular sockets not sharply delimited distally,<br />
ending in wide groove in ventral surface of front,<br />
groove reaching margin of front, but less distinct<br />
distally than proximally.<br />
Basal antennal segment (Figure 84a) short, distinctly<br />
failing to reach orbit and separated from<br />
orbit by almost entire length of second segment.<br />
Outer angle of merus of third maxilliped somewhat<br />
triangularly produced laterally.<br />
Chelipeds markedly different, resembling those<br />
of Parthenope massena. Right cheliped heaviest,<br />
somewhat swollen. Upper surface of palm slightly<br />
convex, almost smooth; outer margin bearing<br />
ridge with 3 large blunt teeth, more pronounced<br />
in smaller than in larger cheliped. Inner margin<br />
of the upper surface of palm bearing about 5<br />
FIGURE 84.—Parthenope expansa (Miers), female, cl 7.5 mm,<br />
Pillsbury Sta 283: a, basal antennal segment; b, second pereiopod;<br />
c, fifth pereiopod.
NUMBER 306 329<br />
blunt teeth, middle largest. Outer surface of palm<br />
with some longitudinal rows of tubercles. Inner<br />
surface of palm smooth, slightly concave. Fingers<br />
somewhat more than half as long as palm, upper<br />
surface of dactylus bearing few large and several<br />
smaller tubercles proximally. Cutting edge of<br />
fixed finger of large cheliped with single large<br />
molariform tooth, occupying greater part of edge,<br />
flanked by small denticle distally. Cutting edge<br />
of fixed finger in small chela with 3 large distal<br />
and 3 small proximal teeth. Edges of dactylus of<br />
both chelipeds bearing 3 or 4 low small teeth.<br />
Carpus short, cup-shaped, with small irregular<br />
tubercles, forming more or less distinct longitudinal<br />
rows. Merus short and wide, inner margin<br />
bearing distinct larger and smaller teeth, on outer<br />
margin teeth less conspicuous. Lower margin of<br />
merus with longitudinal row of large tubercles,<br />
inner surface bearing some scattered tubercles,<br />
outer surface almost smooth.<br />
In following legs (Figure 84b,c) lower surface of<br />
merus bearing 2 rows of granules, 1 on either<br />
margin; upper margin of merus armed with tubercles<br />
in extreme proximal part only. Carpus<br />
with lower surface unarmed, upper carrying angular<br />
tubercle; in addition, last leg with few distal<br />
tubercles on upper margin. Propodus with some<br />
ventral and dorsal denticles; denticles and tubercles<br />
far more distinct in posterior than in anterior<br />
legs. Dactylus of second pereiopod (= first walking<br />
leg) about as long as, but narrower than,<br />
propodus, somewhat longer than carpus; merus<br />
about twice as long as, and broader than, carpus.<br />
Third and fourth pereiopods similar to second.<br />
Fifth pereiopod (Figure 84c) shortest of all legs.<br />
Dactylus longer than, but half as high as, propodus;<br />
carpus about as long as propodus, and half<br />
or less than half as long as merus.<br />
Female abdomen with all somites free.<br />
MEASUREMENTS.—The two examined females<br />
have cl 6 and 7.5 mm and cb 7 and 10 mm. The<br />
holotype had cb 11 mm (Miers, 1879b). The<br />
measurements given in the literature are: cl 10<br />
mm, cb 11 mm (Miers, 1886); cl 8 mm, cb 10<br />
mm (A. Milne Edwards and Bouvier, 1894); for<br />
123 cl 4.0-9.5 mm, cb 5.0-12.0 mm, for 2 oviger-<br />
ous 9 cl 5.0 and 6.0 mm, cb 5.5 and 7.5 mm, and<br />
2 non-ovigerous $ cl 5.0 and 6.5 mm, cb 5.8 and<br />
7.5 mm (Pastore, 1975:150; in Pastore's table,<br />
length and width have evidently been interchanged<br />
by accident). The known size range for<br />
the species thus is cl 4-10 mm, cb 5-12 mm.<br />
REMARKS.—Capart (1951:106, fig. 36) described<br />
and figured a female of what he considered<br />
to be a juvenile Daldorfia bouvieri (A. Milne<br />
Edwards). This specimen (cl 9 mm, cb 12 mm)<br />
originated from Boa Vista, Cape Verde Islands.<br />
The description and figure given by Capart show<br />
a great similarity to the present species, and a<br />
reexamination of Capart's specimen might be of<br />
interest. Our specimens differ from that described<br />
by Capart in having the carapace uniformly and<br />
evenly minutely pitted and not eroded. Capart's<br />
specimen might be abnormal in having two equal<br />
chelipeds, both corresponding with the smaller<br />
cheliped in the present species.<br />
BIOLOGY.—The species has been found in<br />
depths between 30 and 185 m. The bottom at the<br />
localities where it was found was noted in some<br />
instances: gravel, sand and broken shell (A. Milne<br />
Edwards and Bouvier, 1894, 1899); coral, rock<br />
and sand, sand and rock, sand, shell and coral,<br />
sand and shell (A. Milne Edwards and Bouvier,<br />
1900); calcareous and other algae (Forest and<br />
Guinot, 1966); nullipores and coarse sand (Adensamer,<br />
1898).<br />
Ovigerous females have been collected in June<br />
and August (Monod, 1956; Pastore, 1975).<br />
DISTRIBUTION.—Eastern Atlantic, from the<br />
Mediterranean, Madeira, Seine Seamount, the<br />
Azores, the Canary Islands, and <strong>West</strong> Africa from<br />
Spanish Sahara to Sao Tome and Annobon islands<br />
in the Gulf of Guinea. This species has not<br />
been recorded previously from Annobon. The<br />
Mediterranean records are: NW of Crete, 36°-<br />
03'N, 23°06'E (Adensamer, 1898; Pastore, 1975)<br />
and Acitrezza, Bay of Catania, Sicily (Pastore,<br />
1975). Monod (1956) summarized earlier records<br />
and reported material from Madeira, the Canary<br />
Islands, and the Cape Verde Islands. <strong>West</strong> <strong>African</strong><br />
records since 1956 include the following:
330<br />
Canary Islands: No specific locality (Pastore, 1975).<br />
Sao Tome: OO°25'4O"N, 06°40'10"E, 50 m (Forest and<br />
Guinot, 1966).<br />
* Parthenope massena (Roux, 1830)<br />
?Lambrus massena.—Capart, 1951:105, fig. 35.<br />
Lambrus massena. —Sourie, 1954b: 147, 150.—Monod, 1956:<br />
572, 632, figs. 840-856.—Gauld, 1960:72.—Rossignol,<br />
1962:123.—Crosnier, 1964:31.—Forest and Guinot, 1966:<br />
118— Zariquiey Alvarez, 1968:441, fig. 147 [Spain; references].<br />
Lambrus sp.—Monod, 1956:583, figs. 857, 858.<br />
Lambrus massenae.—Longhurst, 1958:89 [erroneous spelling].<br />
SYNONYMS.—PParthenope contracta Costa, 1840;<br />
?Parthenope hexacantha Costa, 1840; Lambrus pumilus<br />
Costa, 1851; Lambrus rugosus Stimpson, 1857; Lambrus<br />
setubalensis De Brito Capello, 1866; Lambrus<br />
pulchellus A. Milne Edwards, 1868; Lambrus massena<br />
var. atlanticus Miers, 1881; Lambrus massena<br />
var. spinifer Miers, 1881; Lambrus massena var.<br />
goreensis Miers, 1881; Lambrus bicarinatus Miers,<br />
1881.<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 70, 33 m, branched Foraminifera, 2$ (L).<br />
Ghana: Sta 22, 51 m, rough bottom, 2c5, 1 juv (L). Sta 23,<br />
42 m, foliate brown to orange bryozoans, 1
NUMBER 306 331<br />
material reported upon here was taken between<br />
24 and 51 m. The species was taken on the<br />
following types of bottom: sand with Palythoa and<br />
Molgula (Monod, 1956); coarse shelly sand, bottom<br />
with Area and Pyura (Sourie, 1954b); sand<br />
and shells, sand and mud (Longhurst, 1958); mud<br />
and shells (Longhurst, 1958; Forest and Guinot,<br />
1966); sand with Foraminifera on rocky bottom<br />
with gorgonians (Crosnier, 1964); rock and shells,<br />
mud, calcareous algae and shells, calcareous algae,<br />
calcareous algae, sand and shells, sand and<br />
calcareous and other algae, mud and calcareous<br />
algae (Forest and Guinot, 1966).<br />
Ovigerous females have been recorded in all<br />
months but January, April, and August, suggesting<br />
that off <strong>West</strong> Africa the species spawns all<br />
year (Monod, 1956; Forest and Guinot, 1966;<br />
Pillsbury).<br />
DISTRIBUTION.—Eastern Atlantic, from Brittany,<br />
Atlantic coast of France, southward to the<br />
Congo, including the Mediterranean. Monod<br />
(1956) summarized the literature and reported<br />
material from Senegal, Guinea, Sierra Leone,<br />
Ghana and Principe. Other records in the literature<br />
include the following:<br />
Senegal: Anse Bernard and Anse de Hann, Baie de Dakar,<br />
0-17 m (Sourie, 1954b).<br />
Sierra Leone: No specific locality, in 10-106 m (Longhurst,<br />
1958).<br />
Ghana: Off Accra, 20-40 m (Gauld, 1960). 04°37'N,<br />
00°50'W, 90-100 m (Forest and Guinot, 1966).<br />
Nigeria: Off the mouths of the Niger River, 04°03'N,<br />
06°12'E, 32 m (Forest and Guinot, 1966).<br />
Cameroon: No specific locality, in 10-50 m (Crosnier,<br />
1964).<br />
Principe: 01°38'25"N, O7°22'O5"E, 31 m; 01°43'10"N,<br />
07°28'20"E, 73 m;.01°43'N, 07°28'55"E, 37 m; [Cais de]<br />
Santana, 11 m; Praia Grande, 3-12 m (all Forest and Guinot,<br />
1966).<br />
Sao Tome: 00°25'15"N, 06°43'05"E, 8-30 m; Baia de<br />
Ana de Chaves, 5 m (Forest and Guinot, 1966).<br />
Annobon: 01°27.5'S, 05°36.5'E, 35 m (Forest and Guinot,<br />
1966).<br />
Gabon: W of Pointe Gombe, 40 m (Rossignol, 1962).<br />
Congo: W of Pointe-Noire, 10 m (Rossignol, 1962).<br />
Parthenope miersii (A. Milne Edwards and<br />
Bouvier, 1898)<br />
Lambrus miersi— Capart, 1951:105 [discussion].—Monod,<br />
1956:583 [references].<br />
Parthenope miersi.—Zariquiey Alvarez, 1968:439 [Spain; references].<br />
DISTRIBUTION.—Eastern Atlantic, from Portugal,<br />
the Bay of Cadiz, Seine Seamount, and the<br />
Cape Verde Islands; sublittoral, in depths between<br />
91 and 240 m.<br />
* Parthenope notialis, new species<br />
FIGURES 85, 86a,*<br />
Lambrus mediterraneus.—Studer, 1882:335 [not Lambrus mediterraneus<br />
Roux, 1828 = Cancer macrochelos Herbst, 1790].<br />
Lambrus Mediterraneus.—Studer, 1883:9.<br />
Lambrus macrochelos.— Rathbun, 1900a:295.—Monod, 1956:<br />
585, figs. 859-861.—Longhurst, 1958:89.—Gauld, 1960:<br />
72.—Guinot and Ribeiro, 1962:80.—Rossignol, 1962:<br />
123.—Crosnier, 1964:34. [Not Cancer macrochelos Herbst,<br />
1790.]<br />
Lambrus macrocheles.— Doflein, 1904:87 [part], pi. 32: fig. 5.—<br />
Balss, 1921:54.—Odhner, 1923:20.—Capart, 1951:102,<br />
fig. 34, pi. 2: figs. 5, 6.—Forest and Guinot, 1966:119.—<br />
?Maurin, 1968a:59, 62; 1968b:480, 486, 489.—Crosnier,<br />
1970:1215 [listed], 1219. [Not Cancer macrochelos Herbst,<br />
1790.]<br />
Lambrus (Lambrus) macrocheles.—Monod, 1933b:498 [not Cancer<br />
macrochelos Herbst, 1790].<br />
Parthenope.—Vass, 1966:19, 22.<br />
Lambrus.— ?Maurin, 1968b, fig. 9.<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia:<br />
Sta 68, 70 m, broken shell, 66, 12? (2 ov) (L).<br />
Ivory Coast: Sta 42, 62-75 m, mud with brown branched<br />
Foraminifera, 66*, 4$ (W). Sta 46, 38-42 m, mud with dense<br />
Jullienella, 1(5, 29 (W). Sta 50, 128-192 m, 1$ ov (L). Sta 59,<br />
55-64 m, mud with dense branched Foraminifera, 1
332<br />
it was not distinguished by the authors that have<br />
dealt with it. We found, however, a number of<br />
characters that made it impossible to consider the<br />
present <strong>West</strong> <strong>African</strong> specimens and typical Med-<br />
iterranean specimens of P. macrochelos as belonging<br />
to the same species or subspecies.<br />
The shape of the carapace (Figure 85), which<br />
in adult specimens is distinctly wider than long<br />
(proportion length:width being about 5:6), has<br />
the same general shape as in P. macrochelos. The<br />
rostrum is narrow and bears a tooth at either side.<br />
The inner orbital angle bears two teeth and a<br />
strong tubercle. The upper margin of the orbit is<br />
provided with a strong tubercle (Figure 86a),<br />
behind which sometimes a trace of a smaller<br />
tubercle may be visible; in P. macrochelos (Figure<br />
86c) this second tubercle is quite distinct. Some<br />
distance behind the orbits, in the median area of<br />
the mesogastric region, there are four tubercles<br />
placed in a quadrangle. The anterior pair stands<br />
close to the posterior pair and its tubercles are<br />
placed slightly wider apart than those of the<br />
posterior pair. The four tubercles thus form a<br />
trapezium with the widest margin anteriorly. In<br />
P. macrochelos, the posterior tubercles are wider<br />
apart than the anterior, so that the trapezium<br />
formed by the four tubercles has the narrowest<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
margin anteriorly; in P. macrochelos the two pairs<br />
of tubercles are separated from each other by a<br />
greater distance than in P. notialis. Behind the<br />
group of four tubercles there is a strong median<br />
gastric spine. Behind the cervical groove there is<br />
a median row of 3 strong spines (1 urogastric and<br />
2 cardiac); the posterior of these is the largest. In<br />
P. notialis these spines are relatively longer than<br />
in P. macrochelos; the arrangement of the spines is<br />
the same in the two species. The median gastric<br />
region is swollen and so is the branchial region.<br />
In the depressed area between these two swollen<br />
regions there is a row of three blunt tubercles<br />
close to and parallel with the median row of<br />
spines. The branchial region ends posterolaterally<br />
in three large teeth, between which there are<br />
small tubercles. In P. macrochelos, instead of these<br />
tubercles, there are teeth that often are only<br />
slightly smaller than the large teeth. The middle<br />
of the three large teeth is placed at the posterior<br />
end of a blunt oblique ridge, which carries another<br />
spine more anteriorly. The anterolateral<br />
margin of the carapace bears a row of about 7<br />
teeth, which are distinctly shorter and narrower<br />
than the outer posterolateral tooth; in P. macrochelos<br />
these anterolateral teeth are much larger,<br />
reaching almost the size of the outer posterolat-<br />
FIGURE 85.—Parthmope notialis, new species (from Monod, 1956, fig. 859).
NUMBER 306 333<br />
FIGURE 86.—Parthenope notialis, new species, paratype, male,<br />
cl 17.5 mm, Pillsbury Sta 260: a, front; b, abdomen. Parthenope<br />
macrochelos (Herbst), male, cl 32.8 mm, Naples: c, front; d,<br />
abdomen.<br />
eral. The general shape and tuberculation of the<br />
carapace of P. notialis is very similar to that of P.<br />
macrochelos.<br />
The epistome, oral field, and third maxilliped<br />
are like in P. macrochelos, only the spines are<br />
sharper and more distinct. The subhepatic region<br />
just lateral of the oral cavity is smooth in P.<br />
notialis, uniformly tuberculated in P. macrochelos.<br />
In P. notialis the chelipeds of the adult males<br />
are relatively distinctly longer than in P. macrochelos.<br />
The second pereiopod in P. notialis does not<br />
attain the end of the merus of the chela, while in<br />
P. macrochelos it reaches beyond. The fingers of the<br />
chelipeds in P. notialis are more laterally compressed<br />
than in P. macrochelos, where they are more<br />
cylindrical. The teeth on the palm in P. notialis<br />
also are more compressed and have the margins<br />
with small serrations, sometimes small tubercles<br />
may be seen on the lateral surfaces of these teeth.<br />
In P. macrochelos the teeth on the palm are more<br />
**<br />
conical and have spinules on all sides. The upper<br />
surface of the palm of the chelipeds in P. notialis<br />
is almost smooth, only a few indistinct tubercles<br />
are seen; in P. macrochelos this surface is very rough<br />
with many spines and tubercles. The lower outer<br />
surface of the palm is slightly convex with a few<br />
rows of small tubercles, of which the median is<br />
the most conspicuous. In P. macrochelos this surface<br />
is more convex and the tubercles are larger. In P.<br />
notialis the merus of the cheliped has the slightly<br />
convex upper surface almost smooth, except for<br />
a median row of spinules; in P. macrochelos the<br />
surface is more uniformly spinuliferous. The<br />
spines on the margins of the upper surface of the<br />
merus are more flattened and less numerous in P.<br />
notialis than in P. macrochelos.<br />
The following pereiopods are very similar in P.<br />
notialis and P. macrochelos. The dactylus is covered<br />
with a very short felt-like pubescence; it is practically<br />
as long as the propodus in the second<br />
pereiopod, becoming gradually relatively longer<br />
in the following legs. The carpus also is about as<br />
long as the propodus in pereiopod 2, but it becomes<br />
gradually shorter in the following legs. The<br />
merus is the longest segment and is as long as<br />
propodus and carpus combined. A row of spinules<br />
is present on the upper and the lower margins of<br />
the merus, and on the upper margin of the carpus<br />
and propodus of all legs. The spinules are distinct<br />
on the merus, faint to very faint on the carpus<br />
and propodus; in the posterior legs they are more<br />
distinct than in the anterior.<br />
The shape and armament of the male thoracic<br />
sternum and abdomen is about the same in the<br />
two species. The second abdominal segment<br />
shows one median and two lateral teeth with<br />
tubercles in between; in P. notialis (Figure 86A)<br />
these teeth are large and bluntly lobiform, flattened<br />
and higher than wide, the tubercles are<br />
few, small and inconspicuous; in P. macrochelos<br />
(Figure 86W) the teeth are placed wider apart, are<br />
less flattened, lower and more triangular, while<br />
the tubercles are distinct and rather large. The<br />
general shape of the abdomen is the same in the<br />
two species, but the last somite is narrower in P.<br />
notialis. The third to fifth somites of the abdomen
334<br />
are fused: sutures rather than articulations indicate<br />
the lines between the somites.<br />
Figures: Figures of the whole animal were published<br />
by Doflein (1904, pi. 32: fig. 5), Capart<br />
(1951, fig. 34), and Monod (1956, fig. 859).<br />
Male Pleopod: Figures of the male pleopods are<br />
provided by Capart (1951, pi. 2: figs. 5, 6) (Zaire)<br />
and Monod (1956, figs. 860-861) (Senegal). They<br />
do not essentially differ from those of P. macrochelos.<br />
MEASUREMENTS.—The carapace length of the<br />
specimens seen by us varies between 5 and 21<br />
mm, that of the ovigerous females between 14<br />
and 18 mm. Odhner's (1923) specimen had the<br />
carapace width between 11 and 19 mm. Both<br />
Capart's (1951) largest male and largest female<br />
had the carapace 18.5 mm long and 21 mm wide.<br />
Monod (1956:585, 586) gave the following measurements<br />
for six of his males: cl 12 to 18 mm, cb<br />
13 to 23 mm; and for three ovigerous females: cl<br />
10 to 14 mm, cb 11 to 16 mm. The single specimen<br />
reported upon by Guinot and Ribeiro (1962:80)<br />
was a male with the carapace length and width<br />
both 5 mm. It is clear that the present species is<br />
a small one compared to P. macrochelos, which<br />
may attain a carapace length of 39 mm. The eggs<br />
of P. notialis are numerous and small, they are<br />
spherical and are 0.3 to 0.35 mm in diameter.<br />
Zariquiey Alvarez (1968:441), in citing ovigerous<br />
females of P. macrochelos with cl 10 mm, based his<br />
information on Monod's (1956) data of the present<br />
new species.<br />
REMARKS.—Parthenope notialis very strongly resembles<br />
P. macrochelos, and it is not surprising that<br />
the two species have always been confused. They<br />
may finally prove to be only subspecifically distinct.<br />
Our extensive material of the present species,<br />
however, differs consistently from the specimens<br />
of P. macrochelos, with which we could compare<br />
it.<br />
So far as we could ascertain all previous records<br />
of/*, macrochelos from tropical <strong>West</strong> Africa pertain<br />
to the present species.<br />
Studer (1883), in discussing his Liberian specimens<br />
(males and females), described the upper<br />
surface of the palm of the cheliped (which he<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
inadvertently named "Carpus") as "fast glatt,<br />
nur mit wenig Hockern besetzt" and also described<br />
the outer lower surface of the palm with<br />
its median row of tubercles, and remarked that<br />
in these features his specimens differed from Mediterranean<br />
P. macrochelos. This clearly proves the<br />
identity of his material with the present species.<br />
Doflein (1904:87) mentioned two lots of"Lambrus<br />
macrocheles" from <strong>West</strong> Africa. One of these<br />
consisted of a single damaged male from Seine<br />
Seamount, NE of Madeira at 33°43.8'N, 14°<br />
20'W, ca. 150 m deep. This male might well<br />
belong to Parthenope massena (Roux). Doflein remarked<br />
that it had "auffallend kiirzere Scheren<br />
mit dunkel-rotbraunen Fingerenden," which<br />
would fit P. massena quite well, as we examined<br />
numerous specimens of that species which, after<br />
long preservation, still showed distinct reddish<br />
brown finger tips. Bouvier (1922:76, pi. 2: fig. 3),<br />
under the name Lambrus Miersi, described and<br />
figured (in color) P. massena, remarking that "le<br />
bout des doigts des pinces [sont] d'un brun noiratre,"<br />
which color also is clearly shown on the<br />
figure. Doflein's specimen had the rostrum broken<br />
("dessen Rostrum abgebrochen ist"), and thus<br />
lacked a character that undoubtedly would have<br />
made Doflein recognize the true identity of the<br />
species.<br />
Doflein's second lot came from off the mouth<br />
of the Congo River and the male figured by him<br />
(1904, pi. 32: fig. 5) clearly belongs to the present<br />
species.<br />
Odhner (1923) considered his specimens to be<br />
juveniles of P. macrochelos as none had a carapace<br />
width of more than 19 mm; it is more probable<br />
that his specimens are P. notialis.<br />
Capart's (1951:102, fig. 34) illustration of a<br />
female from off Angola identified as "Lambrus<br />
macrocheles" clearly shows that it belongs to P.<br />
notialis as probably does also Capart's other material,<br />
since he remarked that the specimens in his<br />
collection "ne montrent pas entre eux de variations<br />
notables." As shown above the measurements<br />
of Capart's specimens fall entirely within<br />
the range that we found for the present species.<br />
Monod (1956:585, figs. 859-861) gave excellent
NUMBER 306 335<br />
figures of the present species, while the measurements<br />
given by him check well with those of P.<br />
notialis (p. 334). Of one of his lots (from S of Cap<br />
Vert) Monod stated specially "petite forme, face<br />
superieure des chelipedes plus ou moins lisses<br />
entre les cretes." There is thus no indication that<br />
any of his specimens do not belong to P. notialis.<br />
Longhurst's (1958) and Gauld's (1960) specimens<br />
were identified by Monod.<br />
We examined a specimen from Guinea-Bissau<br />
reported upon by Forest and Guinot (1966) as P.<br />
macrochelos and found that it belongs to P. notialis.<br />
The characters used in various keys (Bouvier,<br />
1940:309, 310; Monod, 1956:572; Zariquiey Alvarez,<br />
1968:437, 438) to distinguish between P.<br />
macrochelos and P. miersii (A. Milne Edwards and<br />
Bouvier), also serve to distinguish P. macrochelos<br />
from P. notialis. This notwithstanding, we believe<br />
that P. notialis is closer to P. macrochelos than to P.<br />
miersii. Parthenope miersii has the carapace narrower<br />
than in either P. macrochelos or P. notialis<br />
and the chelipeds are shorter. Parthenope verrucosa<br />
(Studer, 1883) from Ascension has the rostrum<br />
similar to that of P. macrochelos and P. notialis, but<br />
its chelipeds are shorter than in these two species,<br />
and the tuberculation of the carapace seems to<br />
be different. Additional information on both P.<br />
miersii and P. verrucosa is badly needed; material<br />
of neither species was available to us.<br />
TYPE-LOCALITY.—The male holotype was collected<br />
at Pillsbury Sta 260, off Cameroon, 03°45'N,<br />
09°05'E to 03°43'N, 09° 10'E, depth 46 m.<br />
DISPOSITION OF TYPES.—The holotype (Crust.<br />
D. 31545) is deposited in the collection of the<br />
Rijksmuseum van Natuurlijke Historie, Leiden.<br />
Some of the paratypes are in the Leiden Museum<br />
and some in the National Museum of Natural<br />
History, <strong>Smithsonian</strong> <strong>Institution</strong>, Washington,<br />
DC.<br />
ETYMOLOGY.—The specific epithet is from the<br />
Latin notialis (southern).<br />
BIOLOGY.—The species has been reported from<br />
depths between 18 and 162 m (one of Maurin's<br />
uncertain records is from 300 to 500 m); however,<br />
more than 80% of the records are from depths<br />
between 30 and 110 m. The species usually is<br />
found on bottoms of mud, sandy mud or sand,<br />
almost always mixed with broken shells, bryozoans,<br />
branched or foliate Foraminifera, corals<br />
or rocks. The records in the literature are the<br />
following: shelly mud (Longhurst, 1958; Forest<br />
and Guinot, 1966); sand, mud, coral and rock;<br />
sand, brown mud and coral; sand, mud and rock<br />
(Capart, 1951); sand, mud and shells; mud, shells<br />
and Cidaris; mud, stones, calcareous algae, sand<br />
and Foraminifera; gravel, shells and Foraminifera<br />
(Forest and Guinot, 1966); sandy mud with shells;<br />
mud with dense Jullienella (Voss, 1966); bottom<br />
with Jullienella foetida Schlumberger (Monod,<br />
1956); sand and broken shells; sand, clayish sand<br />
and broken shells (Odhner, 1923); rock and shells<br />
(Forest and Guinot, 1966); sand and mud or<br />
sandy mud (Capart, 1951; Guinot and Ribeiro,<br />
1962; Forest and Guinot, 1966); and mud (Capart,<br />
1951; Forest and Guinot, 1966). The present<br />
material has been taken on similar types of bottom.<br />
Ovigerous females were observed in February,<br />
May, July, and September (Monod, 1956),<br />
March (Crosnier, 1970), May (Forest and Guinot,<br />
1966), May and June (present paper), November<br />
(Capart, 1951).<br />
DISTRIBUTION.—Most of the records of P. macrochelos<br />
from tropical <strong>West</strong> Africa are based on the<br />
present species. Whether Maurin's (1968a, 1968b)<br />
material from off Spanish Sahara and Mauritania<br />
belongs here, to P. macrochelos, or to P. miersii is<br />
not clear; this can only be decided after reexamination<br />
of his material. The range of the present<br />
species extends at least from Senegal to Angola,<br />
but it might go north as far as the Spanish<br />
Sahara. If we include Maurin's material in the<br />
present species, the published records for it are<br />
the following:<br />
Spanish Sahara: Medano de Aaiiin and W of Cabo<br />
Bojador, 300-500 m (Maurin, 1968b).<br />
Mauritania: Bane d'Arguin, 40-60 m (Maurin, 1968b),<br />
and 90-100 m (Maurin, 1968a,b). Tamzak, 70-75 m<br />
(Maurin, 1968a, 1968b).<br />
Senegal: Off Saint-Louis, 100-300 m; off Kayar; S of<br />
Cap Vert, 97-98 m; SE of lie de la Madeleine, 48 m; off<br />
Dakar, 140 m; near Goree, 40-41 m, 50 m, % m, 132 m (all
336<br />
Monod, 1956). 13°01'N, 17°24'W, 51-55 m; 12°55.5'N,<br />
17°33'W, 65-75 m (Forest and Guinot, 1966).<br />
Guinea-Bissau: 10°19'N, 16°34'W, 60-73 m (Forest and<br />
Guinot, 1966).<br />
Guinea: 09°40'N, 14°05'W, 18 m (Forest and Guinot,<br />
1966).<br />
Sierra Leone: No specific locality, 54-106 m (Longhurst,<br />
1958).<br />
Liberia: 04°40'N, 09°40.6'W, 90 m (Studer, 1882, 1883).<br />
04°34.5'N, 08°31'W, 64 m (Forest and Guinot, 1966).<br />
Ivory Coast: O5°O7'N, 04°32'W to O5°O7'N, 04°36'W,<br />
38-42 m, and 04°35'N, 06°40'W to 04°35'N, 06°41'W, 64<br />
m (Voss, 1966).<br />
Ghana: 04°40'N, 02°08'W to 04°39'N, 02°05'W, 50 m;<br />
04°36.5'N, 01°31'W, 50 m; 04°37'N, 00°50'W, 90-100 m<br />
(Forest and Guinot, 1966). Accra, 43 m, 44 m, 65 m (Monod,<br />
1956; Gauld, 1960).<br />
Cameroon: No specific locality, depth more than 50 m<br />
(Crosnier, 1964).<br />
Gabon: 00°25'N, 09°00'E, 73 m (Forest and Guinot,<br />
1966). Off Pointe Banda, 03°57.5'S, 10°36.5'E, 85 m (Capart,<br />
1951).<br />
Cabinda: <strong>West</strong> of Landana, 50-65 m (Rossignol, 1962).<br />
Angola: Off Moita Seca, 06° 16'S, 12°O7'E, 50 m; 06°21'S,<br />
H°53'12"E, 100 m (Capart, 1951). Off the mouth of the<br />
Congo River, 06°18.7'S, 12°02.1'E, 44 m (Donein, 1904).<br />
09°47'S, 13° 1 l'E, 30-35 m; 09°40'S, 13°02'E, 80 m (Capart,<br />
1951). Baia Farta, Benguela, 22-28 m (Guinot and Ribeiro,<br />
1962). Baia dos Elefantes, 13°05'S, 12°46'E, 100-110 m;<br />
13°05'S, 12°45'E, 100-110 m; Baia de Salinas, 14°05'S,<br />
12°17'E, 110 m (Capart, 1951). Off Porto Alexandre, 72 m,<br />
108 m (Odhner, 1923). 16°37'S, 11°22'E, 126 m; 16°41'S,<br />
11°21'E, 162 m (Crosnier, 1970).<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
(L). Sta 49, 73-77 m, 26, 5? (4 ov) (L). Sta 50, 128-192 m,<br />
26, 1? (W). Sta 59, 55-64 m, mud with dense branched<br />
Foraminifera, Id (L). Sta 60, 79-82 m, coral or rock, lcJ (L).<br />
Nigeria: Sta 237, 101 m, 49 (1 ov) (W).<br />
Undaunted Material: Angola: Sta 96, 162 m, 19 ov (L).<br />
Other Material: Congo: Off Pointe-Noire, 04°56'S, 11°-<br />
31'E, 95 m, muddy sand, 3 Sep 1965, A. Crosnier, Id (W).<br />
DESCRIPTION.—Capart, 1951:108.<br />
Figures: Capart, 1951, fig. 37; Monod, 1956,<br />
593, figs. 868-870.<br />
Male Pleopod: Capart, 1951, pi. 2: figs. 14, 15<br />
(Zaire).<br />
MEASUREMENTS.—The ovigerous females in the<br />
present collection had the carapace length 6 to 9<br />
mm. Capart (1951) mentioned ovigerous females<br />
with cl 7 and 10 mm, cb 7 and 11 mm; Monod's<br />
(1956) ovigerous females had cl 7 to 9 mm and<br />
cb 8 to 10 mm. The carapace length in the rest of<br />
our material varies between 6 and 12 mm. Capart's<br />
(1951) largest specimen had cl 11 and cb<br />
11.5 mm; Monod's (1956) smallest and largest<br />
male had cl 5 and 11.5 and cb 4.5 and 12 mm,<br />
respectively, while Crosnier's (1967) males had cl<br />
6.6 to 8.7 mm and cb 7.0 to 9.1 mm.<br />
BIOLOGY.—The species is known from depths<br />
between 64 and 215 m; the shallowest record is<br />
55-64 m (present material), the deepest 215-220<br />
m (Capart, 1951; Monod, 1956). Of the records<br />
90% are from between 70 and 140 m. It has been<br />
reported from mud (Capart, 1951; Crosnier,<br />
Genus Solenolambrus Stimpson, 1871 1967), sandy mud (Capart, 1951; Crosnier, 1967),<br />
Solcnolambrus Stimpson, 1871a: 132 [type-species: Solenolambrus and shelly mud (Longhurst, 1958).<br />
typicus Stimpson, 1871, by use of typicus; gender: mascu- Ovigerous females have been found in the<br />
line].<br />
months of February, March, May, July, and No-<br />
Pisolambrus A. Milne Edwards, 1878, in 1873-1881:157 [typevember (Capart, 1951; Monod, 1956; Crosnier,<br />
species: Pisolambrus nitidus A. Milne Edwards, 1878, by<br />
1970; present paper).<br />
monotypy; gender: masculine].<br />
DISTRIBUTION.—Tropical <strong>West</strong> Africa, from localities<br />
between Senegal and Angola. It has not<br />
* Solenolambrus noordendei (Capart, 1951)<br />
been recorded previously from Liberia or Nigeria,<br />
Heterocrypta noordendei Capart, 1951:108, fig. 37, pi. 2: figs. 14, but these records are well within its known range.<br />
15.<br />
Records since Monod's (1956) include the follow-<br />
Solenolambrus noordendei.—Monod, 1956:593, figs. 868-870. ing:<br />
Longhurst, 1958:89.—Gauld, 1960:72.—Crosnier, 1967:<br />
340; 1970:1215 [listed], 1219.<br />
Sierra Leone: No specific locality, 72 m (Longhurst, 1958).<br />
Ivory Coast: No specific locality (Crosnier, 1967).<br />
MATERIAL EXAMINED.—Pillsbury Material: Liberia: Ghana: Off Accra, 80 m (Gauld, 1960).<br />
Sta 68, 70 m, broken shell, 1
RV Pillsbury COLLECTIONS<br />
Collections made in the Gulf of Guinea in 1964<br />
and 1965 (data from Bayer, 1966; OT = otter<br />
trawl).<br />
5<br />
Appendix I: Station Data<br />
1. Nigeria. Lagos harbor, 06°28'N, 03°23'E, shore collecting,<br />
23 May 1964: Callinectes marginatus, Cyclograpsus<br />
integer, Geograpsus lividus, Goniopsis pelii, Metagrapsus curvatus,<br />
Pachygrapsus gracilis, Panopeus africanus, Sesarma<br />
(Chiromantes) buettikoferi, Uca tangeri<br />
2. Nigeria. Lagos harbor, 06°28'N, O3°23'E, dipnet at<br />
surface, outgoing tide, 23 May 1964: Callinectes amnicola,<br />
Callinectes pallidus<br />
16. Ghana. 05°40'N, 00°30'E to 05°40'N, 00° 17'E, 46 m,<br />
mud with Foraminifera, shells, 6' OT, 26 May 1964:<br />
Capartiella longipes, Eurynome parvirostris, Ilia spinosa, Inachus<br />
angolensis, Inachus nanus, Machaerus oxyacantha, Macropodia<br />
gilsoni, Macropodia spinulosa, Medorippe lanata<br />
17. Ghana. 05°35'N, 00°10'E to 05°36'N, 00°11.5'E, 48<br />
m, fine sand and green mud, 40' OT, 26 May 1964:<br />
Achaeus buderes, Achaeus foresti, Calappa pelii, Calyps-<br />
v 3Q<br />
A :<br />
N H<br />
pis<br />
/' /• /' ^3<br />
%\)<br />
1*64 CRUISE *<br />
achaeus calypso, Ebalia affinis, Inachus biceps, Macropodia<br />
spinulosa, Parthenope notialis, Pisa carinimana<br />
18. Ghana. [05°04'N, 00°12'E] to 05°01'N, 00°12'E,<br />
3047-3129 m, soft dark gray clay, Blake Trawl, 26<br />
May 1964: Ethusina beninia<br />
22. Ghana. 05°25'N, 00°01'W to 05°22'N, 00°02'W, 51<br />
m, rough bottom, 6' OT, 27 May 1964: Achaeus buderes,<br />
Calappa pelii, Calypsachaeus calypso, Ebalia tuberculata,<br />
Ethusa vossi, Herbstia condyliata, Ilia spinosa, Inachus angolensis,<br />
Inachus nanus, Macropodia hesperiae, Nanocassiope<br />
melanodactyla, Parthenope massena, Parthenope notialis,<br />
Stenorhynchus lanceolatus, Stemodromia spinirostris<br />
23. Ghana. 05°10'N, 00°25'W to 05°08'N, 00°28'W, 42<br />
m, foliate brown to orange bryozoans, 6' OT, 28 May<br />
1964: Achaeus buderes, Calappa pelii, Capartiella longipes,<br />
Ebalia affinis, Ebalia tuberculata, Ethusa vossi, Eurynome<br />
parvirostris, Heterocrypta maltzami, Ilia spinosa, Inachus<br />
biceps, Inachus nanus, Macropipus rugosus, Macropodia gils&ni,<br />
Macropodia spinulosa, Medorippe lanata, Nanocassiope<br />
melanodactyla, Parthenope massena, Parthenope notialis, Pilumnus<br />
perrieri, Pisa carinimana, Stenorhynchus lanceolatus<br />
24. Ghana. 04°56'N, 00°47.5'W to 04°56'N, 00°50 / W,<br />
35-37 m, dark red bryozoans, 6' OT, 28 May 1964:<br />
GULF<br />
t<br />
/ ^<br />
OF GUINEA . IMS C«UIS£ tf<br />
Z 300 • /<br />
\% /<br />
A? 98 (J Sae Tom<br />
FIGURE 87.—Cruise track of the RA^ Pillsbury in the Gulf of Guinea, 1964 and 1965<br />
(from Voss, 1966, fig. 2).<br />
337<br />
\<br />
* 1
338<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Achaeus buderes, Calappa rubroguttata, Capartiella longipes, gilsoni, Monodaeus rouxi, Pseudomyra mbizi, SoUnolambrus<br />
Ebalia qffinis, Ebalia tuberculata, Ethusa vossi, Heterocrypta noordendei<br />
maltzami, Ilia spinosa, Inachus biceps, Machaerus atlanticus, 46. Ivory Coast. O5°O7'N, 04°32'W to O5°O7'N, 04°36'W,<br />
Macropipus rugosus, Macropodia spinulosa, Medorippe lan- 38-42 m, mud with dense Jullienella, 6' OT, 30 May<br />
ata, Nanocassiope melanodactyla, Palicus caronii, Parthenope 1964: Calappa pelii, Capartiella longipes, Cronius ruber,<br />
massena, Parthenope notialis, Pilumnus stebbingi, Pisa cari- Ethusa vossi, Heterocrypta maltzami, Ilia spinosa, Leopoldius<br />
nimana, Portunus inaequalis, Sttnorhynchus lanceolatus, Ster- pisifer, Machaerus atlanticus, Machaerus oxyacantha, Macronodromia<br />
spinirostris, Typhlocarcinodes integrifrons<br />
podia spinulosa, Medorippe lanata, Nanocassiope melanodac-<br />
26. Ghana. 04°57'N, 01°16'W to 04°59'N, 01°16.5'W, 27 tyla, Parthenope notialis, Phyllodorippe armata, Pisa carini-<br />
m, shell bottom (scallops), 6' OT, 28 May 1964: mana, Stenorhynchus lanceolatus, Stemodromia spinirostris<br />
Dromia monodi, Ethusa vossi, Ilia spinosa, Inachus biceps, 47. Ivory Coast. 05°04.5'N, 04°51.5'W, 37 m, bottom<br />
Parthenope massena, Portunus inaequalis, Stenorhynchus lan- with Jullienella, 6' OT, 31 May 1964: Calappa pelii,<br />
ceolatus<br />
Calappa rubroguttata, Ebalia affinis, Ethusa vossi, Leopoldius<br />
27. Ghana. 04°48'N, 01°42'W to 04°49'N, 01°47'W, 33 pisifer, Machaerus oxyacantha, Macropodia gilsoni, Macro-<br />
m, 6' OT, 28 May 1964: Ebalia tuberculata, Pilumnus podia spinulosa, Medorippe lanata, Nanocassiope melanodac-<br />
perrieri, Pisa carinimana, Portunus inaequalis<br />
tyla, Neodorippe armata, Philyra laevidorsalis, Phyllodorippe<br />
28. Ghana. 04°40'N, 02°00'W to [04°39'N, 02'02'W], 49- armata, Pisa carinimana, Portunus inaequalis, Stemodromia<br />
53 m, 6' OT, 28 May 1964: Capartiella longipes, Inachus spinirostris<br />
angolensis, Machaerus oxyacantha, Macropodia gilsoni, Ma- 48. Ivory Coast. [05°05'N, 04°59.5'W], 22 m, 6' OT, 31<br />
cropodia spinulosa, Medorippe lanata, Pseudomyra mbizi, May 1964: Cronius ruber, Ilia spinosa, Machaerus oxyacan-<br />
Raninoides bouvieri, Stenorhynchus lanceolatus, Stemodromia tha, Macropodia spinulosa, Phyllodorippe armata, Stenorhyn-<br />
spinirostris<br />
chus lanceolatus<br />
29. Ghana. [04°38'N, 02°02'W to 04°36'N, 02°00'W], 49. Ivory Coast. 05°00'N, 05°00'W to 04°59'N, 05°00'W,<br />
58-60 m, 40' OT, 28 May 1964: Inachus nanus, Pseudo- 73-77 m, 6' OT, 31 May 1964: Calappa pelii, Ethusa<br />
myra mbizi.<br />
rugulosa, Inachus angolensis, Inachus nanus, Monodaeus rouxi,<br />
30. Ghana. 04°46'N, O2°3O'W to 04°45'N, 02°33'W, 61- Phyllodorippe armata, Pseudomyra mbizi, SoUnolambrus noor-<br />
64 m, coral, 40' OT, 28 May 1964: Macropodia gilsoni, dendei<br />
Medorippe lanata, Stenorhynchus lanceolatus<br />
50. Ivory Coast. 04°58'N, 05°00'W to 04°57'N, 05°01'W,<br />
32. Ghana. 04°37'N, O2°32'W to 04°38'N, 02°35'W, 110<br />
128-192 m, 6' OT, 3 May 1964: Ethusa rugulosa, Inachus<br />
m, 40' OT, 28 May 1964: Macropodia gilsoni, Pseudomyra angolensis, Macropodia gilsoni, Monodaeus rouxi, Parthenope<br />
mbizi<br />
notialis, Pseudcmyra mbizi, SoUnolambrus noordendei<br />
34. Ghana. 03°53'N, 02°33'W to [03°47'N, 02°33'W], 51. Ivory Coast. 04°56'N, 05°01'W to 04°56.6'N, 05°-<br />
1948-1984 m, mud, Blake Trawl, 29 May 1964: Ethu- 03'W, 329-494 m, 6' OT, 31 May 1964: Acanthocarpus<br />
sina beninia<br />
brevispinis, Bathynectes piperitus, Carcinoplax bamardi, Ger-<br />
41. Ivory Coast. 04°47'N, O3°33'W to 04°47'N, 03°35'W, yon maritae<br />
641-842 m, 6' OT, 30 May 1964: Ethusa rosacea, Geryon<br />
maritae<br />
42. Ivory Coast. 05°02.5'N, 03°49.5'W to 05°05'N, 03°-<br />
59. Ivory Coast. 04°57.5'N, 05°- 22W to 04°57'N, 05°-<br />
30'W, 55-64 m, mud with dense branched Foraminifera,<br />
6' OT, 1 Jun 1964: Macropodia gilsoni, Parthenope<br />
52^, 62-75 m, mud with brown, branched Forami- notialis, Pseudomyra mbizi, SoUnolambrus noordendei<br />
nifera, 6' OT, 30 May 1964: Atlantotlos rhombifer, Ca- 60. Ivory Coast. 04°55'N, 05°34.5'W to 04°- 54'N, 05°lappa<br />
pelii, Capartiella longipes, Ilia spinosa, Inachus ango- 37^, 79-82 m, coral or rock, 6' OT, 1 Jun 1964:<br />
lensis, Inachus nanus, Macropipus rugosus, Macropodia gil- Capartiella longipes, Ethusa rugulosa, Inachus nanus, Macrosoni,<br />
Macropodia straeleni, Medorippe lanata, Nanocassiope podia hesperiae, Macropodia straeleni, Medorippe lanata,<br />
melanodactyla, Parthenope notialis, Pisa carinimana, Pseudo- Parthenope notialis, Pisa carinimana, Pseudomyra mbizi, Somyra<br />
mbizi, SoUnolambrus noordendei, Stenorhynchus lanceo- Unolambrus noordendei<br />
latus<br />
62. Ivory Coast. 04°45'N, 06°13.5'W to 04°44'N,<br />
43. Ivory Coast. Surface tow during Sta. 42, 0.5 m net, 30 06° 16^, 46 m, brown, branched and foliate Fora-<br />
May 1964: Callinectes pallidus<br />
minifera, 6' OT, 1 Jun 1964: Calappa pelii, Ilia spinosa,<br />
44. Ivory Coast. 05°05'N, 04°00'W to 05°04'N, 04°02'W, Inachus angolensis, Machaerus atlanticus, Macropodia gilsoni,<br />
403-586 m, hard dark gray mud, 6' OT, 30 May Medorippe lanata, Monodaeus rouxi, Parthenope notialis, Pisa<br />
1964: Balhynectes piperitus, Carcinoplax bamardi, Ethusa carinimana, Pseudomyra mbizi, Raninoides bouvieri, Stemo-<br />
rosacea, Geryon maritae<br />
dromia spinirostris<br />
45. Ivory Coast. 05°05'N, 04°04.5'W to 05°06'N, 04°-<br />
06'W, 73-97 m, 40' OT, 30 May 1964: Macropodia<br />
63. Ivory Coast. 04°35'N, 06°40'W to 04°35'N, 06°4l'W,<br />
64 m, sandy mud with shells, 6' OT, 2 Jun 1964:
NUMBER 306 339<br />
Atlantotlos rhombifer, Ilia spinosa, Inachus angolensis, Ma230.<br />
Nigeria. 06° 1 l'N, 03°36'E to 06° 10'N, 03°38'E, 82cropodia<br />
gilsoni, Parthenope notialis, Pseudomyra mbizi, Ster- 97 m, hard ground, with gorgonians, coral, rock, 40'<br />
nodromia spinirostris<br />
OT, 11 May 1965: Macropodia hesperiae, Pisa armata<br />
64. Ivory Coast. 04°23'N, 07°06.5'W to 04°22'N, 232. Nigeria. 05°56'N, 04°27'E to 05°54'N, 04°27'E, 101-<br />
07°08.5'W, 68 m, 6' OT, 2 Jun 1964: Calappa pelii,<br />
132 m, green mud, 40' OT, 11 May 1965: Pseudomyra<br />
Inachus angolensis, Inachus nanus, Macropipus rugosus, Ma- mbizi<br />
cropodia gilsoni, Medorippe lanata, Parthenope notialis, Pisa236.<br />
Nigeria. 05°20'N, 04°45'E to 05° 19'N, 04°48'E, 101carinimana,<br />
Pseudomedaeus africanus, Raninoides bouvieri 128 m, coral ground, rough, 40' OT, 12 May 1965:<br />
65. Ivory Coast. 04°15'N, 07°32'W to 04°12'N, Pseudomyra mbizi<br />
O7°35.5'W, 46-49 m, 40' OT, 2 Jun 1964: Macropipus 237. Nigeria. 05°19'N, 04°48'E to O5°O7'N, 04°55'E, 101<br />
rugosus, Macropodia hesperiae, Macropodia spinulosa, Ma- m, 10' OT, 12 May 1965: Calappa pelii, Macropodia<br />
cropodia straeleni, Medorippe lanata, Stenorhynchus lanceola- gilsoni, Monodaeus rouxi, Pseudomyra mbizi, Solenolambrus<br />
tus<br />
noordendei<br />
68. Liberia. 04°23'N, 08°05.5'W to 04°24'N, 08°07.5'W, 239. Nigeria. 04°56'N, 05°00'E to 04°54'N, 05°05'E, 73 m,<br />
70 m, broken shell, 6' OT, 3 Jun 1964: Achaeus foresti, 10' OT, 13 May 1965: Atlantotlos rhombifer, Capar-<br />
Atlantotlos rhombifer, Calappa pelii, Calypsachaeus calypso, tiella longipes, Inachus angolensis, Inachus nanus, Macropodia<br />
Capartiella longipes, Ebalia affinis, Ethusa vossi, Eurynome gilsoni, Macropodia straeleni, Monodaeus rouxi, Pseudomyra<br />
aspera, Heterocrypta maltzami, Homola barbata, Ilia spinosa, mbizi<br />
Inachus nanus, Macropipus rugosus, Macropodia gilsoni, 240. Nigeria. 04°44'N,05°17'E to04°41'N,05°19'E, 37 m,<br />
Macropodia straeleni, Medorippe lanata, Monodaeus rouxi, 10' OT, 13 May 1965: Capartiella longipes<br />
Parthenope notialis, Pseudomyra mbizi, Raninoides bouvieri, 241. Nigeria. 04°35'N, 05°18'E to 04°34'N, 05°19'E, 59-<br />
Solenolambrus noordendei, Stemodromia spinirostris<br />
63 m, mud and shell, 10' OT, 13 May 1965: Atlantotlos<br />
69. Liberia. 04°29.5'N, 08°06'W to 04°29.5'N, rhombifer, Calappa pelii, Capartiella longipes, Homola bar-<br />
08°07.5'W, 29 m, coral or rock, 6' OT, 3 Jun 1964: bata, Inachus angolensis, Machaerus oxyacantha, Macropipus<br />
Inachus nanus, Pisa carinimana, Pseudomyra mbizi<br />
rugosus, Macropodia gilsoni, Medorippe lanata, Portunus<br />
70. Liberia. 04°30'N, 08°09'W to 04°29.5'N, 08°09'W, validus, Pseudomyra mbizi, Raninoides bouvieri<br />
33 m, branched Foraminifera, 6' OT, 3 Jun 1964: 245. Nigeria. 04°32'N, 05°07'E to °31'N, 05°13'E, 64-<br />
Apiomithrax violaceus, Ethusa vossi, Nanocassiope melano- 119 m, mud, 40' OT, 13 May 1965: Pseudomyra mbizi<br />
dactyla, Paractaea margaritaria, Parthenope massena, Pisa246.<br />
Nigeria. 04° 13'N, 05°30'E to 04° 10'N, 05°33'E, 37 m,<br />
carinimana, Stenorhynchus lanceolatus<br />
40' OT, 13 May 1965; Ilia spinosa, Portunus inaequalis,<br />
73. Liberia. ?04°38'N, ?09°20'W to 04°40'N, 09°20'W, Pseudomyra mbizi, Stenorhynchus lanceolatus<br />
311-366 m, 40' OT, 4 Jun 1964: Bathynectes piperitus 248. Nigeria. 04°03'N, 05°4l'E to 04°07'N, O5°4O'E, 33 m,<br />
74. Liberia. 04°20'N, 09°26'W to 04°30'N, 09°22'W,<br />
10' OT, 13 May 1965: Achaeus turbator, Atlantotlos<br />
641-733 m, 40' OT, 4 Jun 1964: Ethusa rosacea, Geryon rhombifer, Calappa rubroguttata, Calypsachaeus calypso, Ca-<br />
maritae<br />
partiella longipes, Dromia monodi, Ebalia affinis, Ebalia<br />
82. Liberia. 04°57'N, 09°30'W to 04°58'N, 09°32'W, tuberculata, Ethusa vossi, Heterocrypta maltzami, Ilia spinosa,<br />
146-150 m, 40' OT, 5 Jun 1964: Macropodia macrocheles Inachus biceps, Leopoldius pisifer, Macropodia spinulosa,<br />
83. Liberia. 04°59'N, 09°37'W to 04°57.5'N, 09°33'W, Medorippe lanata, Parthenope massena, Phyllodorippe armata,<br />
156-220 m, 40' OT, 5 Jun 1964: Macropipus rugosus<br />
Pilumnus perrieri, Pisa carinimana, Portunus inaequalis,<br />
224. Nigeria. Lagos, 06°28'N, 03°23'E, shore, sand beach, Stenorhynchus lanceolatus<br />
ichthyocide, 9 May 1965: Ocypode cursor, Pilumnopeus 250. Nigeria. 04°06'N, 05°58'E to 04°02'N, 06°04'E, 24 m,<br />
africanus<br />
brackish water, mud, 10' OT, 14 May 1965: Callinectes<br />
225. Nigeria. Lagos harbor, 06°28'N, 03°23'E, shore at pallidus, Dromia monodi, Machaerus oxyacantha, Nanocas-<br />
dock, 9 May 1965: Plagusia depressa<br />
siope melanodactyla, Phyllodorippe armata, Parthenope mas-<br />
226. Nigeria. Lagos harbor, 06°28'N, O3°23'E, surface at<br />
sena, Portunus inaequalis<br />
dock, night light, 9 May 1965: Callinectes pallidus 251. Nigeria. O4°O3'N, 06°03'E to 04°04'N, 06°04'E, 27 m,<br />
227. Nigeria. Lagos harbor, 06°28'N, O3°23'E, shore, on<br />
mud, 10' OT, 14 May 1965: CallinecUs pallidus, Ma-<br />
rocks and seawall, by hand, 10 May 1965: Goniopsis chaerus oxyacantha, Phyllodorippe armata<br />
pelii, Pachygrapsus gracilis<br />
252. Nigeria. 04°04'N, 06° 18'E to 04°05'N, 06°22'E, 30 m,<br />
228. Nigeria. Lagos harbor, 06°28'N, 03°23'E, surface at<br />
mud, 10' OT, 14 May 1965: Callinectes pallidus, Ilia<br />
dock, 10 May 1965: Callinectes pallidus<br />
spinosa, Machaerus oxyacantha, Phyllodorippe armata, Por-<br />
229. Nigeria. Lagos harbor, 06°28'N, O3°23'E, surface at tunus inaequalis, Portunus validus, Stenorhynchus lanceolatus<br />
dock, dip net, 10 May 1965: Callinectes amnicola, Calli- 253. Nigeria. 04°04'N, 06°35'E to 04°03'N, 06°38'E, 33nectes<br />
pallidus<br />
40 m, mud, 10' OT, 14 May 1965: Calappa pelii,
340<br />
Leopoldius pisifer, Parthenope massena, Phyllodorippe armata,<br />
Stenorhynchus lanceolatus<br />
254. Nigeria. 03°50'N, 07°08'E to 03°51'N, 07° 12'E, 148-<br />
174 m, 40' OT, 14 May 1965: Pseudonym mbizi<br />
255. Nigeria. 03°49'N, 07°38'E to 03°48'N, 07°42'E, 264-<br />
269 m, 40' OT, 14 May 1965: Acanthocarpus brevispinis,<br />
Carcinoplax bamardi, Inachus grallator, Macropodia macrocheles<br />
256. Nigeria. 03°45'N, 08°03'E to 03°45'N, 08°02'E, 409-<br />
485 m, 40' OT, 14 May 1965: Acanthocarpus brevispinis,<br />
Carcinoplax bamardi<br />
257. Fernando Poo. 03°45'N, 08°48'E, shore, ichthyocide,<br />
15 May 1965: Callinectes marginatus, Callinectes pallidus,<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Eurypanopeus blanchardi, Pachygrapsus transversus, Xanthod- Pilumnus stebbingi, Sakaila africana<br />
ius inaequalis inaequalis<br />
316. Nigeria. Lagos, 06°28'N, 03°23'E, shore, sand beach,<br />
258. Fernando Poo. 03°45'N, 08°48'E, shore, ichthyocide,<br />
ichthyocide, 28 May 1965: Ocypode cursor, Pachygrapsus<br />
15 May 1965: Callinectes pallidus, Epixanthus hellerii,<br />
Eurypanopeus blanchardi, Grapsus grapsus, Pachygrapsus<br />
transversus, Percnon gibbesi, Xanthodius denticulatus, Xan-<br />
transversus<br />
thodius inaequalis inaequalis<br />
Geronimo COLLECTIONS<br />
259. Cameroon. 03°53'N, 08°53'E to 03°51'N, 08°54'E,<br />
59 m, mud and broken shell, 10' OT, 16 May 1965:<br />
Calappa pelii, Inachus angolensis, Macropipus rugosus, Macropodia<br />
gilsoni, Pseudomyra mbizi, Raninoides bouvieri<br />
260. Cameroon. 03°45'N, 09°05'E to 03°43'N, 09°10'E,<br />
46 m, 10' OT, 16 May 1965: Atlantotlos rhombifer,<br />
Calappa pelii, Calypsachaeus calypso, Ilia spinosa, Inachus<br />
minus, Macropipus rugosus, Macropodia gilsoni, Medorippe<br />
lanata, Parthenope notialis, Raninoides bouvieri<br />
271. Annobon. 01°25'S, 05°38'E, NE coast between Punta<br />
Yoyo and Punta Pedrinha, shore, ichthyocide, 19 May<br />
1965: Acanthonyx minor, Grapsus grapsus, Nanocassiope<br />
melanodactyla, Pachygrapsus transversus, Percnon gibbesi,<br />
Plagusia depressa, Sirpus gordonae, Xanthodius denticulatus,<br />
Xanthodius inaequalis inaequalis<br />
273. Annobon. 01°24'S, O5°37'E, N coast, near Islote Piramide,<br />
shore, ichthyocide, 19 May 1965: Acanthonyx<br />
minor, Domecia acanthophora africana, Grapsus grapsus, Ocypode<br />
cursor, Pachygrapsus transversus, Plagusia depressa<br />
275. Annobon. 01°24'S, O5°37'E to 01°24'S, 05°38'E, 9-<br />
69 m, rubble of coralline algae, dredge, 20 May 1965:<br />
Cataleptodius Jloridanus, Cronius ruber, Dynomene filholi,<br />
Euryozius pagalu, Globopilumnus stridulans, Herbstia nitida,<br />
Nanocassiope melanodactyla, Nanopilumnus boletifer, Paractaea<br />
margaritaria, Paractaea rufopunctata africana, Pisa calva,<br />
Ranilia constricta, Stenorhynchus lanceolatus, Xanthodius denticulatus<br />
278. Annobon. 01°24'S, 05°37'E, shore [collection made in<br />
transit to Crater Lake], 20 May 1965: Ocypode cursor<br />
281. Annobon. 01°24'S, O5°37'E, shore, ichthyocide, 20<br />
May 1965: Callinectes marginatus, Grapsus grapsus, Ocy-<br />
pode cursor<br />
282. Annobon. 01°28'S, 05°36'-37'E to 01°29'S, 05°36'E,<br />
18-37 m, nodular coralline algae, dredge, 21 May<br />
1965: Domecia acanthophora africana, Dynomene filholi,<br />
Euryozius pagalu, Globopilumnus stridulans, Glyptoxanthus<br />
angolensis, Herbstia nitida, Microcassiope rufopunctata, Nanocassiope<br />
melanodactyla, Paractaea margaritaria, Paractaea<br />
rufopunctata africana, Pisa calva<br />
283. Annobon. 01°29'S, 05°35'E, 51-55 m, nodular coral-<br />
line algae, dredge, 21 May 1965: Dynomene filholi,<br />
Ebalia tuberculata, Euryozius pagalu, Herbstia nitida, Homola<br />
barbata, Nanocassiope melanodactyla, Paractaea margaritaria,<br />
Parthenope expansa, Parthenope massena, Pilumnus<br />
stebbingi, Pisa calva<br />
284. Annobon. 01°30'S, 05°36'E, 73 m, black basaltic<br />
rocks, dredge, 21 May 1965: Nanocassiope melanodactyla,<br />
Collections made by National Marine Fisheries<br />
Service vessel Geronimo off Gabon in 1963.<br />
179. Off Gabon River, OO°O2.2'S, 08°50.2'E, 161 fm (293<br />
m), 31 Aug 1963: Acanthocarpus brevispinis, Bathynectes<br />
piperitus, Carcinoplax bamardi<br />
185. 00°32'S, 08°42'E, 110 fm (200 m), 1 Sep 1963:<br />
Pseudomyra mbizi<br />
187. OO°32'S, 08°40'E, 165 fm (300 m), 1 Sep 1963:<br />
Pseudomyra mbizi<br />
191. 01°28'S, 08°24.5'E, 165 fm (300 m), 3 Sep 1963:<br />
Bathynectes piperitus, Geryon maritae<br />
197. 01°30'S, 08°27.5'E, 110 fm (200 m), Sep 1963: Macropodia<br />
macrocheles<br />
198. 01°28'S, 08°24.5'E, 165 fm (300 m), 3 Sep 1963:<br />
Acanthocarpus brevispinis, Bathynectes piperitus, Carcinoplax<br />
bamardi, Geryon maritae<br />
199. 01°26.4'S, 08°24'E, 220 fm (400 m), 3 Sep 1963:<br />
Bathynectes piperitus<br />
202. 02°00'S, 08°55'E, 55 fm (100 m), 4 Sep 1963: Acanthocarpus<br />
brevispinis, Macropipus rugosus<br />
203. 02°01'S, 08°50.5'E, 110 fm (200 m), 4 Sep 1963:<br />
Acanthocarpus brevispinis, Bathynectes piperitus, Carcinoplax<br />
bamardi, Geryon maritae<br />
206. 02°00'S, 08°46.5'E, 250-335 fm (455-610 m), 4 Sep<br />
1963: Bathynectes piperitus<br />
211. 02°32'S, 09°05'E, 55 fm (100 m), 5 Sep 1963: Atelecyclus<br />
rotundatus, Euchirograpsus liguricus, Macropipus rugosus,<br />
Pseudomedaeus africanus<br />
212. 02°30'S, 08°58'E, 110 fm (200 m), 5 Sep 1963: Calappa<br />
pelii<br />
213. 02°31'S, 08°51'E, 165 fm (300 m), 5 Sep 1963: Bathynectes<br />
piperitus, Macropipus rugosus
NUMBER 306 341<br />
214. 02°30'S, 08°52'E, 300 fm (546 m), 5 Sep 1963: Bathynecles<br />
piperitus<br />
220. 03°02'S, 09°21'E, 165 fm (300 m), 6 Sep 1963: Bathynectes<br />
piperitus<br />
228. 03°31'S, 09°53'E, 165 fm (300 m), 7 Sep 1963: Macropipus<br />
rugosus<br />
235. 04°01'S, 10°35'E, 55 fm (100 m), 8 Sep 1963: Calappa<br />
pelii, Macropodia gilsoni, Parthenope notialis, Sakaila africana<br />
247. 04°38.4'S, ll°01.2'E, 220 fm (400 m), 9 Sep 1963:<br />
Bathynectes piperitus<br />
Undaunted COLLECTIONS<br />
Collections made by National Marine Fisheries<br />
Service vessel Undaunted off Angola and South-<br />
<strong>West</strong> Africa (below 17°15'S) in 1968.<br />
94. 16°27'S, 11°35'E, 90 m, 18 Mar 1968: Goneplax rhomboides,<br />
Inachus angolensis, Macropipus australis<br />
95. 16°37'S, 11°22'E, 126 m, 18 Mar 1968: Calappa pelii,<br />
Goneplax rhomboides, Macropipus australis, Macropodia gilsoni,<br />
Macropodia spinulosa, Parthenope notialis<br />
96. 16°41'S, 11°21'E, 162 m, 18 Mar 1968: Calappa pelii,<br />
Eurynome aspera, Inachus angolensis, Inachus grallator, Macropipus<br />
australis, Parthenope notialis, Pisa armata, Solenolambrus<br />
noordendei<br />
102. 17°O2'S, ll°40'E, 54 m, 24 Mar 1968: Calappa pelii,<br />
Medorippe lanata<br />
103. 17°O6'S, 11°35'E, 90 m, 24 Mar 1968: Calappa pelii,<br />
Inachus angolensis, Macropipus australis, Medorippe lanata,<br />
Pseudomyra mbizi<br />
104. 17°09'S, ll°30'E, 126 m, 24 Mar 1968: Macropipus<br />
australis<br />
105. 17°13'S, 11 O 27'E, 155 m, 24 Mar 1968: Macropipus<br />
australis<br />
106. 17° 18'S, 11 °24'E, 225 m, 24 March 1968: Euchirograpsus<br />
liguricus, Macropipus australis, Monodaeus couchii<br />
107. 17°23'S, 11 °20'E, 359 m, 24 Mar 1968: Euchirograpsus<br />
liguricus, Geryon maritae<br />
111. 10°36'S, 13°12'E, ca 366 m, 12 Apr 1968: Bathynectes<br />
piperitus
Localities from the literature or from specimenassociated<br />
data cited in the species accounts are<br />
listed here, alphabetically by country, along with<br />
coordinates. Spellings and coordinates are from<br />
gazetteers of the United States Board on Geographic<br />
Names. Alternate spellings and coordinates<br />
from sources other than those gazetteers are<br />
given in brackets. Localities that we have not<br />
been able to locate are identified in the text by a<br />
question-mark in brackets, [?], and are cross-referenced<br />
in this gazetteer by species name. Specific<br />
localities identified in the text as "near" another<br />
locality (e.g., Boulbinet, near Conakry) or as in<br />
another locality, (e.g., Samba, Luanda) are not<br />
listed separately here if they could not be located<br />
in the Board on Geographic Names gazetteers.<br />
Localities in the literature or from specimen labels<br />
originally accompanied by coordinates are not<br />
repeated here; in the list, coordinates added by<br />
us are set off in brackets. We have anglicized<br />
country names but not names for localities within<br />
countries. We have used older names for two of<br />
the offshore islands of the Gulf of Guinea, now<br />
part of Equatorial Guinea: Annobon, instead of<br />
Pagalu, and Fernando Poo, instead of Macias<br />
Nguema Biyogo.<br />
Other sources, which have been helpful in identifying<br />
<strong>West</strong> <strong>African</strong> localities either by listing<br />
coordinates or by giving maps, include: Forest<br />
and Games, 1960 (Morocco); Guinot and Ribeiro,<br />
1962 (Cape Verde Islands, Angola);<br />
Longhurst, 1958 (Sierra Leone and Guinea<br />
Shelf); Rathbun, 1921 (a list of <strong>West</strong> <strong>African</strong><br />
localities, mostly in the Congo and Zaire); Rossignol,<br />
1962 (localities between Nigeria and<br />
southern Angola); Sourie, 1954a (Senegal, especially<br />
around Dakar); and Tiirkay, 1976b (Madeira).<br />
Appendix II: Gazetteer<br />
342<br />
Angola<br />
Baia do Ambriz<br />
Baia de Ambrizete<br />
Baia de Benguela<br />
Baia da Caota, Benguela<br />
Baia dos Elefantes<br />
Baia Farta, Benguela<br />
Baia do Lobito<br />
Baia de Luanda<br />
Baia de Mocamedes<br />
Baia de Porto Amboim<br />
Baia de Santa Marta<br />
Baia dos Tigres<br />
Baia das Vacas [Ponta das<br />
Vacas]<br />
Benguela<br />
Cacuaco<br />
Chiloango<br />
Egito Praia<br />
Ilha de Luanda<br />
Luanda [St. Paul do<br />
Loanda]<br />
Lucira<br />
Mocamedes<br />
Morro da Cruz<br />
Musserra [Mussera,<br />
Massera]<br />
Mussulo Grande<br />
Ponta da Caruita<br />
Ponta da Moita Seca<br />
Ponta do Morro [Cap<br />
Morro?]<br />
Ponta de Sao Jose<br />
Ponta do Sombreiro<br />
Porto Alexandre<br />
Porto Amboim<br />
Praia Amelia, Mocamedes<br />
Praia das Conchas, Mocamedes<br />
Quicembo [Kinsembo]<br />
Rio Chiloango<br />
Rio Cuanza<br />
Rio Cunene [Kunene River]<br />
Santo Antonio do Zaire [San<br />
Antonio, Saint-Antoine]<br />
07° 50'S, 13<br />
07°<br />
12°<br />
12°<br />
13°<br />
12°<br />
12°<br />
08°<br />
15°<br />
10°<br />
13°<br />
16°<br />
12°<br />
C<br />
13'S, 12 C<br />
35'S, 13 C<br />
36'S, 13 C<br />
13'S, 12°<br />
36'S, 13'<br />
20'S, 13 C<br />
47'S, 13 C<br />
10'S, 12'<br />
42'S, 13'<br />
51'S, 12 C<br />
'06'E<br />
'51'E<br />
'21'E<br />
'16'E<br />
'44'E<br />
'13'E<br />
'34'E<br />
'16'E<br />
"08'E<br />
'45'E<br />
'28'E<br />
(<br />
38'S, 11' 46'E<br />
(<br />
37'S, 13' 14'E<br />
12° 35'S, 13<br />
08°<br />
05°<br />
11°<br />
08°<br />
08°<br />
C(<br />
25'E<br />
47'S, 13'>22'E<br />
12'S, 12''08'E<br />
59'S, 13''46'E<br />
48'S, 13''13'E<br />
48'S, 13''14'E<br />
13° 51'S, 12''31'E<br />
15° 10'S, 12'WE<br />
08° 57'S, 13''04'E<br />
07° 29'S, 12''58'E<br />
08° ll'S, 13'<br />
D<br />
12° '35'S, 13' 16'E<br />
06° 07'S, 12''16'E<br />
10° 45'S, 13'>43'E<br />
12 C<br />
12c<br />
15 C<br />
10 c<br />
15 C<br />
15 C<br />
07 c<br />
05 c<br />
09 c<br />
17 C<br />
06 c<br />
5 17'E<br />
3 12'E<br />
36'S, 13'<br />
3<br />
35'S, 13' 18'E<br />
3<br />
49'S, 11' 53'E<br />
'44'S, 13'°44'E<br />
"12'S, 12'°06'E<br />
'07'S, 12°07'E<br />
'44'S, 13°03'E<br />
'12'S, 12°07'E<br />
'19'S, 13°08'E<br />
'20'S, 11 °50'E<br />
'07'S, 12°18'E
NUMBER 306<br />
Annobon [Pagalu, Equatorial<br />
Guinea]<br />
Isla Tortuga<br />
Islote Piramide<br />
Punta Pedrinha<br />
Punta Yoyo<br />
San Antonio<br />
Santa Cruz<br />
Ascension Island<br />
Azores<br />
Ilha do Corvo<br />
Ilha do Faial [Fayal]<br />
Almoxarife. See Acanthonyx<br />
brevifrons<br />
Caldeirinhas [Caldeira Inferno]<br />
Feteira<br />
Horta<br />
Pasteleiro<br />
Ilha das Flores<br />
Ilha da Muda. See Dromia<br />
marmorea<br />
Ilha do Pico<br />
Madalena<br />
Ilha de Sao Miguel<br />
Caloura<br />
Ponta Delgada<br />
Ponta da Galera<br />
Ilha Terceira<br />
Ponta Sao Diego. See<br />
Dromia marmorea<br />
Ilheu da Praia<br />
Rochas dos Capelinhos [Volcano<br />
Capelinhos]<br />
Cabinda<br />
Cabinda<br />
Chinchoxo [Tschintschotscho,<br />
Quinchoxo]<br />
Landana<br />
Quila [River]<br />
Cameroon<br />
Batanga<br />
Bibundi<br />
Douala<br />
Kribi<br />
Souellaba [Souelaba]<br />
Wouri River<br />
Yaounde<br />
Canary Islands<br />
Estrecho de la Bocaina<br />
Isla de Fuertaventura<br />
Puerto Cabras<br />
Punta Morro Jable [Ponta<br />
da Matorra, Punta del<br />
Mattorral]<br />
01°25'S, 05°36'E<br />
01°24'S, 05°38'E<br />
01°24'S, O5°37'E<br />
[01°24'35"S, O5°37'25"E]<br />
[01°24'45"S, 05°37'35"E]<br />
O1°27'S, 05°37'E<br />
O1°27'S, 05°37'E<br />
O7°57'S, 14°22'W<br />
38°30'N, 28°0O'W<br />
39°42'N, 31°06'W<br />
38°34'N, 28°42'W<br />
38°30'N, 28°37'W<br />
38°31'N, 28°41'W<br />
38°32'N, 28°38'W<br />
38°31'N, 28°38'W<br />
39°26'N, < i\°\3'^M<br />
38°28'N, 28°20'W<br />
38°32'N, 28°32'W<br />
37°47'N, 25°30'W<br />
37°42'N, 25°30'W<br />
37°44'N, 25°40'W<br />
37°42'N, 25°31'W<br />
38°43'N, 27°13'W<br />
39°03'N, 27°57'W<br />
38°36'N, 28°50'W<br />
05°33'S, 12°12'E<br />
[05°09.24'S, 12°03.75'E]<br />
05°13'S, 12°08'E<br />
05°58'S, 14°47'E<br />
[ca 02°54'N, 09°55'E]<br />
04°13'N, 08°59'E<br />
04°03'N, 09°42'E<br />
02°57'N, 09°55'E<br />
03°49'N, 09°33'E<br />
04°06'N, 09°43'E<br />
03°52'N, 11°31'E<br />
28°00'N, 15°3O'W<br />
28°47'N, 13°50'W<br />
28°20'N, 14°00'W<br />
28°29'N, 13°51'W<br />
28°02'N, 14°20'W<br />
Isla de la Gomera<br />
Isla de Gran Canaria<br />
Isla de Lanzarote<br />
Arrecife<br />
Playa Quemada<br />
Isla de La Palma<br />
Santa Cruz de La Palma<br />
Isla de Tenerife<br />
Ensenada de Crist ianos<br />
Playa de los Abrigos<br />
Puerto de la Cruz<br />
Puerto de Orotava<br />
Cape Verde Islands<br />
Baixo Joao Leitao<br />
Boa Vista, Ilha da<br />
Ilheu de Sal Rei<br />
Ponta Rodrigo<br />
Porto de Sal Rei<br />
Ribeira do Rabil<br />
Branco, Ilheu<br />
Brava, Ilha<br />
Baia de Faja di Agua<br />
[Porto da Faja]<br />
Ponta Garbeiro. See Dorrucia<br />
acanthophora africana<br />
Porto da Furna<br />
Fogo, Ilha do<br />
Ponta da Areia<br />
Ilheu Luis Carneiro<br />
Maio, Ilha de<br />
Porto Ingles<br />
Sal, Ilha do<br />
Baia da Murdeira [Mordeira]<br />
Pedra Lume<br />
Santo Antao, Ilha de<br />
Baia do Monte Trigo<br />
Ponta do Esbarradeiro<br />
Ponta da Praia Formosa<br />
Ponta do Sol<br />
Porto dos Carvoeiros [Baio<br />
do Porto Novo]<br />
Santa Luzia, Ilha de<br />
Sao Nicolau, Ilha de<br />
Tarrafal<br />
Sao Tiago, Ilha de [Santiago]<br />
Baia de Santa Clara<br />
Porto da Praia [La Praya,<br />
Praia]<br />
Porto de Sao Francisco<br />
Tarrafal<br />
Sao Vicente, Ilha de<br />
28°06'N, 17°08'W<br />
28°00'N, 15°36'W<br />
29°00'N, 13°40'W<br />
28°57'N, 13°32'W<br />
28°54'N,13°43'W<br />
28°40'N, 17°52'W<br />
28°41'N, 17°45'W<br />
28°19'N, 16°34'W<br />
28°02'N, 16°42'W<br />
28°01'N, 16°35'W<br />
28°24'N, 16°33'W<br />
28°46'N, 17°45'W<br />
16°00'N, 24°00'W<br />
15°48'N, 23°!!^<br />
16°05'N, 22°50'W<br />
16°10'N,22°56'W<br />
16°12'N, 22°43'W<br />
16°10'N, 22°56'W<br />
16°09'N, 22°59'W<br />
16°39'N, 24°41'W<br />
14°52'N, 24°43'W<br />
14°52'N, 24°45'W<br />
14°53'N, 24°41'W<br />
14°55'N, 24°25'W<br />
14°53'N, 24°31'W<br />
14°58'N, 24°40'W<br />
15°15'N,23 o 10'W<br />
15°08'N,23°13'W<br />
16°45'N, 22°55'W<br />
16°41'N, 22°57'W<br />
16°46'N, 22°54'W<br />
WOb'W, 25°10'W<br />
17°OrN,25°2O'W<br />
16°55'N, 25°14'W<br />
16°55'N, 25°13'W<br />
17°12'N, 25°06'W<br />
17°O2'N, 25°04'W<br />
16°46'N, 24°45'W<br />
16°35'N, 24°15'W<br />
16°34'N, 24°22'W<br />
15°05'N, 23°40'W<br />
15°01'N, 23°45'W<br />
14°54'N,23°31'W<br />
14°58'N, 23°28'W<br />
15°17'N, 23°46'W<br />
16°50'N, 25°00'W<br />
343
344<br />
Baia da Fateixa<br />
Baia das Gat as<br />
Ilheu Raso<br />
Ponta da Calheta [Baia de<br />
Calheta]<br />
Porto Grande<br />
Praia da Matiota [Baia da<br />
Matiota]<br />
Conception Bank<br />
Congo<br />
Bahua, See Cardisoma armatum<br />
Baie de Loango<br />
Baie de Pointe-Noire<br />
Banga [Banda]<br />
Djeno<br />
Loango<br />
Loya River<br />
Pointe Indienne<br />
Pointe Kounda<br />
Pointe-Noire<br />
Songololo River [Songolo<br />
River]<br />
Dahomey<br />
Cotonou<br />
Grand-Popo<br />
Lac Nokoue<br />
Zogbo, NW of Cotonou on<br />
Lac Nokoue<br />
Fernando Poo [Macias Nguema<br />
Biyogo, Equatorial Guinea]<br />
Mongola<br />
San Carlos<br />
Gabon<br />
Baie du Cap Lopez<br />
Barre des Portugais<br />
Cap Lopez<br />
Cap Santa Cfara [Pointe<br />
Santa Clara]<br />
Crique Banjia [Banya]<br />
Ivindo River<br />
Libreville<br />
Mayumba [Mayoumba]<br />
Nyanga<br />
Ogooue [stream]<br />
Owendo<br />
Pointe Banda<br />
Pointe Claire<br />
Pointe Gombe<br />
Pointe Panga<br />
Port-Gentil<br />
Sette Cama<br />
Gambia<br />
Bathurst<br />
Gambia River<br />
16°52'N, 25°04'W<br />
16°12'N, 22°43'W<br />
16°37'N, 24°36"W<br />
16°47'N, 24°58'W<br />
16°53'N, 25°O1'W<br />
16°53'N, 25°00'W<br />
29°30'N, 12°45'W<br />
04°36'S, 11°44'E<br />
04°47'S, 11°51'E<br />
04°34'S, 12°23'E<br />
04°55'S, 11°57'E<br />
04°39'S, 11°48'E<br />
04°49'S, 12°52'E<br />
04°40'S, 11°47'E<br />
04°ll'S, 11°23'E<br />
04°48'S, 11°51'E<br />
04°45'S, 11°51'E<br />
06°21'N, 02°26'E<br />
06°17'N, 01°50'E<br />
06°26'N, 02°27'E<br />
[06°24'N, 02°25'E]<br />
O3°3O'N, 08°42'E<br />
03°46'N, 08°44'E<br />
03°27'N, 08°33'E<br />
00°40'S, 08°50'E<br />
01°16'S, 09°00'E<br />
00°37'S, 08°43'E<br />
00°30'N, 09°19'E<br />
00°14'N, 09°40'E<br />
00°09'S) 12°09'E<br />
OO°23'N, 09°27'E<br />
03°25'S, 10°39'E<br />
02°59'S, 10°17'E<br />
00°49'S, 09°00'E<br />
00°17'N, 09°30'E<br />
03°46'S, ll°00'E<br />
01°08'S)09°26'E<br />
00°18'N, 09°18'E<br />
03°15'S,.10°32'E<br />
00°43'S, 08°47'E<br />
02°32'S, 09°45'E<br />
13°27'N, lG'^'W<br />
13°28'N, 16°34'W<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Gunjur<br />
Ghana<br />
Accra<br />
Ada<br />
Angaw Lagoon<br />
Ankobra River [Ancobra]<br />
Apam<br />
Axim<br />
Baya River<br />
Butre [Boutry, Butry]<br />
Chorkor, near Accra<br />
Christiansborg<br />
Densu<br />
Denu<br />
Dixcove<br />
Elmina [St. George<br />
d'Elmina]<br />
Hwini River<br />
Komenda<br />
Kumasi<br />
Labadi<br />
Ningo<br />
Prampram<br />
Sakumo Lagoon<br />
Sekondi<br />
Shama Bay<br />
Takoradi<br />
Tema harbor<br />
Tenkpobo[Tenpobo]<br />
Teshi [Teshie]<br />
Volta River delta<br />
Winneba<br />
Guinea<br />
Conakry<br />
Fotoba<br />
lie Blanche<br />
lie de Corail<br />
lie Kassa [Cassa]<br />
lies de Los<br />
lie Marara<br />
lie Roume<br />
lie Tamara<br />
Matakong<br />
Rio Pongo<br />
Tanene<br />
Guinea-Bissau [Portuguese<br />
Guinea]<br />
Arquipelago dos Bijagos<br />
[Bissagos Islands]<br />
Bissau<br />
Ilha de Bissau<br />
Ilha de Bubaque<br />
Ilha Caravela<br />
Ilha de Rubane [Rouban]<br />
l'N, 16°46'W<br />
05°33'N, 00°13'W<br />
05°46'N, OO°37'E<br />
05°48'N, OO°47'E<br />
04°53'N, O2°17'W<br />
05°17'N, 00°44'W<br />
04°52'N, 02°14'W<br />
[05°05'N ,01<br />
04°49'N,<br />
[05°33'N<br />
O5°33'N,<br />
05°31'N,<br />
06°05'N,<br />
04°48'N,<br />
05°05'N,<br />
o 22'W]<br />
01°55'W<br />
, 00°15'W]<br />
00°ll'W<br />
00°19'W<br />
01°08'E<br />
O1°57'W<br />
01°21'W<br />
04°53'N,<br />
05°03'N,<br />
06°41'N,<br />
05°33'N,<br />
06°03'N,<br />
05°42'N,<br />
05° 3 l'N,<br />
04°56'N,<br />
05°00'N,<br />
04°53'N,<br />
05°38'N,<br />
05°43'N,<br />
05°35'N,<br />
05°55'N,<br />
05°20'N,<br />
09° 3 l'N,<br />
09°30'N,<br />
09°26'N,<br />
09°26'N,<br />
09°29'N,<br />
09°30'N,<br />
10°02'N,<br />
09°28'N,<br />
09°29'N,<br />
09° 16'N,<br />
10°03'N,<br />
10°03'N,<br />
01°47"W<br />
01°29'W<br />
O1°37'W<br />
00°09'W<br />
00°ll'W<br />
00°07'E<br />
00°19'W<br />
01°42'W<br />
01°36'W<br />
01°45'W<br />
00°01'E<br />
00°08'W<br />
00°06'W<br />
01°00'E<br />
OO°37'W<br />
13°43'W<br />
13°48'W<br />
13°46'W<br />
13°45'W<br />
13 "48^<br />
14°02'W<br />
13°48'W<br />
13°49'W<br />
13°25'W<br />
14°04'W<br />
13°54'W<br />
, 16°05'W<br />
11°51'N, 1<br />
11°52'N, 15°46'W<br />
11°15'N, 15°52'W<br />
11°32'N, 16°20'W<br />
ll°20'N, 15°49'W
NUMBER 306<br />
Uha de Soga [Soja]<br />
Ivory Coast<br />
Fresco<br />
Grand-Bassam<br />
Grand-Lahou<br />
Jacqueville<br />
Lagune Ebrie<br />
Port-Bouet<br />
San-Pedro<br />
Sassandra<br />
Tabou<br />
Vridi<br />
Josephine Seamount<br />
La Chapelle Bank [Bane de la<br />
Chapelle]<br />
Liberia<br />
Bromley. See Sesarma (Chiromantes)<br />
buettikoferi<br />
Bushrod Island<br />
Cape Mesurado<br />
Farmington River<br />
Grand Cape Mount, near<br />
Robertsport<br />
Harbel [Habel]<br />
Junk River<br />
Lake Piso [Fisherman's<br />
Lake]<br />
Mesurado River<br />
Monrovia<br />
Robertsport<br />
St. Paul River<br />
Upper Buchanan<br />
Madeira Islands<br />
Agua de Pena<br />
Canical<br />
Enseada da Abra<br />
Funchal<br />
Ilha de Porto Santo<br />
Ilhas Desertas<br />
Deserta Grande<br />
Ilheu do Bugio<br />
Ilheu do Gorgulho<br />
Machico Bay<br />
Ponta da Garajau<br />
Ponta da Oliveira<br />
Ponta dos Reis Magos<br />
Ponta de Sao Lourenco<br />
Prainha. See Portunus<br />
hastatus<br />
Ribeira Brava<br />
Mauritania<br />
Baie de Cansado<br />
Baie de l'Ouest [on Cap<br />
Blanc peninsula]<br />
H°21'N, 15°52'W<br />
05°05'N, 05°34'W<br />
O5°12'N, 03°44'W<br />
05°08'N, 05°01'W<br />
05°12'N, 04°25'W<br />
05°14'N, 04°26'W<br />
05 o 15'N,03 o 58'W<br />
04°44'N, 06°37'W<br />
04°57'N, 06°05'W<br />
04°24'N, 07°21'W<br />
05°15'N, 04°02'W<br />
36°52'N, 14°20'W<br />
47°37'N, 07°18'W<br />
06°21'N, 10°47'W<br />
06°19'N, 10°49'W<br />
06°08'N, 10°22'W<br />
[06°44'N, 11°24'W]<br />
06°16'N, 10°21"W<br />
06°09'N, 10°23'W<br />
06°45'N, 11°16'W<br />
06°20'N, 10°48'W<br />
06°19'N, 10°48'W<br />
06°45'N, 11°22'W<br />
06°23'N, 10°48'W<br />
05°56'N, 10°04'W<br />
32°44'N,17°00'W<br />
32°42'N, 16°46'W<br />
32°44'N, 16°44'W<br />
32°44'N, 16°42'W<br />
32°38'N, 16°54'W<br />
33°04'N, 16°20'W<br />
32°30'N, l6 o 3ffYf<br />
32°30'N, lG^CW<br />
32°26'N, 16°29'W<br />
32°37'N, le^^<br />
32°42'N, 16°46'W<br />
32°38'N, 16°51'W<br />
32°38'N, ^"SCW<br />
32°38'N, \6°A:9rW<br />
32°44'N, 16°41'W<br />
32°39'N) 17°O4'W<br />
20°53'N, 17°01'W<br />
[ca. 20°47'N, 19°24<br />
Baie du Repos<br />
Baie de Saint-Jean<br />
Bane d'Arguin<br />
Bilaouakh. See Eurypanopeus<br />
blanchardi<br />
Cansado<br />
Cap Blanc<br />
Cap Timiris<br />
Nouakchott<br />
Pointe de Cansado<br />
Portendick [Portendic]<br />
Port-Etienne<br />
Tamzak [Tamxat]<br />
Morocco<br />
Agadir<br />
Asilah [Arzila]<br />
Bane de Spartel [off Cap<br />
Spartel]<br />
Cap Draa [Cap Noun]<br />
Cap Juby<br />
Cap Rhir [Cap Ghir]<br />
Cap Spartel<br />
David<br />
El Hank. See Pachygrapsus<br />
transversus<br />
Essaouira<br />
Fosse de Rabat<br />
Foum Agouitir [Puerto-Cansado]<br />
Foum Assaka<br />
Larache<br />
Lucien Saint. See Percnon<br />
gibbesi<br />
Mannesman<br />
Oued Bou Regrcg<br />
Oued Cherrat<br />
Plage David<br />
Pointe d'El Hank<br />
Rabat<br />
Sidi Bou Knadel [Bouknadel]<br />
Sidi Moussa<br />
Skhirat<br />
Tangier<br />
Temara<br />
Temara—plage<br />
Tillet [PTillelt]. See Acanthonyx<br />
lunulatus<br />
Nigeria<br />
Adu River<br />
Ajudaibo<br />
Bonny River<br />
Brass<br />
Forcados<br />
20°55'N, 17°O2'W<br />
19°27'N, 16°22'W<br />
20°10'N, 16°45'W<br />
20°51'N, 17°O2'W<br />
20°46'N, 17°03'W<br />
19°23'N, 16°32'W<br />
18°06'N, 15°57'W<br />
20°51'N, 17°02'W<br />
18°35'N, 16°O7'W<br />
20°54'N, 17°04'W<br />
17°26'N, 16°02'W<br />
30°24'N, 09°36'W<br />
35°28'N, 06°02'W<br />
[ca. 35°54'N, 06°14'W]<br />
28°44'N, ll'WW<br />
27°57'N, 12°55'W<br />
30°38'N, 09°54'W<br />
35°48'N, 05°56'W<br />
[33°50'N, O7°O7'W]<br />
31°31'N, 09°46'W<br />
[ca. 34°N, 07°W]<br />
28°04'N, 12° 14^<br />
29°08'N, 10°24'W<br />
35°12'N, 06°09'W<br />
[33°37'N, O7°33'W]<br />
34°03'N, Oe'SCW<br />
33°50'N, O7°O7'W<br />
[33°50'N, O7°O7'W]<br />
33°37'N, 07°40'W<br />
34°02'N, 06°50'W<br />
34°08'N, 06°44'W<br />
32°53'N, 08°51'W<br />
33°52'N, O7°O3'W<br />
35°48'N, 05°48'W<br />
33°55'N, Wbb'VJ<br />
33°56'N, 06°58'W<br />
345<br />
07°38'N, 10°05'E<br />
[05°34.5'N, 05°11.75'E]<br />
04°23'N, 07°06'E<br />
04°19'N, 06°14'E<br />
05°22'N, 05°26'E
346<br />
Lagos<br />
Ogudu River [creek]<br />
Port Harcourt<br />
Tarkwa Bay<br />
Victoria Beach<br />
Yaba<br />
Princesse Alice Bank<br />
Principe, Ilha do<br />
Baia das Agulhas<br />
Baia de Santo Antonio<br />
Ilheu Caroco<br />
Ilheu Santana [Sta. Ana]<br />
Hheus dos Mosteiros<br />
Morro de Praia Grande<br />
Ponta Capitao<br />
Ponta da Mina<br />
Ponta Novo Destino<br />
Ponta do Pico Negro<br />
Ponta da Praia Salgada<br />
Praia Pequena<br />
Rio Papagaio<br />
Santana<br />
Santo Antonio<br />
Tinhosa Grande [Hermano<br />
Grande]<br />
Rio Muni [Equatorial Guinea]<br />
Cabo San Juan<br />
Corisco Bay<br />
Isla de Corisco<br />
Islas de Elobey<br />
Rio Muni mouth<br />
Saint Helena<br />
James Bay<br />
Ruperts Bay<br />
Sao Tome, Ilha de<br />
Angra de Sao Joao dos Angolares<br />
Baia do Almoxarife<br />
Baia de Ana de Chaves<br />
Baia de Iogoiogo<br />
Baia de Sao Miguel<br />
Binda<br />
Ilheu das Cabras<br />
Ilheu Gago Coutinho [Ilheu<br />
das Rolas]<br />
Ilheu Macaco [ilot dos<br />
Cocos]<br />
Ilheu de Sant'Ana<br />
Morro Peixe<br />
Oquedelrei<br />
Ponta Diogo Nunes<br />
Ponta Diogo Vaz<br />
Ponta Furada<br />
Ponta de Sao Sebastiao<br />
06°27'N, 03°23'E<br />
06°34'N, 03°24'E<br />
04°46'N, 07°01'E<br />
06°24'N, 03°24'E<br />
06°25'N, 03°24'E<br />
06°32'N, 03°23'E<br />
38°00'N, 29°15'W<br />
O1°37'N, 07°25'E<br />
O1°37'N, 07°21'E<br />
01°39'N, O7°27'E<br />
01 o 31'N,07°26'E<br />
01°40'N, O7°27'E<br />
01°41'N,07°29'E<br />
01°41'N, O7°27'E<br />
01°40'N, 07°28'E<br />
01°39'N, 07°26'E<br />
Ol^N, 07°26'E<br />
01°32'N,07°24'E<br />
01°39'N, O7°27'E<br />
01°38'N, O7°27'E<br />
01°39'N, 07°25'E<br />
01°40'N, 07°26'E<br />
01°39'N, 07°26'E<br />
01°21'N, 07°18'E<br />
0ri0'N,09°21'E<br />
00°55'N, 09°25'E<br />
00°55'N, 09°19'E<br />
00°59'N, 09°30'E<br />
01°01'N, 09°36'E<br />
15°57'S, 05°42'W<br />
15°55'S, 05°43'W<br />
15°55'S, 05°43'W<br />
00°12'N, 06°39'E<br />
00°07'N, 06°39'E<br />
00°17'N, 06°45'E<br />
00°21'N, 06°44'E<br />
00°02'N, 06°33'E<br />
00°08'N, 06°30'E<br />
00°13'N, 06°28'E<br />
00°24'N, 06°43'E<br />
00°01'S, 06°32'E<br />
00 o 06'N,06°3rE<br />
00°15'N, 06°46'E<br />
00°24'N, 06°39'E<br />
OO°22'N, 06°44'E<br />
00°23'N, 06°43'E<br />
00°19'N, 06°30'E<br />
00°15'N, 06°28'E<br />
00°21'N, 06°44'E<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
Praia das Conchas<br />
Praia de Fernao Dias<br />
Praia Lagarto<br />
Praia Melao<br />
Praia Pantufo<br />
Praia de Santa Catarina<br />
Ribeira Peixe<br />
Sant'Ana<br />
Seine Seamount [Bane de la<br />
Seine]<br />
Senegal<br />
Anse Bernard<br />
Anse de Hann<br />
Baie de Dakar<br />
Baie de Goree<br />
Bakoy [?=Bako or<br />
Bakor River]<br />
Baling [?=Bafing]<br />
Bane du Seminole<br />
Bargny Gouddou<br />
Bel-Air<br />
Camberene<br />
Cap Manuel<br />
Cap Vert<br />
Casamance River<br />
Dakar<br />
Faleme<br />
Faleme River. See Callinectes<br />
amnicola<br />
Fosse de Kayar [Cayar]<br />
Goree<br />
lie de Goree<br />
lie de la Madeleine<br />
Joal<br />
Lac de Nguer [N'Guer]<br />
Lagoba. See Dromia monodi<br />
Leybar<br />
Mbao<br />
Mboro<br />
Mbour<br />
Ngaparou<br />
Ngazobil<br />
Ngor<br />
Pointe des Almadies [?Les<br />
Almadies]<br />
Pointe de Sangomar<br />
Pointe Thionk<br />
Presqu'ile du Cap Vert<br />
Rosso<br />
Rtombo. See Cardisoma<br />
armatum<br />
Rufisque<br />
Saint-Louis<br />
Saloum<br />
00°24'N, 06°37'E<br />
OO°25'N, 06°41'E<br />
0O°22'N, 06°43'E<br />
00°19'N, 06°45'E<br />
00°19'N,06°45'E<br />
00°16'N, 06°29'E<br />
00°05'N, 06°37'E<br />
00°16'N, 06°45'E<br />
33°45'N, 14°25'W<br />
14°40'N, 17°26'W<br />
14°43'N, 17°25'W<br />
14°41'N, 17°25'W<br />
14°40'N, 17°2O'W<br />
14°00'N, 15°16'W<br />
12°55'N, 14°44'W<br />
14°23'N, 16°57'W<br />
14°36'N, 17°28'W<br />
14°42'N, 17°14'W<br />
14°42'N, 17°25"W<br />
14°46'N, 17°26'W<br />
14°39'N, 17°26'W<br />
14°43'N, 17°28'W<br />
12°33'N, 16°46'W<br />
14°40'N, 17°26'W<br />
14°46'N, 12°14'W<br />
14°57'N, 17°14'W<br />
14°40'N, 17°24'W<br />
14°40'N, 17°24'W<br />
14°39'N, 17°28'W<br />
14°10'N, 16°51'W<br />
14°52'N, 14°53'W<br />
15°58'N, lG^CW<br />
14°44'N, 17°19'W<br />
15°09'N, 16°54'W<br />
14°24'N, 16°58'W<br />
14°28'N, 17°04'W<br />
14°12'N, 16°52'W<br />
14°45'N, 17°31'W<br />
14°45'N, 17°32'W<br />
13°50'N, 16°46'W<br />
12°40'N, 16°40'W<br />
14°43'N, 17°28'W<br />
16°30'N, 15°49'W<br />
14°43'N, 17°17'W<br />
16°02'N, 16°30'W<br />
13°50'N, 16°45'W
NUMBER 306<br />
Senegal River<br />
Soumb-Dioun<br />
Thiaroye-sur-Mer [Tiaroye]<br />
Sierra Leone<br />
Aberdeen<br />
Bullom Shore<br />
Cline Bay [Kline]<br />
Freetown<br />
Great Scarcies River<br />
Kissy<br />
Murray Town<br />
Robene Point<br />
Rokupr<br />
Sierra Leone River<br />
Spanish Sahara<br />
Angra da Cintra (Bay)<br />
Cabo Barbas<br />
Cabo Blanco<br />
Cabo Bojador<br />
Cabo Corbeiro [Corveiro]<br />
Islote Virginia<br />
Medano de Aaiun<br />
15° 48'N, 16°32'W<br />
3<br />
[14' 41'N, 17°28'W]<br />
14° 44'N, 17°23'W<br />
08° 30'N, 13°17'W<br />
[08' >34'N, 13°12'W]<br />
08° 29'N, 13°12'W<br />
08° 30'N, 13°15'W<br />
08° 55'N, 13°08'W<br />
08° 28'N, 13°12'W<br />
08° 30'N, 13°16'W<br />
a<br />
[08' 29'N, 13°06'W]<br />
08° 40'N, 12°23'W<br />
08° 30'N, ^"lS'W<br />
23° 00'N, 16°15'W<br />
22° 18'N, 16°41'W<br />
20° 46'N, 17°O3'W<br />
26° 08'N, 14°30'W<br />
21° 48'N, 16°58'W<br />
22° 09'N, 16°50'W<br />
27° 07'N, 13°25'W<br />
Morro Garnet [Cabo<br />
Garnet]<br />
Pulpito Bay [?Buen Jardin]<br />
Villa Cisneros<br />
Togo<br />
Lome<br />
Zaire<br />
Banana<br />
Boma<br />
Congo River<br />
lie de Mateba<br />
Katala<br />
Kunga<br />
Malela<br />
Moanda<br />
Shiloango River [Loango R.<br />
(Congo), Chiloango R.<br />
(Angola)]<br />
Tumuna. See Sesarma (Chinmantes)<br />
angolense<br />
Vista<br />
Zambi<br />
24°45'N, 14°55'W<br />
[24°41'N, 14°51'W]<br />
23°42'N, 15°56'W<br />
06°08'N, 01°13'E<br />
06°01'S, 12°24'E<br />
05°51'S, 13°03'E<br />
06°04'S, 12°24'E<br />
05°54'S, 12°50'E<br />
05°58'S, 12°44'E<br />
[ca. 05°55'S, 12°35'E]<br />
05°59'S, 12°37'E<br />
[05°55'S, 12°25'E]<br />
05°12'S, 12°O7'E<br />
347<br />
05°52'S, 12°17'E<br />
[ca. 05°52'S, 12°50'E]
Addendum<br />
After this manuscript went to the printer, Mr. C. B. Powell forwarded to us<br />
representatives of what proved to be a fourth new genus and species of<br />
camptandriine crab from <strong>West</strong> Africa; it is described here. This species raises<br />
the number of East Atlantic Ocypodidae to six genera and seven species; six<br />
of the species occur off tropical <strong>West</strong> Africa.<br />
Genus Lillyanella, new genus<br />
TYPE-SPECIES.—Lillyanella plumipes, new species.<br />
ETYMOLOGY.—The name Lillyanella is chosen in<br />
recognition of the important contribution to the<br />
present study made by Lilly King Manning, to<br />
whom we are indebted for preparation of the<br />
illustrations. Gender is feminine.<br />
DIAGNOSIS.—A genus of Camptandriinae. Carapace<br />
slightly convex, subhexagonal, broader<br />
than long, widest in the middle. Regions poorly<br />
indicated, surface even, with sparse, scattered,<br />
very short, stiff, dark pubescence; in posterolateral<br />
part some small granules. Front wide, anterior<br />
margin broadly truncate, slightly sinuous.<br />
Epigastric lobes barely visible. H-shaped groove<br />
distinct.<br />
Anterolateral margins with two large and one<br />
or more small to very small teeth far behind outer<br />
orbital angle. Posterior margins of the orbits<br />
slightly curved, almost in one line, outer half<br />
somewhat granular.<br />
Cornea well developed, slightly narrower than<br />
eyestalk. Antennules obliquely folded. Antennae<br />
short, flagellum consisting of a single segment<br />
with a long apical seta. Lower orbital margin<br />
distinct, with several distinct denticles and granules.<br />
Suborbital ridge and pterygostomian region<br />
with granules.<br />
Epistome deeply concave with raised anterior<br />
and posterior margins. Third maxilliped not fill-<br />
348<br />
ing oral field. Merus slightly longer than ischium<br />
and partly covering exopod; anteromedian part<br />
of merus with quadrangular excision. Anterior<br />
margin of ischium oblique.<br />
Chelipeds equal. Those of adult male much<br />
larger and more robust than those of female, and<br />
with molariform tooth on dactylus. Walking legs<br />
short and robust. Merus with dorsal row of teeth,<br />
of which distal and subdistal large, the following<br />
smaller but still distinct; lower surface of merus<br />
with teeth or tubercles. The other segments without<br />
teeth or denticles, some with a few granules.<br />
All legs covered with dense short dark pubescence,<br />
especially dorsally. Propodus of third leg of<br />
adult male with conspicuous tuft of long soft<br />
hairs.<br />
Male abdomen elongate triangular, only<br />
slightly constricted at fifth somite, but entirely<br />
covering the pleopods; second to fourth somites<br />
of male abdomen fused. First somite not wider<br />
than second and failing to reach the coxae of the<br />
fifth pereiopods. Male gonopods strongly recurved,<br />
slightly swollen apically with some distinct<br />
recurved spines.<br />
REMARKS.—Lillyanella differs from all other<br />
genera of Camptandriinae in the shape and armament<br />
of the ambulatory legs. In this respect it<br />
resembles the genus Leipocten most closely, in<br />
which the rneri likewise have spines on the lower<br />
surface. However, in Leipocten the large distal and<br />
subdistal dorsal teeth of the merus are absent,<br />
and the other segments of the legs are different.
NUMBER 306 349<br />
Lillyanella plumipes, new species boundary of freshwater and oligohaline area, un-<br />
FIGURE 88<br />
der bark of large sticks, 5 February 1980, C. B.<br />
Powell, \6 holotype (L), 86, 99 ov, 4 non-oviger-<br />
MATERIAL EXAMINED.—Pillsbury Material: None. ous 9, 1 juv, paratypes (L); 76, 169 (8 ov), 3 juv,<br />
Other Material: Nigeria: New Calabar River at paratypes(W).<br />
Choba bridge, 04° 54'N, 06°54'E, intertidal zone, DESCRIPTION.—Carapace subhexagonal to al-<br />
9<br />
FIGURE 88.—Lillyanella plumipes, new genus, new species. Ovigerous female: a, carapace, dorsal<br />
view; b, pterygostomian region, orbit, and third maxilliped, ventral view; c, second leg, d, third<br />
leg; e, fourth leg; /, fifth leg. Male: g, carapace, dorsal view; k, abdomen; i, first pleopod.<br />
Smaller male: j, first leg. Male holotype: *, first leg, /, propodus and dactylus of third leg. Nonovigerous<br />
female: m, first leg; n, chela.<br />
f
350<br />
most quadrangular, somewhat (about 1.2 times)<br />
broader than long. Surface slightly convex but<br />
even, without conspicuous elevations. H-shaped<br />
groove distinct, but not very deep. Epigastric<br />
lobes low and inconspicuous. Posterior half of<br />
anterolateral margin with two large teeth, between<br />
which 1 to 3 very small. Outer orbital<br />
angle sharp, about as large as the larger anterolateral<br />
teeth. Posterolateral margins with several<br />
small granules. Posterior margin almost straight,<br />
about as wide as the base of front. Front wide<br />
and not very prominent, anterior margin broad<br />
and somewhat sinuous: slightly concave in the<br />
middle, slightly convex laterally; anterolateral<br />
angle widely rounded; lateral margin evenly<br />
merging, under a broad curve, with the posterior<br />
orbital margin. Posterior orbital margins almost<br />
straight, the two lying in a practically single line;<br />
no incisions on this margin, but granules are<br />
present in its greater part. Dorsal surface of carapace<br />
with a sparse pubescence of short, stiff,<br />
curved, black hairs. The posterolateral portions<br />
of carapace sparsely granulated.<br />
Eyes almost filling the orbits, with the cornea<br />
well developed, rounded, slightly narrower than<br />
the eyestalk. Eyestalk with short, curved, stiff,<br />
dark hairs on anterior surface. Lower margin of<br />
orbit with distinct denticles and granules over its<br />
full length, and with a denticulated process near<br />
the base of the antenna. A short suborbital row<br />
of granules below outer half of orbit. Pterygostomian<br />
region granulate.<br />
Antennules covered by front, folding obliquely.<br />
Antennae short, entering orbit, the latter widely<br />
open at base. Antennal flagellum consisting of a<br />
single segment with a very long apical seta.<br />
Epistome wide, deeply concave, with anterior<br />
and posterior margins elevated.<br />
Third maxillipeds not filling oral cavity. Merus<br />
rectangularly excavated anteromedially; outer<br />
anterior angle broadly rounded. Ischium longer<br />
than merus, inner anterior angle somewhat produced<br />
forward. Exopod at base about 1/3 as wide<br />
as ischium; distal part, including multi-articulated<br />
flagellum, hidden behind merus.<br />
First pereiopods equal left and right, in adult<br />
males much larger and more robust than in fe-<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
males. Fingers of adult male cheliped somewhat<br />
shorter than palm, curved and gaping, tips ending<br />
in horn-colored hoofs; cutting edges with<br />
small denticles, in addition to which a large<br />
molar-like tooth in the dactylus. Palm in adult<br />
males almost as high as long. Surface of palm and<br />
fingers smooth, except for some minute granules<br />
on lower surface of palm. Carpus short, cupshaped,<br />
upper anterior margin with tubercles.<br />
Merus about twice as long as carpus, but still<br />
short and robust; lower surface with granulae,<br />
both inner and outer lower margin with a longitudinal<br />
row of teeth; also dorsal margin with a<br />
row of teeth; upper part of anterior margin granulate.<br />
Ischium short, lower margin with teeth.<br />
First leg of female more slender than that of male,<br />
palm more than twice as long as high; fingers as<br />
long as palm, slightly gaping with denticles along<br />
cutting edge, but no molariform tooth, tips of<br />
fingers hooved; rest of leg as in male, but more<br />
slender.<br />
First three walking legs of about the same size,<br />
the third leg (P3) slightly longer than the other<br />
two; the fifth leg shortest of all legs. Dactylus of<br />
walking legs simple with sharp, curved tip; about<br />
3/4 of length of propodus. Propodus two to three<br />
times as long as high (relatively longest in third<br />
leg), unarmed. Carpus short, about as long as<br />
dactylus, narrowing proximally, upper anterior<br />
margin sometimes with granules. Merus with dorsal<br />
margin ending in a large distal tooth and a<br />
subdistal tooth of about same size. Behind subdistal<br />
tooth a longitudinal row of up to 10 rather<br />
small but quite distinct teeth. Lower surface of<br />
merus with teeth or tubercles. Sometimes tubercles<br />
on outer surface near upper or lower margin.<br />
In adult males propodus of third pereiopod with<br />
a conspicuous large brush of very long soft hairs.<br />
In all specimens a distinct pubescence of short,<br />
curved, stiff, dark hairs on all segments of the<br />
legs, especially dorsally; here and there a few<br />
longer and softer hairs may be present.<br />
Female abdomen wide, reaching the coxae of<br />
the pereiopods; all segments free; distal segment<br />
widely triangular with broadly rounded slightly<br />
protruding top.<br />
Male abdomen elongate triangular. First som-
NUMBER 306 351<br />
ite about as wide as second, failing to reach coxae<br />
of fifth pereiopods. Second, third, and fourth<br />
somites fused; indentation in lateral margin<br />
marking separation between second and third<br />
somites. Second somite very short, only slightly<br />
longer than first. Third and fourth somites of the<br />
same length, about twice as long as second and<br />
as wide; fourth segment narrowing slightly distally.<br />
Fifth and sixth somites free. Fifth longer<br />
than fourth, constricted in the basal half, but not<br />
strongly enough to expose the gonopods. Sixth<br />
somite about as long as fifth, narrowing distally.<br />
Seventh somite as long as sixth, but much narrower,<br />
broadly rounded at apex.<br />
Male gonopod with the recurved shape characteristic<br />
of Camptandriinae, narrowing distally,<br />
but widening slightly just before the tip. Apex<br />
broadly rounded, provided with three large<br />
spines; inner part of widened subapical portion<br />
with a group of recurved spines; further numerous<br />
very small spinules in distal part of gonopod.<br />
Second pleopod of male very small, L-shaped.<br />
MEASUREMENTS.—In the adult male holotype<br />
cl is 4.2, cb 5.2 mm. The carapace width in the<br />
ovigerous females is 4.3 to 6.3 mm, in the nonovigerous<br />
females 4.0 to 6.5 mm, in the males 3.7<br />
to 6.5 mm, and in the juveniles 1.3 to 4.5 mm.<br />
The eggs are about 0.2 mm in diameter.<br />
REMARKS.—Lillyanella plumipes differs from all<br />
other <strong>West</strong> <strong>African</strong> Camptandriinae by the presence<br />
of a row of teeth on the dorsal margin of the<br />
merus of the walking legs, and by having teeth or<br />
tubercles on the lower surface of that segment.<br />
Also the peculiar brush of long setae on the<br />
propodus of the third leg of the adult males is<br />
quite characteristic. In the presence of anterolateral<br />
teeth on the carapace, Lillyanella differs from<br />
Ecphantor and resembles the two other <strong>West</strong> <strong>African</strong><br />
genera. The wide and broadly truncated<br />
front it has in common with Telmatothrix; in Calabarium<br />
the front is produced in two broadly<br />
triangular teeth separated by a V-shaped incision.<br />
The gonopods also are different from those of the<br />
other genera.<br />
TYPE-LOCALITY.—New Calabar River, slightly<br />
NW of Port Harcourt at Choba bridge, Nigeria,<br />
4°54'N 6°54'E.<br />
DISPOSITION OF TYPES.—The holotype male<br />
(Crust. D 32752) is placed in the collection of the<br />
Rijksmuseum van Natuurlijke Historie, Leiden.<br />
The paratypes are in the same museum and in<br />
the National Museum of Natural History, <strong>Smithsonian</strong><br />
<strong>Institution</strong>, Washington, D.C.<br />
ETYMOLOGY.—The name plumipes is inspired by<br />
the presence of a brush of long hairs on the<br />
propodus of the third leg of adult males.<br />
BIOLOGY.—The site at which the species was<br />
collected was described by Mr. Powell (in litt., 5<br />
February 1980) as follows:<br />
Its proper habitat is under the bark of dead tree branches<br />
etc., in the intertidal zone. The same site has Telmatothrix<br />
(common or abundant), Calabarium, Sesarma buettikoferi (abundant)<br />
and juveniles of S. angolense and S. alberti. The main<br />
vegetation is Pandanus and freshwater swamp trees, with the<br />
aquatic Crinum (subtidal) and Nymphaea (intertidal); a few<br />
specimens ofRhizophora occur, but these are small individuals<br />
which probably will not flower—there is no mature Rhizophora<br />
within sight of the bridge. The two other invertebrates<br />
occurring under the bark of the same branches are Potamalpheops<br />
monodi (Sollaud) (Alpheidae) and the large burrowing<br />
and filter-feeding mayfly nymph Povilla. The site is slightly<br />
beyond the upstream limit of isopods, amphipods and snails<br />
and bivalves. The New Calabar River is a soft-water river,<br />
lacking a freshwater mollusc fauna. Once I did find a single<br />
specimen of the oligohaline (?) snail Potamopyrgus ciliatus<br />
(Gould) at the collecting site, but have not been able to find<br />
any more. Occasionally the small and very thin specimen of<br />
the polychaete Namalycastis is found; also a red nemertean.<br />
The shrimps present are Potamalpheops monodi (Sollaud), an<br />
undescribed species of Potamalpheops, juveniles of Macrobrachium<br />
felicinum Holthuis (uncommon), M. vollenhovenii (Herklots),<br />
M. macrobrachion (Herklots) (most common) and Caridina,<br />
but no Palaemonetes.<br />
Mr. Powell remarked that the species "has the<br />
same distinctive odor as does Telmatothrix, when<br />
preserved in formalin. Also it sheds its legs much<br />
more readily than any other crab I've met."<br />
Ovigerous females were collected in February.<br />
DISTRIBUTION.—Known only from the type locality.
352 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
ADDENDUM TO KEY<br />
Our key to the genera of Camptandriinae (p. 193) can be altered to include<br />
this additional new genus by substituting the following two couplets for<br />
couplet 8 and changing couplets 9-11 to 10-12.<br />
8. Merus of pereiopods with blunt spines or teeth on lower (flexor)<br />
surface 9<br />
Merus of pereiopods without ventral spines or teeth 10<br />
9. Upper margin of merus of pereiopods granular, without large distal or<br />
subdistal teeth; lower margin of propodus of these legs ending in a<br />
tooth. Carapace granular all over, without large anterolateral teeth<br />
Leipocten<br />
Upper margin of merus of pereiopods ending in a large distal tooth and<br />
a subdistal tooth of the same size; lower margin of propodus without a<br />
distal tooth. Carapace smooth for the larger part, only granular in the<br />
posterolateral area; anterolateral margin with 2 large and some small<br />
teeth Lillyanella, new genus
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Halfte 1895/96-1897/98. Zoologische Ergebnisse
354<br />
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Klasse, 102:1-30, figures 1-9, plate 1.<br />
1932. Uber einige systematisch interessante Xanthidae<br />
(Crustacea Decapoda Brachyura) der Harmsschen<br />
Reisen nach dem Sundaarchipel. Zeitschrift for wissenschaftliche<br />
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1933a. Uber zwei interessant Xanthidae (Crustacea Dekapoda)<br />
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1954. New Records and New Species of Crustacea from<br />
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1888. Catalogue des Crustaces marins recueillis aux Acores<br />
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1835. Some Account of the Crustacea of the Coasts of<br />
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1855. A Monograph of the Leucosiadae, with Observations<br />
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1893. Notice of the Crustaceans Collected by the United<br />
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1968c. Les Carpilinae: Recherches preliminaires sur les<br />
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1969b. Les Goneplacidae (suite): Recherches preliminaires<br />
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1969c. Les Goneplacidae (suite et fin): Recherches preliminaires<br />
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1969d. Sur divers Xanthidae notamment sur Actaea de<br />
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1963. Remarques sur les Genres Cleistostoma, Paracleistostoma<br />
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1958. An Annotated List of the Decapod Crustacea of<br />
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1957. The Genus Cancrincola (Copepoda, Harpacticoida)<br />
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1913. Dromiacea: Die Decapoda Brachyura der Siboga-<br />
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1970. Invertebrate Zoology. Volume 3, 523 pages. New<br />
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1915. Crustacea Decapoda: Fauna of the Chilka Lake.<br />
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1970. Some Decapod Crustacea from Northern South-<br />
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1856. Remarks on the Habits and Distribution of Marine<br />
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1880a. Synopsis of the Grapsidae: Carcinological Notes,<br />
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1880b. On a Collection of Crustacea from Virginia, North<br />
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1967. A Find of Geryon affinis Milne-Edwards & Bouvier,<br />
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1892. Beitrage zur Kenntnis der Crustaceen der Canarischen<br />
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1878. Kurze Notizen iiber einige neue Crustaceen sowie<br />
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1818. Histoire naturelle des animaux sans vertebres, presentant<br />
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1810. Considerations generates sur Vordre naturel des animaux<br />
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1815a. A Tabular View of the External Characters of<br />
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1963. Premieres donnees sur la regulation osmotique et<br />
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218.<br />
1966. Variations du serum et de 1'urine en fonction de<br />
la salinite chez le crabe terrestre Cardisoma armatum<br />
Herklots. Memoires de la Socie'te des sciences naturelles<br />
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figures 1-11.<br />
Le Loeuff, P., and A. Intes<br />
1968. La faune benthique du plateau continental de<br />
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saisonnieres (Mars 1966-<br />
Fevrier 1967). Centre de Recherches Oceanographiques,<br />
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1-63.<br />
1969. Premieres observations sur la faune benthique du<br />
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Le Loeuff, P., A. Intes, and J. C. Le Guen<br />
1974. Note sur les premiers essais de capture du crabe<br />
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1910. Dekapode Crustaceen Aquatorialafrikas. Wissenschaftliche<br />
Ergebnisse der Deutschen Zentral-Afrika-Expedition<br />
1907-1908 unter Fuhrung Adolf Friedrichs,<br />
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121-134, plate 3. [Pages 1-14 on separate.]<br />
Lenz, H., and K. Strunck<br />
1914. Brachyuren und Macruren mit Ausschluss der<br />
Sergestiden: Die Dekapoden der Deutschen Siidpolar-Expedition<br />
1901-1903, I. Deutsche Siidpolar-<br />
Expedition 1901-1903, 15(Zoologie VII): 257-345,<br />
figures 1-5, plates 12-22.<br />
Leone, Charles A.<br />
1949. Comparative Serology of Some <strong>Brachyuran</strong> Crustacea<br />
and Studies in Hemocyanin Correspondence.<br />
Biological Bulletin, 97(3):273-286, figures<br />
1-3.<br />
1951. A Serological Analysis of the Systematic Relationship<br />
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Biological Bulletin, 100(l):44-48.<br />
Lewinsohn, Ch.<br />
1977. Die Dromiidae des Roten Meeres (Crustacea Decapoda.<br />
Brachyura). Zoologische Verhandelingen, uitgegeven<br />
door het Rijksmuseum van Natuurlijke Historie te<br />
Leiden, 151:1-41, figures 1-9, plate 1.<br />
Lewinsohn, Ch., and L. B. Holthuis<br />
1964. New Records of Decapod Crustacea from the<br />
Mediterranean Coast of Israel and the Eastern<br />
Mediterranean. Zoologische Mededelingen, uitgegeven<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
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40(8):45-63, figures 1-5.<br />
Lewis, Elizabeth G., and Paul A. Haefner, Jr.<br />
1978. Intraspecific Setal Variation in Bathynectes superbus<br />
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1747. Wdstgota-Resa, pa Riksens Hogloflige Slanders Befallning<br />
forrattad ar 1746. Med Anmdrkningar uti Oeconomien,<br />
Naturkunnogheten, Antiquiteter, Inwanames Seder<br />
och Lefnads-Sdtt, med tilhorige Figurer. xi + 284 and<br />
20 unnumbered pages, plates 1-5. Stockholm.<br />
1754. Museum S:ae R:ae M:tis Adolphi Friderici Regis Suecorum,<br />
Gothorum, Vandalorumque. Haer. Now. Due. Slesv.<br />
Hols. Storm. Ditm. Com. Oldenb. Delmenhorstiae. &c.<br />
&c. in quo animalia rariora imprimis, et exotica: Quadrupedia,<br />
Aves, Amphibia, Pisces, Insecta, Vermes describuntur<br />
et determinantur, latine et suetice cum Iconibus.<br />
Hans Maj:ts Adolf Frideriks var allernadigste Konungs<br />
Naturalie Samling innehallande sdlsynte och frdmmande<br />
Djur, som bevaras pa Kongl. Lust-Slottet Ulriksdahl;<br />
beskrefne och afritade samt pa nadig Befallning. xxx +<br />
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1758. Systema Naturae per Regna Tria Naturae, Secundum<br />
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Differentiis, Synonymis, Locis. Edition 10, volume 1,<br />
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1761. Fauna Suecica sistens Animalia Sueciae Regni: Mammalia,<br />
Aves, Amphibia, Pisces, Insecta, Vermes. Distributa<br />
per Classes & Ordines, Genera & Species, cum<br />
Differentiis Specierum, Synonymis Auctorum, Nominibus<br />
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pages 1-578.<br />
1767. Systema naturae per regna tria naturae, secundum classes,<br />
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1877. Remarks on the Crustacea of the Pacific Coast of<br />
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1846. Crustaces, Arachnides, Myriopodes et Hexapodes:<br />
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NUMBER 306 367<br />
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1838. On the Brachyurous Decapod Crustacea Brought<br />
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1879. On Some New or Imperfectly Known Podophthalmous<br />
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1881. Carcinological Studies in the Leyden Museum,<br />
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1883. Carcinological Studies in the Leyden Museum,<br />
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1888. Bericht iiber die im indischen Archipel von Dr. J.<br />
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Archivfir Naturgeschichte, 53:215-600, plates<br />
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1892. Decapoden des Indischen Archipels. In Max Weber,<br />
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1895a. Bericht iiber die von Herrn Schiffscapitan Storm<br />
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1895b. Bericht iiber die von Herrn Schiffscapitan Storm<br />
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Borneo und Celebes sowie in der Java-See gesammelten<br />
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1896. Platychirograpsus spectabilis, nov. gen. n. sp., eine<br />
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1897. Bericht iiber die von Herrn Schiffscapitan Storm<br />
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Borneo und Celebes sowie in der Java-See gesammelten<br />
Decapoden und Stomatopoden. Fiinfter<br />
Theil. Zoologische Jahrbiicher, Abtheilung fur Sys-<br />
tematik, Geographie und Biologie der Thiere, 9:725-790,<br />
plates 12-14.<br />
1900. Note sur une petite collection de Crustaces Decapodes,<br />
provenant de la Cote d'Angola (Afrique<br />
Occidentale). Me'moires de la Socie'te' Zoologique de<br />
France, 13:31-65, plates 1, 2.<br />
1902. Description d'une espece nouvelle du genre Heteropanope<br />
Stimpson, provenant de la Cote d'lvoire,<br />
Heteropanope africana n. sp. Bulletin du Museum<br />
d'Histoire naturelle (Paris), 4:254-260, figures 1, 2.<br />
1906. Diagnoses of Five New Species of Decapod Crustacea<br />
and of the Hitherto Unknown Male of<br />
Spirontocans rectirostris (Stimps.) from the Inland<br />
Sea of Japan, as also of a New Species of Palaemon<br />
from Darjeeling, Bengal. Annals and Magazine of<br />
Natural History, series 7, 17:400-406.<br />
1913. Sur une nouvelle observation de Crabes habitant<br />
les coquilles vides des Balanes. Bulletin du Museum<br />
d'Histoire naturelle (Paris), 19:9-11.<br />
Marchand, Lewis J.<br />
1946. The Saber Crab, Platychirograpsus typicus Rathbun,<br />
in Florida: A Case of Accidental Dispersal. Quarterly<br />
Journal of the Florida Academy of Sciences, 9(2):<br />
93-100, figures 1, 2.<br />
Von Martens, E.<br />
1872. Ueber Cubanische Crustaceen nach den<br />
Sammlungen Dr. J. Gundlach's. Archiv fur Naturgeschichte,<br />
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Mason, J., and C. Davidson<br />
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Massuti, M.<br />
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1968b. Les crustaces captures par la "Thalassa" au large<br />
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368<br />
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1875. St. Helena: A Physical, Historical, and Topographical<br />
Description of the Island, including its Geology, Fauna,<br />
Flora, and Meteorology, xiv + 426 pages, frontispiece,<br />
plates 1-56 (not all numbered). London.<br />
Miers, Edward J.<br />
1876. Catalogue of the Stalk- and Sessile-eyed Crustacea of New<br />
Zealand, xii + 136 pages, plates 1-3. London.<br />
1878. Revision of the Plagusiinae. Annals and Magazine of<br />
Natural History, series 5, 1:147-154.<br />
1879a. On the Classification of the Maioid Crustacea or<br />
Oxyrhyncha, with a Synopsis of the Families,<br />
Subfamilies, and Genera. Journal of the Linnean<br />
Society, Zoology, 14:634-673, plates 12, 13.<br />
1879b. Descriptions of New or Little Known Species of<br />
Maioid Crustacea (Oxyrhyncha) in the Collection<br />
of the British Museum. Annals and Magazine of<br />
Natural History, series 5, 4:1-28, plates 4, 5.<br />
1881a. On a Collection of Crustacea made by Baron<br />
Hermann Maltzam [sic] at Goree Island, Senegambia.<br />
Annals and Magazine of Natural History,<br />
series 5, 8:204-220, 259-281, 364-377, plates 13-<br />
16.<br />
1881b. Crustacea. In Report on a Collection Made by Mr.<br />
T. Conry in Ascension Island. Annals and Magazine<br />
of Natural History, series 5, 8:432-434, 2 figures.<br />
1886. Report on the Brachyura Collected by H. M. S.<br />
Challenger during the Years 1873-1876. Report on<br />
the Scientific Results of the Voyage of H. M. S. Challenger<br />
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1861. Etudes zoologiques sur les Crustaces recents de la<br />
famille des Portuniens. Archives du Museum d'Histoire<br />
naturelle (Paris), 10:309-428, 2 unnumbered pages,<br />
plates 28-38.<br />
1862a. Monographic des Crustaces fossiles de la famille<br />
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series 4, 18:31-85, plates 1-10.<br />
1862b. Faune carcinologique de ltle de la Reunion, Annexe<br />
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1863. Monographic des Crustaces fossiles de la famille<br />
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1865. Cancerides, Pirimclides, Carpilides, Premiere Partie:<br />
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Museum d'Histoire naturelle (Paris), 1:177-308, plates<br />
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1867a. Descriptions des especes nouvelles de Crustaces<br />
provenant dc Saint-Vincent du Cap Vert. In L.<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
de Folin and L. Perier, Les Fonds de la Mer, 1867-<br />
1872, volume 1, chapter 9, pages 32-44, plates 21-<br />
21 bis. [According to Rehder (1946:74, 75) volume<br />
1 of Les Fonds de la Mer was published in six parts,<br />
as follows: pages 1-48, 1867; pages 49-112, 1868;<br />
pages 113-176, 1869; pages 177-256, 1870; pages<br />
257-272, 1871; pages 273-316, 1872.]<br />
1867b. Descriptions de quelques especes nouvelles de<br />
Crustaces Brachyures. Annales de la Socie'te entomologique<br />
de France, series 4, 7:263-288.<br />
1868. Observations sur la faune carcinologique des lies<br />
du Cap-Vert. Nouvelles Archives du Museum d'Histoire<br />
naturelle (Paris), 4:49-68, plates 16-18.<br />
1869. Description de quelques especes nouvelles de Crustaces<br />
provenant du voyage de M. A. Bouvier aux<br />
lies du Cap Vert. Revue el Magasin de Zoologie, 21:<br />
350-355, 374-378, 409-412 [pages 1-13 on separate].<br />
1873a. Description de quelques Crustaces nouveaux ou<br />
peu connus provenant du Musee de M. C. Godeffroy.<br />
Journal des Museum Godeffroy, 1(4): 77-88,<br />
plates 1, 2. [Pages 1-12, plates 12, 13 on separate.]<br />
1873b. Recherches sur la faune carcinologique de la Nouvelle-Caledonie,<br />
Deuxieme Partie. Nouvelles Archives<br />
du Museum d'Histoire naturelle (Paris), 9:155-<br />
332, plates 4-18.<br />
1873-1881. Etudes sur les Xiphosures et les Crustaces de<br />
la region Mexicaine. In Mission scientifique au Mexique<br />
et dans CAmerique centrale, Recherches Zoologiques<br />
pour servir a Fhistoire de la faune de VAmerique centrale<br />
et du Mexique, 5:1-368, plates 1-61. [See Monod<br />
(1956:642) for dates of publication. Monod noted<br />
that he could not trace the date of publication of<br />
plate 44. In the edition in the Division of Crustacea<br />
library, <strong>Smithsonian</strong> <strong>Institution</strong>, plate 44 was<br />
published in 1880.]<br />
1878. Description de quelques especes nouvelles de Crustaces<br />
provenant du voyage aux iles du Cap-Vert<br />
de MM. Bouvier et de Cessac. Bulletin de la Socie'te<br />
Philomathique de Paris, series 7, 2:225-232 [pages 6-<br />
13 on separate].<br />
1880. Etudes preliminaires sur les Crustaces, l ere Partie.<br />
In Reports on the Results of Dredging under the<br />
Supervision of Alexander Agassiz, in the Gulf of<br />
Mexico, and in the Caribbean Sea, 1877, '78, '79,<br />
by the U. S. Coast Survey Steamer "Blake,"<br />
Lieut.-Commander C. D. Sigsbee, U.S.N., and<br />
Commander J. R. Bartlett, U.S.N., Commanding.<br />
Bulletin of the Museum of Comparative Zoology at Harvard<br />
College, 8(1): 1-68, plates 1, 2.<br />
1883. Recueil de figures de Crustaces nouveaux ou peu connus.<br />
Part 1, pages 1-3, plates 1-44. Paris.
NUMBER 306 369<br />
Milne Edwards, A., and E.-L. Bouvier<br />
1894. Brachyures et Anomoures. In Crustaces decapodes<br />
provenant des campagnes du yacht I'Hirondelle<br />
(1886, 1887, 1888), Premiere Partie. Resultats des<br />
Campagnes Scientifiques accomplies sur son yacht par<br />
Albert F r , Prince Souverain de Monaco, 7:1-112, 8<br />
figures, plates 1-11.<br />
1897. Crustaces nouveaux provenant des campagnes du<br />
Travailleur et du Talisman. Bulletin du Museum<br />
d'Histoire naturelle (Paris), 3:297-301.<br />
1898. Crustaces nouveaux provenant des campagnes du<br />
Travailleur et du Talisman. Bulletin du Museum<br />
d'Histoire naturelle (Paris), 4:32-35, 75-77, 152-154,<br />
183-190, 234-238.<br />
1899. Crustaces Decapodes provenant des campagnes de<br />
I'Hirondelle (Supplement) et de la Princesse-Alice<br />
(1891-1897). Resultats des Campagnes Scientifiques<br />
accomplies sur son yacht par Albert f r , Prince Souverain<br />
de Monaco, 13:1-106, plates 1-4.<br />
1900. Brachyures et Anomoures. In Crustaces Decapodes,<br />
Premiere Partie. Expeditions Scientifiques du<br />
Travailleur et Talisman pendant les Annies 1880,<br />
1881, 1882, 1883. 396 pages, 32 plates. Paris.<br />
1902. Les Dromiaces et Oxystomes. In Reports on the<br />
Results of Dredging, under the Supervision of<br />
Alexander Agassiz, in the Gulf of Mexico (1877-<br />
78), in the Caribbean Sea (1878-79), and along<br />
the Atlantic Coast of the United States (1880), by<br />
the U.S. Coast Survey Steamer "Blake," Lieut.-<br />
Com. C. D. Sigsbee, U.S.N., and Commander J.<br />
R. Bartlett, U.S.N., Commanding, XXXIX. Memoirs<br />
of the Museum of Comparative Zoology at Harvard<br />
College, 27(1): 1-127, plates 1-25.<br />
1923. Les Porcellanides et des Brachyures. In Reports on<br />
the Results of Dredging, under the Supervision of<br />
Alexander Agassiz, in the Gulf of Mexico (1877-<br />
78), in the Caribbean Sea (1878-79), and along<br />
the Atlantic Coast of the United States (1880), by<br />
the U.S. Coast Survey Steamer "Blake," Lieut.-<br />
Com. C. D. Sigsbee, U.S.N., and Commander J.<br />
R. Bartlett, U.S.N., Commanding. XLVII. Memoirs<br />
of the Museum of Comparative Zoology at Harvard<br />
College, 47(4):281-395, figures 1-23, plates 1-12.<br />
Milne Edwards, H.<br />
1834. Histoire naturelle des Crustaces, comprenant Vanatomic, la<br />
physiologie et la classification de ces animaux. Volume<br />
1: xxxv + 468 pages. Paris.<br />
1836-1844. Les Crustaces. In G. Cuvier, Le regne animal,<br />
distribue dapres son organisation, pour servir de base a<br />
rhistoire naturelle des animaux, et a"introduction a<br />
Canaiomie comparee, 278 pages. Atlas, plates 1-80.<br />
Paris. [According to Cowan (1976:60), the text<br />
and plates of this work were published in 23 parts<br />
(livraisons) between September 1836 and March<br />
1844. These parts are summarized here as follows:<br />
main part number (equivalent Crustacea part<br />
number): pages, plates, (date): 8(1): 1-8, pis. 1, 7,<br />
22, 26, September 1836. 20(2):9-16, pis. 2, 17, 24,<br />
41, March 1837. 33(3): 17-24, pis. 25, 42, 47, 48,<br />
October 1837. 38(4):25-32, pis. 37, 38, 44, 50,<br />
January 1838. 51(5):33-4O, pis. 8, 20, 29, 33, July<br />
1838. 60(6) :41-48, pis. 9, 21, 27, 40, November<br />
1838. 70(7):49-56, pis. 23, 30, 31, 45, April 1839.<br />
73(8):57-64, pis. 43, 46, 58, 61, June 1839. 87(9):<br />
65-72, pis. 28, 49, 65, 66, January 1840. 90(10):<br />
73-80, pis. 3, 36, 71, IXbis, February 1840. 95(11):<br />
81-88, pis. 19, 32, 34bis, 67, May 1840. 99(12) :89-<br />
96, pis. 10, llbis, 34, 51, July 1840. 107(13):97-<br />
104, pis. 4, 12, 76, November 1840. 110(14): 105-<br />
112, pis. 52, 53, 55, 59, December 1840. 118(15):<br />
113-120, pis. 5, 18, 54, bbbis, April 1841. 120(16):<br />
121-128, pis. 16, 39, 56, 77, May 1841. 149(17):<br />
129-136, pis. 6, 15, 35, 60, July 1842. 157(18):<br />
137-144, pis. 11, 14, 64, 69, November 1842.<br />
176(19): 145-176, pis. 13, 68, 70, July 1843.<br />
180(20): 177-184, pis. 57, 62, 62bis, 63, August<br />
1843. 184(21): 185-216, pis. 54bis, 74, 75, October<br />
1843. 188(22):217-248, pis. 70bis, 72, 78, December<br />
1843. 194(23):249-278 + 5, pis. 73, 79, 80,<br />
March 1844.]<br />
1837. Histoire naturelle des Crustaces, comprenant Panatomie, la<br />
physiologie et la classification de ces animaux. Volume<br />
2, 532 pages. Atlas [1834, 1837, 1840]: 32 pages,<br />
plates 1-14, 14bis, 15-25, 25bis, 26-42. Paris. [See<br />
Holthuis, 1979c for dates of publication.]<br />
1852. De la famille des ocypodides (Ocypodidae). Second<br />
Memoire. In Observations sur les affinites zoologiques<br />
et la classification naturelle des Crustaces.<br />
Annales des Sciences Naturelles, series 3 (Zoology), 18:<br />
128-166, plates 3, 4. [Reprinted with separate<br />
pagination in undated Melanges Carcinologiques<br />
(pages 73-128), along with other articles; p. 92 in<br />
Melanges = page 128 of original.]<br />
1853. Memoires sur la famille des Ocypodiens, suite.<br />
Annales des Sciences Naturelles, series 3 (Zoology), 20:<br />
163-228, plates 6-11. [A continuation of H. Milne<br />
Edwards, 1852, and reprinted with it in undated<br />
Melanges Carcinologiques, pages 129-1%.]<br />
1854. Notes sur quelques Crustaces nouveaux ou peu<br />
connus conserves dans la collection du Museum<br />
d'Histoire Naturelle. Archives du Museum d'Histoire<br />
naturelle (Paris), 7:145-192, plates 9-16. [Although<br />
volume 7 of the Archives is dated 1854-1855, the<br />
reprint of this article carries the date 1854, and<br />
the paper must have been published in that year,<br />
even though it is cited by page in H. Milne<br />
Edwards (1853). In Neave's Nomenclator Zoologicus<br />
the present paper is dated 1853, and the generic
370<br />
names proposed in it are given precedence over<br />
those published in the previous paper (H. Milne<br />
Edwards, 1853). Apart from the fact that H. Milne<br />
Edwards in his 1853 paper cited the pages of his<br />
1854 paper, we cannot find any evidence that the<br />
latter was published earlier than indicated both<br />
on the volume and on the reprint. H. Milne<br />
Edwards in 1853 may have had page proofs of the<br />
article that was published in 1854 and cited page<br />
numbers from those proofs.]<br />
Milne Edwards, H., and H. Lucas<br />
1843. Crustaces. In A. d'Orbigny, Voyage dans PAme'rique<br />
meridionale (le Bre'sil, la republique orientate de<br />
FUruguay, la republique Argentine, la Patagonie, la<br />
republique du Chili, la republique de Bolivia, la republique<br />
du Perm), execute pendant les ounces 1826, 1827, 1828,<br />
1829, 1830, 1831, 1832 et 1833, 6(1): 1-37, plates<br />
1-17. Strasbourg. [Plates 10-17 possibly published<br />
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Monod, Th.<br />
1927. Decapoda (excl. Palaemonidae, Atyidae et Potamonidae),<br />
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624, figures 1-3.<br />
1928. L'industrie des peches au Cameroun, Commissariat<br />
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Mission Monod (1925-1926), 1 (Generalites): 1-504,<br />
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1933b. Sur quelques Crustaces de PAfrique occidentale<br />
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1946. Sur la presence du genre Acanthocarpus dans<br />
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1956. Hippidea et Brachyura ouest-africains. Memoires de<br />
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1967. Crevettes et crabes de la cote occidentale<br />
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1970. John Cranch, Zoologiste de PExpedition du Congo<br />
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Descrizione de'Crostacei, de'Testacei e de' Pesci che<br />
abitano le lagune e golfo Veneto rappresentati in figure, a<br />
chiaroscuro ed a colori dalV Abate Stefano Chiereghini<br />
Ven. Clodiense applicata per commissione govemativa. xi<br />
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1847b. Prospetto della fauna marina volgare del Veneto<br />
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1869. Annotazioni illustranti cinquantaquattro specie di<br />
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1973. First Records of the Chinese Mitten Crab, Eriocheir<br />
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Neumann, R.<br />
1878. Systematische Uebersicht der Gattungen der Oxyrhynchen:<br />
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1936. Crustaceos Decapodes e Stomatopodes Marinhos<br />
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1965. Notes on Hexapus sexpes (Fabricius) Obtained from<br />
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Krebse des Strassburger Museums, mit besonderer<br />
Beriicksichtigung der von Herrn Dr. Doderlein bei<br />
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Formen, VI. Theil. Zoologische Jahrbucher, Abtheilung<br />
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1893b. Abtheilung: Brachyura (Brachyura genuina Boas),<br />
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1. Gruppe: Cyclometopa. Die Decapoden-<br />
Krebse des Strassburger Museums, mit besonderer<br />
Beriicksichtigung der von Herrn Dr. Doderlein bei<br />
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1894. Abtheilung: Brachyura (Brachyura genuina<br />
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Cancrinea, 2. Gruppe: Catametopa. Die Decapoden-Krebse<br />
des Strassburger Museums, mit<br />
besonderer Beriicksichtigung der von Herrn Dr.<br />
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gesammelten und zur Zeit im Strassburger Museum<br />
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1899. Crustacea, Zweite Halfte: Malacostraca. In H. G.<br />
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del Golfo e delle Lagune di Venezia; preceduto da una<br />
Folkslags Sprdk, Seder, Hushallning, m. m. 6 unnumbered<br />
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Sciencias mathematicas, physicas e naturaes (Lisboa),<br />
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Museums, mit besonderer Beriicksichtigung der<br />
d'Afrique Occidentale dans les collections du Mu-<br />
von Herrn Dr. Doderlein bei Japan und bei den<br />
seum d'Histoire naturelle de Lisbonne (Suite).<br />
Liu-Kiu-Inseln gesammelten und z.Z. im Strass-<br />
Jornal de Sciencias mathematicas, physicas e naturaes<br />
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faune carcinologique des ties Saint Thome et du<br />
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Prince. Jornal de Sciencias mathematicas, physicas e<br />
1893a. Abtheilung: Brachyura (Brachyura genuina<br />
naturaes (Lisboa), series 2, 1:129-139.
372<br />
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY<br />
1890. Note sur quelques especes de Crustaces des lies S.<br />
ricas 1873-1876, 3(1): 1-304, figures, 1 map. Leip-<br />
Thome, du Prince et Ilheo das Rolas. Jomal de<br />
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Sacncias mathematicas, physicas e naturaes (Lisboa), Pennant, T.<br />
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1777. Crustacea, Mollusca, Testacea. British Zoology, edi-<br />
1892. Nova contribuicao para a fauna carcinologica da<br />
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1895a. Crustaceos da Africa Occidental Portugueza. Jor-<br />
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1833. Su di alcuni nuovi crustacei dei mari di Messina.<br />
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1839. Descrizione di due nuovi Crustacei dei Mari di<br />
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1974. Die Unterfamilie Macrophthalminae Dana im<br />
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1977. Die Macrophthalminae des Zoologischen Instituts<br />
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1814. Precis des de'couvertes et travaux somiologiques de Mr. C.<br />
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1815. Analyse de la nature ou tableau de I'univers et de corps<br />
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1818. [Review of] Journal of the Academy of Natural<br />
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Ramadan, Sh. E., and N. M. Dowidar<br />
1976. Brachyura (Decapoda Crustacea) from the Mediterranean<br />
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1840. Catalogue of the Crustacea Brought by Thomas<br />
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New, among Which Are Included Several Species<br />
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Collection of the Academy. Journal of the Academy<br />
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1893a. Catalogue of the <strong>Crabs</strong> of the Family Maiidae in<br />
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1893b. Scientific Results of Explorations by the U. S. Fish<br />
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1896. The Genus Callinectes. Proceedings of the United States<br />
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1897a. The <strong>African</strong> Swimming <strong>Crabs</strong> of the Genus Callinectes.<br />
Proceedings of the Biological Society of Washington,<br />
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1897b. A Revision of the Nomenclature of the Brachyura.<br />
Proceedings of the Biological Society of Washington, 11:<br />
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1898. The Brachyura of the Biological Expedition to the<br />
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