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Abstract and Figures

The Caribbean waters of Venezuela are composed by a large variety of habitats, with over 2800 km of coastline, islands, and islets. This area is a transitional zone between two main biogeographic provinces, the Caribbean and the Brazilian, separated by the fresh water outflows of the Orinoco and Amazon rivers, and is therefore expected to be an area of high species diversity. However, concerning the study of molluscs, Venezuela is probably the poorest known region in the Caribbean. The best compilation of opisthobranch species known in Venezuela was produced almost a decade ago, mentioning the occurrence of 57 species, plus seven determined only to genus level. In this work, 134 species are reported for Venezuela (71 are illustrated), representing about 40 % of the entire diversity of opisthobranchs known in the Caribbean. Among the species occurring in Venezuela, 49 have here the southern limit of their distribution range and only one the northern limit. Forty-six species are recorded for the first time to the country and one is a new record for the Caribbean Sea, namely Placida cremoniana. In addition, the distribution and ecology of the species are given based in literature and new data.
Content may be subject to copyright.
Accepted by M. Schroedl: 28 Sept. 2015; published: 29 Oct. 2015
ZOOTAXA
ISSN 1175-5326 (print edition)
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Copyright © 2015 Magnolia Press
Zootaxa 4034 (2): 201
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The opisthobranch gastropods (Mollusca: Heterobranchia) from Venezuela:
an annotated and illustrated inventory of species
MANUEL CABALLER GUTIÉRREZ
1,2
, JESÚS ORTEA
3
, NELSY RIVERO
4
, GABRIELA CARIAS TUCKER
5
,
MANUEL ANTÓNIO E. MALAQUIAS
6
& SAMUEL NARCISO
7
1
Muséum national d’Histoire naturelle. 55 rue Buffon, 75005 Paris. E-mail: mcaballergutierrez@mnhn.fr
2
Department of Oceanology and Coastal Sciences, Venezuelan Institute for Scientific Research, Carretera Panamericana km 11,
Miranda, Venezuela. E-mail: manuelcaballergutierrez@hotmail.com
3
Department BOS, Faculty of Biology, University of Oviedo, Asturias, Spain. E-mail: jortea@uniovi.es
4
Environmental Sciences Headship, Academic Vice-Chancellorship, Caribbean Maritime Experimental National University, Ave. El
Ejercito, Catia La Mar, Vargas, Venezuela. E-mail: nelsyrivero@gmail.com
5
Clinical laboratory, Policlínica Metropolitana Urbanización Caurimare, Calle A-1, Caracas, Venezuela.
E-mail: gabycariast@gmail.com
6
Phylogenetics Systematics and Evolution Research Group, Section of Taxonomy and Evolution, Department of Natural History, Uni-
versity Museum Bergen, University of Bergen, PB 7800, 5020-Bergen, Norway. E-mail: Manuel.Malaquias@uib.no
7
Centro de Investigación y Atención Comunitaria (CIAC) de la Fundación para la Defensa de la Naturaleza (FUDENA), Calle Cara-
bobo s/n, Chichiriviche, Falcon, Venezuela. E-mail: samuelnarciso@gmail.com
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 206
Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .207
Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Class Gastropoda Cuvier, 1795 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .209
SubClass Heterobranchia Burmeister, 1837 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Clade Tectipleura Schrödl, Jörger, Klussmann-Kolb & Wilson, 2011 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Superorder Euopisthobranchia Jörger, Stöger, Kano, Fukuda, Knebelsberger & Schrödl, 2010 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Order Cephalaspidea P. Fischer, 1883 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Family Aglajidae Pilsbry, 1895 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Genus Migaya Ortea, Caballer & Espinosa, 2014. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
1*. Migaya felis (Er. Marcus & Ev. Marcus, 1970). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210
Genus Navanax Pilsbry, 1895a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210
2. Navanax gemmatus (Mörch, 1863) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210
Genus Chelidonura A. Adams, 1850. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210
3*. Chelidonura hirundinina (Quoy & Gaimard, 1833) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210
4*. Chelidonura cubana Ortea & Martínez, 1997 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
Family Cylichnidae H. Adams & A. Adams, 1854 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
Genus Cylichnella Gabb, 1873 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
5. Cylichnella bidentata (d´Orbigny, 1841) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
Genus Acteocina J. E. Gray, 1847 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
6. Acteocina candei (d´Orbigny, 1841) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
Genus Cylichna Lovén, 1846 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
7. Cylichna krebsii Mörch, 1875 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
Family Retusidae Thiele, 1925 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
Genus Retusa Brown, 1827 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
8. Retusa sulcata (d´Orbigny, 1841) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212
Genus Pyrunculus Pilsbry, 1895 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212
9. Pyrunculus caelatus (Bush, 1885) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212
Family Rhizoridae Dell, 1952 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212
Genus Vo l vu l e l l a Newton, 1891 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212
10. Volvulella minuta (Bush, 1885) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212
11. Volvulella persimilis (Mörch, 1875) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212
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12. Volvulella texasiana Harry, 1967. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212
Family Gastropteridae Swainson, 1840. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212
Genus Gastropteron Kosse, 1813 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212
13. Gastropteron chacmol Gosliner, 1989 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212
Family Philinidae Gray, 1850 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
Genus Philine Ascanius, 1772 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .213
14. Philine sagra (d'Orbigny, 1841) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
15. Philine buchensis Caballer & Ortea, 2015 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
Family Scaphandridae Sars, 1878 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
Genus Scaphander Montfort, 1810 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
16. Scaphander watsoni Dall, 1881 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
17. Scaphander nobilis Verril, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
Family Bullidae Gray, 1827. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
Genus Bulla Linnaeus, 1758 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
18. Bulla occidentalis A. Adams, 1827 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
Family Haminoeidae Pilsbry, 1895b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214
Genus Haminoea Turton & Kingston, 1830 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .214
19. Haminoea cf. elegans (Gray, 1825). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214
20. Haminoea antillarum (d´Orbigny, 1841) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214
21. Haminoea succinea (Conrad, 1846) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
22. Haminoea petitii (d'Orbigny, 1841) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
Genus Atys Montfort, 1810 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
23. Atys caribaeus (d´Orbigny, 1841) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
24. Atys guildingi Sowerby II, 1869 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
25. Atys riiseanus Mörch, 1875 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .215
26. Atys sharpi Vanatta, 1901 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
Order Runcinacea Burn, 1963 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
Family Runcinidae H. Adams & A. Adams, 1854 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
Genus Lapinura Er. Marcus & Ev. Marcus, 1970 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
27*. Lapinura divae (Ev. Marcus & Er. Marcus, 1963) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
Order Umbraculida Odhner, 1939 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .216
Family Umbraculidae Dall, 1889 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
Genus Umbraculum Schumacher, 1817 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .216
28. Umbraculum umbraculum (Lightfoot, 1786) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .216
Order Anaspidea Fischer, 1883 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
Family Aplysiidae Lamarck, 1809 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
Genus Aplysia Linnaeus, 1767 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
29. Aplysia dactylomela Rang, 1828. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
30. Aplysia brasiliana Rang, 1828 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
31. Aplysia cervina (Dall & Simpson, 1901) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
32. Aplysia juliana Quoy & Gaimard, 1832 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
33. Aplysia morio (A. E. Verrill, 1901) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
34. Aplysia parvula Guilding in Mørch, 1863 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
Genus Bursatella Blainville, 1817. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
35. Bursatella leachii Blainville, 1817 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .218
Genus Dolabrifera Gray, 1847 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
36. Dolabrifera dolabrifera (Rang, 1828) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
Genus Petalifera Gray, 1847 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
37. Petalifera petalifera (Rang, 1828) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
38*. Petalifera ramosa Baba, 1959. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
Genus Phyllaplysia Fischer, 1872 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
39*. Phyllaplysia engeli Er. Marcus, 1955 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .219
Genus Syphonota H. Adams & A. Adams, 1854 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .219
40*. Syphonota geographica (Adams & Reeve, 1850). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
Genus Stylocheilus Gould, 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
41. Stylocheilus striatus (Quoy &Gaimard, 1832) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
42*. Stylocheilus longicauda (Quoy & Gaimard, 1825) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
Superorder Panpulmonata Jörger, Stöger, Kano, Fukuda, Knebelsberger & Schrödl, 2010 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
Order Sacoglossa Ihering, 1876 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
Family Oxynoidae Stoliczka, 1868 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .219
Genus Lobiger Krohn, 1847 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
43. Lobiger souverbii Fischer, 1857 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .220
Genus Oxynoe Rafinesque, 1814 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
44. Oxynoe antillarum Mörch, 1863 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
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45*. Oxynoe aguayoi Jaume, 1945 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
Family Boselliidae Ev. Marcus, 1982 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
Genus Bosellia Trinchese, 1891 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
46*. Bosellia curasoae Er. Marcus & Ev. Marcus, 1970 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
Family Plakobranchidae Gray, 1840 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221
Genus Elysia Risso, 1818 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .221
47. Elysia crispata Mörch, 1863 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221
48. Elysia ornata (Swainson, 1840) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221
49. Elysia papillosa Verril, 1901 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221
50. Elysia flava Verril, 1901 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222
51. Elysia serca Verrill, 1901 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222
52*. Elysia cauze Er. Marcus, 1957 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .222
53*. Elysia tuca Ev. Marcus & Er. Marcus, 1967 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222
54*. Elysia nisbeti Thompson, 1977 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222
55*. Elysia zuleicae Ortea & Espinosa, 2002 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223
56*. Checholysia patina (Ev. Marcus, 1980) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223
57*. Checholysia annedupontae Ortea, Espinosa & Caballer, 2005 (in Ortea et al. 2005) . . . . . . . . . . . . . . . . . . . . . . . . . . .223
Genus Thuridilla Bergh, 1872 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223
58*. Thuridilla malaquita Ortea & Buske, 2014 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223
Family Caliphyllidae Tiberi, 1881 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223
Genus Cyerce Bergh, 1870 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223
59*. Cyerce habanensis Ortea & Templado, 1989 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .224
Family Limapontiidae Gray, 1847. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224
Genus Ercolania Trinchese, 1872 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224
60. Ercolania coerulea Trinchese, 1892 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224
61*. Ercolania fuscata (Gould, 1870) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224
Genus Stiliger Ehrenberg, 1828. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224
62. Stiliger cricetus Er. Marcus & Ev. Marcus, 1970 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224
Genus Placida Trinchese, 1876 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .224
63*. Placida cremoniana (Trinchese, 1892) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224
64. Placida kingstoni T.E. Thompson, 1977 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
65. Placida verticilata Ortea, 1982 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
Genus Costasiella Pruvot-Fol, 1951 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
66*. Costasiella nonatoi Ev. Marcus & Er. Marcus, 1960 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
Family Cylindrobullidae Thiele, 1931. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
Genus Cylindrobulla Fischer, 1857. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
67. Cylindrobulla beauii Fischer, 1857. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
Family Volvatellidae Pilsbry, 1895 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226
Genus Ascobulla Ev. Marcus, 1972 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226
68. Ascobulla ulla (Er. Marcus & Ev. Marcus, 1970). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226
Clade Nudipleura Wägele & Willan, 2000 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226
Order Pleurobranchomorpha Pelseneer, 1906 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226
Family Pleurobranchidae Gray, 1827 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226
Genus Berthellina Gardiner, 1936. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226
69. Berthellina quadridens (Mörch, 1863) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .226
Genus Berthella Blainville, 1824 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .226
70. Berthella stellata (Risso, 1826). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226
71. Berthella agassizii (MacFarland, 1909) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226
Genus Pleurobranchus Cuvier, 1804. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226
72. Pleurobranchus areolatus Mörch, 1863 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226
Genus Pleurobranchaea Leue, 1813 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
73. Pleurobranchaea inconspicua Bergh, 1897 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
Order Nudibranchia Cuvier, 1817 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
Family Aegiridae P. Fischer, 1883 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .227
Genus Aegires Lovén, 1844 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
74*. Aegires sublaevis Odhner, 1932 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
75*. Aegires cf. ortizi Templado, Luque & Ortea, 1987 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .227
Family Goniodorididae H. Adams & A. Adams, 1854 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
Genus Okenia Menke, 1830 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
76. Okenia evelinae Er. Marcus, 1957 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
77. Okenia zoobotryon (Smallwood, 1910) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
Genus Bermudella Odhner, 1941. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 228
78*. Bermudella polycerelloides Ortea & Bouchet, 1983. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 228
Family Polyceridae Alder & Hancock, 1845. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 228
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Genus Polycera Cuvier, 1817 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .228
79. Polycera chilluna Er. Marcus, 1961 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .228
80. Polycera odhneri Er. Marcus, 1955 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 228
Genus Polycerella Verril, 1880 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229
81. Polycerella emertoni Verrill, 1880 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229
Genus Ta m b j a Burn, 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .229
82. Tam b j a cf. stegosauriformis Pola, Cervera & Gosliner, 2005. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229
Family Hexabranchidae Bergh, 1891 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .229
Genus Caribranchus Ortea, Caballer & Moro, 2003 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229
83. Caribranchus morsomus (Ev. Marcus & Er. Marcus, 1962) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229
Family Dorididae Rafinesque, 1815 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229
Genus Aphelodoris Bergh, 1879 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229
84. Aphelodoris antillensis Bergh, 1879 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229
Genus Doris Linnaeus, 1758 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 230
85. Doris verrucosa Linnaeus, 1758. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 230
86. Doris bovena Er. Marcus, 1955. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 230
87. Doris kyolis (Ev. Marcus & Er. Marcus, 1967a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 230
Family Discodorididae Bergh, 1891 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 230
Genus Discodoris Bergh, 1877 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 230
88. Discodoris branneri MacFarland, 1909 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 230
Genus Jorunna Bergh, 1876 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .230
89*. Jorunna spazzola (Er. Marcus, 1955) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 230
Genus Geitodoris Bergh, 1891 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
90. Geitodoris immunda Bergh, 1894 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
Genus Platydoris Bergh, 1877 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
91*. Platydoris angustipes (Mörch, 1863) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
Genus Sclerodoris Eliot, 1904. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
92*. Sclerodoris prea (Ev. Marcus & Er. Marcus, 1967) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
Familiy Dendrodorididae O'Donoghue, 1924 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
Genus Dendrodoris Ehrenberg, 1831 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
93. Dendrodoris warta Ev. Marcus & Gallagher, 1976 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
94. Dendrodoris krebsii (Mörch, 1863). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
Family Cadlinidae Bergh, 1891 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232
Genus Cadlina Bergh, 1878 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232
95*. Cadlina rumia Er. Marcus, 1955 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232
Family Chromodorididae Bergh, 1891 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232
Genus Ty r i n na Bergh, 1898 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232
96. Tyrinna evelinae (Er. Marcus, 1958) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .232
Genus Felimida Ev. Marcus, 1971b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .232
97*. Felimida binza (Ev. Marcus & Er. Marcus, 1963) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .232
98*. Felimida clenchi (Russel, 1935) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232
99. Felimida grahami Thompson, 1980 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233
Genus Felimare (Ev. Marcus & Er. Marcus, 1963). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233
100. Felimare acriba (Ev. Marcus & Er. Marcus, 1967) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .233
101*. Felimare fregona (Ortea & Caballer in Ortea, Espinosa, Buske & Caballer, 2013) . . . . . . . . . . . . . . . . . . . . . . . . . . . 233
102*. Felimare kempfi (Ev. Marcus, 1971a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233
103. Felimare marci (Ev. Marcus, 1971c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .233
104. Felimare molloi (Ortea & Valdés in Ortea, Valdés, & García-Gómez, 1996) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 234
105*. Felimare picta (Schultz in Philippi, 1836) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 234
106. Felimare ruthae (Ev. Marcus & Hughes, 1974) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 234
107. Felimare samueli (Caballer & Ortea, 2012) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 234
Family Arminidae Iredale & O'Donoghue, 1923 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 234
Genus Armina Rafinesque, 1814 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .234
108. Armina muelleri (Ihering, 1886) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 234
Family Flabellinidae Bergh in Carus, 1889 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .234
Genus Flabellina Gray in Griffith & Pidgeon, 1833 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 234
109*. Flabellina dushia (Ev. Marcus & Er. Marcus, 1963) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235
110. Flabellina engeli engeli Ev. Marcus & Er. Marcus, 1968 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235
111. Flabellina verta (Ev. Marcus, 1970) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235
112. Flabellina marcusorum Gosliner & Kuzirian, 1990 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235
Family Facelinidae Bergh in Carus, 1889 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235
Genus Austraeolis Burn, 1962. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235
113* . Austraeolis catina Ev. Marcus & Er. Marcus, 1967 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .235
Genus Dondice Er. Marcus, 1958 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .235
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114. Dondice occidentalis Engel, 1925 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .235
115. Dondice parguerensis Brandon & Cutress, 1985 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236
Genus Godiva Macnae, 1954. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236
116* . Godiva rubrolineata Edmunds, 1964 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236
Genus Favorinus Gray, 1850 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236
117. Favorinus auritulus Er. Marcus, 1955. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236
Genus Nanuca Er. Marcus, 1957. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236
118* . Nanuca sebastiani Er. Marcus, 1957 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236
Genus Phidiana Gray, 1850. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 237
119. Phidiana lynceus Bergh, 1867 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 237
Genus Learchis Bergh, 1896 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 237
120*. Learchis poica Ev. Marcus & Er. Marcus, 1960 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 237
121. Learchis? ignis Crescini, De Sisto & Villalba, 2013a. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 237
Family Aeolidiidae Gray, 1827 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 237
Genus Spurilla Bergh, 1864 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 237
122. Spurilla braziliana MacFarland, 1909 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .237
123*. Spurilla sargassicola Bergh, 1871 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 238
Genus Berghia Trinchese, 1877 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .238
124. Berghia creutzbergi Er. Marcus & Ev. Marcus, 1970 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 238
125. Berghia rissodominguezi Muniain & Ortea, 1999 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 238
Genus Anteaeolidiella Miller, 2001 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 238
126. Anteaeolidiella lurana (Ev. Marcus & Er. Marcus, 1967d) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 238
Family Eubranchidae Odhner, 1934 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 238
Genus Eubranchus Forbes, 1838 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 238
127*. Eubranchus conicla (Er. Marcus, 1958) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 238
Family Tergipedidae Bergh, 1889 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 238
Genus Cuthona Alder & Hancock, 1855 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 238
128*. Cuthona iris Edmunds & Just, 1983 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239
Family Tritoniidae Lamarck, 1809 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239
Genus Marionia Vayssière, 1877 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239
129. Marionia tedi Ev. Marcus, 1983 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239
Genus Tritoniopsis Eliot, 1905 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239
130*. Tritoniopsis frydis Er. Marcus & Ev. Marcus, 1970 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239
Family Hancockiidae MacFarland, 1923 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239
Genus Hancockia Gosse, 1877 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239
131. Hancockia ryrca Er. Marcus, 1957 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239
Family Bornellidae Bergh, 1874 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239
Genus Bornella Gray, 1850 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239
132*. Bornella calcarata Mörch, 1863 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .239
Family Scyllaeidae Alder & Hancock, 1855 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245
Genus Scyllaea Linnaeus, 1758 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .245
133*. Scyllaea pelagica Linnaeus, 1758. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245
Family Dotidae Gray, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245
Genus Doto Oken, 1815 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245
134*. Doto casandra Ortea, 2013 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245
Final remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 246
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247
Abstract
The Caribbean waters of Venezuela are composed by a large variety of habitats, with over 2800 km of coastline, islands,
and islets. This area is a transitional zone between two main biogeographic provinces, the Caribbean and the Brazilian,
separated by the fresh water outflows of the Orinoco and Amazon rivers, and is therefore expected to be an area of high
species diversity. However, concerning the study of molluscs, Venezuela is probably the poorest known region in the Ca-
ribbean. The best compilation of opisthobranch species known in Venezuela was produced almost a decade ago, mention-
ing the occurrence of 57 species, plus seven determined only to genus level. In this work, 134 species are reported for
Venezuela (71 are illustrated), representing about 40 % of the entire diversity of opisthobranchs known in the Caribbean.
Among the species occurring in Venezuela, 49 have here the southern limit of their distribution range and only one the
northern limit. Forty-six species are recorded for the first time to the country and one is a new record for the Caribbean
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Sea, namely Placida cremoniana. In addition, the distribution and ecology of the species are given based in literature and
new data.
Key words: taxonomy, new records, southern Caribbean, benthic fauna, marine biodiversity
Introduction
Opisthobranch sea slugs and snails (sensu Rudman & Willan 1998) have been found to be non-monophyletic and
novel classifications have been established (e.g., Schrödl et al. 2011; Zapata et al. 2014), but the descriptive term
“opisthobranch” is still widely used for some “clades with similar traits” (e.g., Galvão Filho et al. 2015).
Opisthobranchs can be found in most coastal areas of the world, from the poles to the tropics on a broad range of
habitats, from supratidal pools to the deep-sea, inhabiting coral reefs, rocky shelves, seagrass beds, coral rubble,
sandy bottoms and algae. They are regarded as being most diverse in shallow areas between 0–30 m deep,
particularly in tropical regions (Valdés et al. 2006; Gosliner et al. 2008), but deep-sea research is unravelling high
levels of previously unknown diversity of these molluscs at high depths too (Valdés 2008; Oskars et al. 2015).
Highest species richness is found in the Indo-Pacific biogeographic region, which harbors approximately 1400
species (Gosliner et al. 2008). In the Atlantic, the Caribbean Sea is the biogeographic province with highest
number of valid species of opisthobranchs. Presently, about 329 species are known in the area (Valdés et al. 2006;
García & Bertsch 2009).
Despite the milestone book by Valdés et al. (2006) on the opisthobranchs of the Caribbean Sea, several areas
remain poorly studied, including the waters of Venezuela. These are composed by a large variety of habitats (e.g.
coral reefs, mangroves, seagrass meadows, rocky shores, cays, sandy beaches, and coastal lagoons), stretching over
2800 km of coastline, islands, and islets (Petuch 1987). Venezuela is a transitional zone (see discussion) between
two main biogeographic provinces, the Caribbean and the Brazilian (Spalding et al. 2007; Briggs & Bowen 2012),
separated by the fresh water outflows of the Orinoco and Amazon rivers, and is therefore expected to be an area of
high species diversity (Petuch 2013). However, concerning the study of molluscs Venezuela is probably the poorest
known region in the Caribbean, as highlighted by Petuch (1987; 2013) more than 20 years ago.
A few opisthobranch species have been cited in regional faunal catalogs, catalogs of other regions or on
ecological studies (Marcus 1957; Rodríguez 1959; Marcus & Marcus 1960; 1962; 1963; 1964; 1967a; 1970; Work
1969; Marcus 1972; 1977; Abbot 1974; Marcus & Hughes 1974; Thompson 1977; González & Princz 1979; Rios
1985; Mollo et al. 1994; Gavagnin et al. 1996; 1997; 2000; Macsotay & Campos 2001; Diaz & Liñero-Arana 2004;
Fernandez & Jimenez 2007; Bitter et al. 2009; Massemin et al. 2009; Ortea et al. 2009; Rios 2009). The
opisthobranchs are an important source of bioactive chemicals with potential medical applications, and some
species inhabiting Venezuela have been cited in chemical studies: Gavagnin et al. (1996; 1997; 2000) found four
different secondary metabolites in specimens of Lobiger souverbii Fischer, 1857 from Los Roques and Elysia
crispata Mörch, 1863 from Mochima, three of them are polypropionates used as defence to high exposure to
sunlight. Miloslavich et al. (2010), on a bibliographic compilation of the invertebrate species living in the
Caribbean, reported 30 species of opisthobranchs for Venezuela. In fact, very few studies with focus on the
opisthobranchs of Venezuela have ever been published: Marcus & Marcus (1967a) described Hypselodoris acriba
Ev. Marcus & Er. Marcus, 1967a, and Marcus (1971c) described Hypselodoris marci Ev. Marcus, 1971c, both from
Los Roques archipelago; Rivero et al. (1998) recorded Aplysia parvula Guilding in Mørch, 1863 from Venezuela;
Mollo et al. (1994) recorded Stiliger cricetus Er. Marcus & Ev. Marcus, 1970 from the National Park Mochima;
Ortea & Valdés (1996 in Ortea et al. 1996) described Mexichromis molloi Ortea & Valdés, 1996 from Picuda
Island, Mochima; Mariño et al. (2011) recorded Dondice parguerensis Brandon & Cutress, 1985 from Buche;
Caballer & Ortea (2012) described Hypselodoris samueli Caballer & Ortea, 2012 from Morrocoy; Crescini et al.
(2013a) described Learchis ignis Crescini, De Sisto & Villalba, 2013a, from Margarita Island, where Crescini et al.
(2013b) recorded Doto chica Ev. Marcus & Er. Marcus, 1960 and Villaba & Crescini (2013) cited Favorinus
auritulus Er. Marcus, 1955. Lately, Caballer & Ortea (2015) described Philine buchensis Caballer & Ortea, 2015
from Buche and Grune et al. (2015) made an inventory of the opisthobranchs from La Restinga, Margarita.
Overall, Valdés et al. (2006) performed the best synthesis of opisthobranch species known to date in
Venezuela. The authors mentioned the occurrence of 64 species (seven of them determined only to genus level), but
most of the records have unspecified localities.
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This work aims to produce the first comprehensive catalogue of the opisthobranch gastropods known in
Venezuela based on reliable literature records and fieldwork undertaken by the authors along the last 20 years. Both
scientific and technical literature are reviewed, and an annotated checklist is produced discussing whenever
relevant taxonomic problems and including new ecological and geographical data. Illustrations of problematic
species or species depicting unique features are shown.
Material and methods
Sampling took place between the years 1993 to 2015 as a part of a broader research project aiming to survey the
whole diversity of marine molluscs in Venezuela and to unravel new bioactive molecules on these organisms.
Several areas have been sampled, both continental and insular, in a total of 52 sampling sites (Figure 1; Table 1).
FIGURE 1. Sampling points in Venezuela (see abbreviations in Table 1).
Sampling techniques comprised snorkeling and SCUBA diving down to 10 meters deep. Specimens were
obtained by (1) manual collecting after direct observation and (2) collecting suitable habitats (algae, mud, sand,
hydrozoans, etc) followed by in situ sieving through a 1 mm mesh sieve.
Specimens were measured, photographed, and fixed both for DNA extraction (96 % ethanol) and anatomical
studies (frozen over night in sea-water followed by defrosting and preservation in 70 % ethanol). Some species
were observed and photographed on their natural habitat during field campaigns, but not captured.
Species referred in the literature as having wide distribution ranges but lacking specific references to
Venezuela were not considered for the purposes of our inventory (e.g. Corambe obscura Verrill, 1870 cited in
Va ld é s et al. (2006) as “Widely distributed in the Atlantic Ocean. Western Atlantic: from Massachusetts to Brazil”).
Species that have been cited in Venezuela but whose taxonomic identification is uncertain (e.g. Valdés et al. 2006:
Retusa sp. 1, Flabellina sp. 1, sp. 2 and sp. 3 or Discodorididae sp. 1, Facelinidae sp. 3 and sp. 2) were also not
included in the present inventory.
To establish the distribution of the species in the Caribbean we followed, Ortea (2001), Espinosa et al. (2006;
2007), Valdés et al. (2006), Massemin et al. (2009), Rios (2009), Rosenberg (2009), Miloslavich et al. (2010), Yidi
& Sarmiento (2011), Redfern (2001; 2013), Ortea et al. (2012; 2013), Camacho-García et al. (2014b), Galvão Filho
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et al. (2015), Goodheart et al. (2015) and Grune et al. (2015). General records in Venezuela or records to specific
localities in the country are indicated between brackets. Records from outside the Caribbean are indicated with
their corresponding references.
TABLE 1. Sampling sites with acronyms and geographical coordinates.
Sampling sites Acronym Coordinates
Boca de la Laguna, La Restinga National Park, Nueva Esparta BLR 10°58’40’’N 64°10’02’’W
El Horcon, Macanao, Margarita, Nueva Esparta HMM 10°56’41.24’’N 64°17’31.51’’W
Embarcadero, Estacion Biologica UDO, Cubagua, Nueva Esparta ECU 10°49’33.08’’N 64°09’47.66’’W
Playa El Yaque's Lagoon, Isla Tortuga YL 10°57’43’’N 65°13’26’’W
East side of Playa El Yaque's Lagoon, Isla Tortuga EYL 10°57’48’’N 65°13’30’’W
West side of Playa El Yaque's Lagoon, Isla Tortuga WYL 10°57’22’’N 65°13’11’’W
La mano de Dios, Isla Tortuga LMD 10°55’04’’N 65°11’58’’W
Los Mogotes's Lagoon, Isla Tortuga ML 10°54’28’’N 65°12’89’’W
Pedrero de Punta Delgada, Isleta central, Isla Tortuga PPD 10°57’44’’N 65°13’20’’W
Rasa frente al manglar La Gaviota, Isla La Tortuga MG 10°54’14’’N 65°20’49’’W
Punta los negros, Isla La Tortuga PNT 10°54’21.21’’N 65°21’08’’W
Tortuguillo del este, Isla La Tortuga TT 10°58’09’’N 65°24’54’’W
Dique, Cayo Herradura, Isla Tortuga DCH 10°59’42’’N 65°23’04’’W
Norte de los Roques, Archipielago de los Roques AR 11°57’24.79’’N 66°41’41.21’’W
Dos Mosquises este, Archipielago de los Roques MR 11°47’35’’N 66°53’24’’W
Dos Mosquises sur, Archipielago de los Roques DMR 11°47’36’’N 66°53’46’’W
Cumana, Sucre CU 10°28’26’’N 64°08’54’’W
Mochima, Sucre MC 10°20’49’’N 64°20’23’’W
Fundaciencia Scientific Station, Mochima, Sucre FCM 10°20’48’’N 64°20’41’’W
Parque Nacional Mochima, Mochima, Sucre PNM 10°18’28.96’’N 64°28’45.98’’W
Playa Gabarra, Mochima, Sucre PG 10°22’51’’N 64°20’36’’W
Islas Caracas, Mochima, Sucre ICM 10°22’23’’N 64°24’52’’W
Isla Picuda, Chimanas, Mochima, Sucre PCM 10°18’26’’N 64°33’53’’W
Lado Este de El Ocho's Lagoon, Buche, Higuerote EOL 10°32’33’’N 66°05’48’’W
Lado sur de El Ocho's Lagoon, Buche, Higuerote SEL 10°32'28,7"N 66°05'55,3"W
El Ocho Lagoon, Buche, Higuerote OLH 10°32.27'94"N 66°05'55"W
Tunel del amor, Buche, Higuerote TA 10°32’21’’N 66°05’25’’W
Thalassia meadow facing Buche beach, Higuerote PTH 10°32’15’’N 66°05’17’’W
Puerto Cabello, Carabobo PC 10°29’06’’N 68°00’12’’W
Chichiriviche de la Costa, Vargas CHC 10°33’06.24’’N 67°14’19.36’’W
Cayo Playa Azul, Tucacas, Morrocoy National Park PAM 10°50’47’’N 68°13’44’’W
Cayo Norte Morrocoy, Morrocoy National Park CNM 10°46’49.34’’N 68°11’52.25’’W
Playa Varadero reef, Morrocoy APV 10°54’44’’N 68°14’48’’W
Boca del Golfete de Cuare, Morrocoy National Park GFM 10°54’34’’N 68°15’59’’W
Cueva de la Virgen, Golfete de Cuare, Morrocoy National Park CVM 10°54’21.48’’N 68°18’12.52’’W
El Ocho lagoon, Morrocoy National Park OLM 10°51’60’’N 68°13’42’’W
Lagoon near "Boca Grande", Morrocoy National Park BGM 10°51’01’’N 68°14’22’’W
Imparques, South side of Gulfete de Cuare, Morrocoy National Park GCM 10°51’13’’N 68°16’24’’W
Los Juanes, Morrocoy National Park LJM 10°51’54’’N 68°12’59’’W
......continued on the next page
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New records of opisthobranchs for Venezuela are marked with an asterisk (*). Sampling localities are in
abbreviated format; full names as in Table 1. For higher taxonomy arrangements we followed, Wägele & Willan
(2000), Jörger et al. (2010), Schrödl et al. (2011), Wägele et al. (2014), Zapata et al. (2014) and Schrödl (2014).
The pteropod orders Thecosomata and Gymnosomata, because of their common and distinct ecological nature
(they are pelagic), were not considered in this inventory.
The following abbreviations are here used: H—maximum animal length; spc./spcs—live specimen / live
specimens; sh—shell(s); ZMBN—Department of Natural History, University Museum of Bergen, University of
Bergen, Norway; CMIVIC—Marine Collections of the Instituto Venezolano de Investigaciones Cientifícas,
Venezuela; PNC—Photographed by Gaby Carias, but not captured; GCC—Gabriela Carias Collections, Policlínica
Metropolitana Urbanización Caurimare, Calle A-1, Caracas, Venezuela; MCFUDENA—Molluscan Collections of
the Fundacion de Defensa de la Naturaleza, Venezuela, Calle Zamora, Chichiriviche, Venezuela; JOC—Jesus Ortea
Collections, Department BOS, Faculty of Biology, University of Oviedo, Asturias, Spain.; MCC—Manuel
Caballer Collections, Barrio El Tolio 5, 39478 Boo de Pielagos, Cantabria, Spain; NRC—Nelsy Rivero
Collections, Environmental Sciences Headship, Academic Vice-Chancellorship, Caribbean Maritime Experimental
National University, Ave. El Ejercito, Catia La Mar, Vargas, Venezuela.
Results
Class Gastropoda Cuvier, 1795
SubClass Heterobranchia Burmeister, 1837
Clade Tectipleura Schrödl, Jörger, Klussmann-Kolb & Wilson, 2011
Superorder Euopisthobranchia Jörger, Stöger, Kano, Fukuda, Knebelsberger & Schrödl, 2010
Order Cephalaspidea P. Fischer, 1883
Family Aglajidae Pilsbry, 1895
Genus Migaya Ortea, Caballer & Espinosa, 2014
TABLE 1. (Continued)
Sampling sites Acronym Coordinates
Boca Seca, Morrocoy National Park BSM 10°54’21’’N 68°14’07’’W
Cayo Borracho, Morrocoy National Park CBM 10°51’05’’N 68°12’33’’W
Cayo Sombrero, central shoal, Morrocoy National Park CSM 10°56’02’’N 68°12’13’’W
Cayo Muerto, Morrocoy National Park CMM 10°53’07’’N 68°15’35’’W
Cayo Sal, Morrocoy National Park SLM 10°55’50’’N 68°15’31’’W
Cayo Sur, Morrocoy National Park SRM 10°53’01’’N 68°12’14’’W
Playa Norte, Morrocoy National Park NPM 10°53’27’’N 68°16’56’’W
Playa Sur, Morrocoy National Park SPM 10°57’14’’N 68°13’18’’W
Cayo Capuchinos, Morrocoy National Park CM 10°51’25’’N 68°18’13’’W
Las Luisas, Morrocoy National Park LUM 10°49’40’’N 68°17’59’’W
Reef north side of "Cayo Pescadores", Morrocoy National Park CPM 10°52’08’’N 68°12’46’’W
Amuay, Paraguaná, Falcón AM 11°46'24.53"N 70°14'39.96"W
Isla de Aves, Dependencias Federales IA 15°39'53.26"N 63°38'13.45"W
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1*. Migaya felis (Er. Marcus & Ev. Marcus, 1970)
(Figure 2 A)
Material examined. WYL, 5 spcs, H = 7–10 mm, IVICCM000094. WYL, 8 spcs, H = 6–10 mm, IVICCM000102.
WYL, 15 spcs, H = 13 mm, ZMBN 84900.
Ecology. On shallow soft sandy bottoms with high sun exposure, around 1 m deep. The spawn is a coiled
yellow cord filled with small eggs tightly paked together.
Distribution. Bahamas, Cayman Islands, Cuba, Guadeloupe, Jamaica, Mexico, Puerto Rico, Venezuela.
Remarks. Redfern (2001: Fig 661A Plate 115 and Fig. 661B Plate 70) illustrated this species (as Runcina sp.)
based on specimens from the Bahamas. This species was only found in a very restricted area (ca. 10 x 10 m) at El
Yaque's Lagoon, Isla Tortuga, but in high density. This particular area was characterized by soft bottoms with fine
sandy-mud. The animals were usually in clusters of two to five specimens. Ocasionally, lines with 4–8 individuals
were observed. They release a reddish secretion when disturbed.
Aglaja felis Er. Marcus & Ev. Marcus, 1970 has been transferred to the Japanese genus Nakamigawaia Kuroda
& Habe, 1961 (in Habe 1961) by Camacho-García et al. (2014), without a sound explanation and comparison with
the type species of the genus (Nakamigawaia spiralis Kuroda & Habe, 1961, in Habe 1961), which has a different
shell, among other characters. The genus Migaya was erected to host the Caribbean populations of M. felis due to
the unique characters of the species (Ortea et al. 2014). Venezuela represents the southern limit of the distribution
of this species in the Caribbean.
Genus Navanax Pilsbry, 1895a
2. Navanax gemmatus (Mörch, 1863)
(Figure 2 B)
Material examined. WYL, 1 spc., H = 27 mm, IVICCM000114. YL, 2 spcs, H = 20, 50 mm, ZMBN 84912. EYL,
6 spcs, H = 10–30 mm, ZMBN 84908. IA, 1 spc., PNC.
Ecology. 0.5–1 m deep, commonly found under rocks or crawling on rocks, seldom on sand.
Distribution. ABC Islands, Antigua and Barbuda, Bahamas, Barbados, Belize, Brazil, Chile, Colombia, Costa
Rica, Cuba, Curaçao, Grenada, Guadeloupe, Honduras, Jamaica, Martinique, Mexico, St. Barthelemy/St.
Bartholomew, St. Vincent & the Grenadines, USA, Virgin Islands, Venezuela (Valdés et al. 2006, Rosenberg 2009.
Mochima: Gavagnin et al. 2000).
Remarks. Recently, Ornelas-Gatdula et al. (2012), based on molecular evidence, considered all records of
Navanax aenigmaticus (Bergh, 1893) in the Caribbean to be N. gemmatus, which is according to the authors the
only species of the genus occurring in the western Atlantic Ocean. This species is known to feed upon other
“opisthobranchs” (Valdés et al. 2006) and we have observed it feeding on the sacoglossan Elysia crispata during
fieldwork while they were temporarily left together on a tray. This species releases a reddish-yellow substance
when disturbed.
Genus Chelidonura A. Adams, 1850
3*. Chelidonura hirundinina (Quoy & Gaimard, 1833)
(Figure 2 C)
Material examined. WYL, 3 spcs, H = 17–29 mm, IVICCM000103. PNT, 1 spc., H = 15 mm, IVICCM000109.
AR, 2 spcs, H = 15–16 mm, GCC. AR, 1 spc., PNC.
Ecology. Intertidal to shallow on rocky or sandy shores; 1 m deep.
Distribution. ABC Islands, Bahamas, Cayman Islands, Cuba, Guadeloupe, Jamaica, Mexico, Puerto Rico,
USA, Venezuela.
Remarks. This colourful species feeds on flatworms associated to filamentous green algae that are abundant
on sandy-mud bottoms around the beds of Thalassia testudinum. According to a recently published work
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(Camacho-García et al. 2014), this name refers to a complex of species from all around the world, with Indo-
Pacific specimens showing a different affiliation from the Bahamian specimen included by the authors. However,
the Caribbean specimens need further studies to determine if they are a complex or unique species. This is a new
southern limit to the distribution of this species.
4*. Chelidonura cubana Ortea & Martínez, 1997
(Figure 5 I)
Material examined. ALM, 1 spc., PNC.
Ecology. Common on shallow waters, among algae or death corals.
Distribution. Bahamas, Cayman Islands, Cuba, Guadeloupe, Venezuela.
Remarks. This is a new southern limit to the distribution of this species.
Family Cylichnidae H. Adams & A. Adams, 1854
Genus Cylichnella Gabb, 1873
5. Cylichnella bidentata (d´Orbigny, 1841)
Distribution. Brazil, Colombia, Cuba, Guadeloupe, Jamaica, Martinique, Mexico, Panama, Puerto Rico, Uruguay,
USA, Virgin Islands, Venezuela (Valdés et al. 2006; Rosenberg 2009; Massemin et al. 2009: as Acteocina
bidentata; Miloslavich et al. 2010: as Acteocina bidentata; Yidi & Sarmiento 2011).
Remarks. No material of this species was found.
Genus Acteocina J. E. Gray, 1847
6. Acteocina candei (d´Orbigny, 1841)
Material examined. EOL, 1 spc., H = 1 mm, IVICCM000106s.
Ecology. Uncommon on shallow sandy bottoms.
Distribution. Argentina, Bermuda, Brazil, Colombia, Costa Rica, Cuba, Guadeloupe, Guyana, Jamaica,
Martinique, Mexico, Panama, Puerto Rico, St. Croix, USA, Virgin Islands, Venezuela (Valdés et al. 2006;
Rosenberg 2009; Miloslavich et al. 2010. Zulia & Falcon: Princz 1983. Morrocoy: Bitter et al. 2009).
Genus Cylichna Lovén, 1846
7. Cylichna krebsii Mörch, 1875
Distribution. Anguilla, Colombia, Panama, Puerto Rico, St. Barthelemy/St. Bartholomew, St. Martin/St. Maarten,
Venezuela (Miloslavich et al. 2010).
Remarks. No material of this species was found. Venezuela is the southern limit of the distribution of this
species.
Family Retusidae Thiele, 1925
Genus Retusa Brown, 1827
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8. Retusa sulcata (d´Orbigny, 1841)
Distribution. Bahamas, Colombia, Costa Rica, Cuba, Guadeloupe, Panama, USA, Virgin Islands, Venezuela
(Princz 1982; Valdés et al. 2006; Rosenberg 2009; Yidi & Sarmiento 2011).
Remarks. No material of this species was found. Venezuela is the southern limit of the distribution of this
species.
Genus Pyrunculus Pilsbry, 1895
9. Pyrunculus caelatus (Bush, 1885)
Distribution. Argentina, Brazil, Guadeloupe, Guyana, Mexico, Puerto Rico, Surinam, Uruguay, USA, Venezuela
(Valdés et al. 2006: as Retusa caelata; Massemin et al. 2009; Rios 2009; Rosenberg 2009: as Retusa caelata;
Miloslavich et al. 2010. La Blanquilla island & Margarita island: Princz 1977: as Retusa caelata).
Remarks. No material of this species was found.
Family Rhizoridae Dell, 1952
Genus Vol v ul e l l a Newton, 1891
10. Volvulella minuta (Bush, 1885)
Distribution. Cuba, Guadeloupe, Jamaica, Martinique, Mexico, Panama, Puerto Rico, USA, Virgin Islands,
Venezuela (Princz 1982: as Volvulella recta; Valdés et al. 2006: Volvulella acuta; Rosenberg 2009).
Remarks. No material of this species was found.Venezuela is the southern limit of the distribution of this
species.
11. Volvulella persimilis (Mörch, 1875)
Distribution. Argentina, Bahamas, Bermuda, Brazil, Colombia, Costa Rica, Cuba, Guadeloupe, Guyana, Mexico,
Panama, Puerto Rico, Santo Domingo, Surinam, Uruguay, USA, Venezuela (Marcus & Marcus 1960; Rios 1985;
Massemin et al. 2009; Rios 2009; Rosenberg 2009; Miloslavich et al. 2010; Yidi & Sarmiento 2011. Falcon, Sucre
& Margarita Island: Princz 1982).
Remarks. No material of this species was found.
12. Volvulella texasiana Harry, 1967
Distribution. Colombia, USA, Venezuela (Miloslavich et al. 2010; Yidi & Sarmiento 2011: as Paravolvulella
texasiana).
Remarks. No material of this species was found. Venezuela is the southern limit of the distribution of this
species.
Family Gastropteridae Swainson, 1840
Genus Gastropteron Kosse, 1813
13. Gastropteron chacmol Gosliner, 1989
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Distribution. Bahamas, Brazil, Cayman Islands, Cuba, Mexico, Venezuela (Valdés et al. 2006; Rosenberg 2009;
Miloslavich et al. 2010. Caracas Island, Sucre: Gosliner 1989. Vargas: Rosenberg 2009).
Remarks. No material of this species was found.
Family Philinidae Gray, 1850
Genus Philine Ascanius, 1772
14. Philine sagra (d'Orbigny, 1841)
Distribution. Brazil, Colombia, Martinique, Panama, Puerto Rico, USA, Venezuela (Massemin et al. 2009;
Miloslavich et al. 2010; Yidi & Sarmiento 2011. Sucre & Margarita Island: Rosenberg 2009).
Remarks. No material of this species was found.
15. Philine buchensis Caballer & Ortea, 2015
(Figure 2 D)
Material examined. OLH, 2 spcs, H = 3–4 mm, IVICCM000133. OLH, 2 spcs, H = 2–3 mm, ZMBN 84905.
Ecology. Shallow mangrove lagoon (1–2 m deep) with soft bottom and brown and green algae including the
calcareous Halimeda spp.
Distribucion. Venezuela.
Remarks. So far only known from Venezuela (type locality: Buche).
Family Scaphandridae Sars, 1878
Genus Scaphander Montfort, 1810
16. Scaphander watsoni Dall, 1881
Distribution. Brazil, Colombia, Cuba, Guadeloupe, Mexico, Panama, USA, Venezuela (Abbott 1974; Marcus
1977; Massemin et al. 2009; Rios 2009; Rosenberg 2009; Eilertsen & Malaquias 2013. Los Testigos: Marcus &
Marcus 1967b; Marcus 1974; Macsotay & Campos 2001).
Remarks. No material of this species was found.
17. Scaphander nobilis Verril, 1884
Distribution. This is an amphi-Atlantic species occurring in the eastern Atlantic from the Bay of Biscay to Senegal
including the Azores and in the western Atlatic is known from Brazil, Cuba, Haiti, Trinidad & Tobago, Gulf of
Mexico, USA, Canada, and Venezuela (Massemin et al. 2009; Eilertsen & Malaquias 2013).
Remarks. No material of this species was found.
Family Bullidae Gray, 1827
Genus Bulla Linnaeus, 1758
18. Bulla occidentalis A. Adams, 1827
(Figure 2 E–F)
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Material examined. APV, 2 spcs, H = 10–35 mm, NRC. CM, 10 spcs, H = 22–27 mm, NRC. LUM, 4 spcs, H =
28–32 mm, NRC. EOL, 2 spcs, H = 20–30 mm, IVICCM000015. SEL, 11 spcs, H = 15–25 mm, IVICCM000058.
OLH, 6 spcs, H = 20–30 mm, IVICCM000137. OLH, 2 spcs, H = 18–27 mm, IVICCM000144. SEL, 3 spcs, H = 2
mm, IVICCM000169. OLH, 8 spcs, H = 13–17 mm, IVICCM000209. ML, 1 spc., H = 18 mm, IVICCM000001s.
YL, 17 spcs, H = 20–30 mm, IVICCM000051s. GCM, 3 spcs, H = 13–16 mm, IVICCM000072s. YL, 2 spcs, H =
23–33 mm, ZMBN 84919. ML, >50 spcs, H = 15 mm, ZMBN 84902. OLH, >50 spcs, H = 10–15 mm, ZMBN
84901. OLM, 2 spcs, H = 4, 9 mm, ZMBN 84927. BGM, 10 spcs, H = 7–15 mm, ZMBN 84936. IA, 1 spc., PNC.
Ecology. Inside shallow mangrove lagoons (1–2 m deep) with muddy bottom and abundant brown and green
algae. Empty shells and egg-masses were observed in reef lagoons with sand bottoms and meadows of the seagrass
Thalassia spp.
Distribution. Antigua, Bahamas, Barbados, Barbuda, Belize, Bermuda, Brazil, Colombia, Cuba, Costa Rica,
Dominicana, Grenada, Guadeloupe, Honduras, Jamaica, Martinique, Mexico, Panama, St. Thomas, St. Croix, St.
Martin/St. Maarten, St. Lucia, St. Vincent & the Grenadines, Surinam, Trinidad & Tobago, Uruguay, USA, Virgin
Islands, Venezuela (Most authors called this species Bulla striata Brugière, 1792 but see Malaquias & Reid (2008)
for a review: Rios 1985; Valdés et al. 2006; Malaquias & Reid 2008; Rios 2009; Rosenberg 2009; Miloslavich et
al. 2010; Yidi & Sarmiento 2011. Gulf of Paria & Miranda: Marcus & Marcus 1967b. Los Roques (as Bulla
umbilicata): Work 1969, Margarita island, Coche island, Cubagua island & Chacopata Peninsule, Sucre: Macsotay
& Campos 2001, Mochima: Diaz & Liñero-Arana 2004. La Restinga, Margarita: Grune et al. 2015: as B. striata).
Remarks. This species shows remarkable phenotypic and ecotypic variability with specimens from sandy
bottoms and seagrass having larger and lighter coloured shells in comparison with those from mangrove
environments that show in average smaller and darker shells. Malaquias & Reid (2008), based on DNA and
morphology, have shown that despite these differences, only a single species of the “striata” complex is present in
the western Atlantic. Egg-masses were observed tangled on leaves of the seagrass Thalassia spp.
Family Haminoeidae Pilsbry, 1895b
Genus Haminoea Turton & Kingston, 1830
19. Haminoea cf. elegans (Gray, 1825)
(Figure 2 G)
Material examined. EOL, 2 spcs, H = 4–7 mm, IVICCM000018. OLH, 1 spc., H = 6 mm, IVICCM000151. OLH,
5 spcs, H = 5–8 mm, ZMBN 84903. OLM, 1 spc., H = 4 mm, ZMBN 84926.
Ecology. Inside mangrove lagoons (1–2 m deep) with soft bottom and brown and green algae including
Halimeda spp.
Distribution. ABC Islands, Bermuda, Brazil, Colombia, Costa Rica, Cuba, Grenada, Guadeloupe, Jamaica,
Mexico, Puerto Rico, St. Vincent & the Grenadines, USA, Virgin Islands, Venezuela (Princz 1982; Valdés et al.
2006; Rosenberg 2009; Miloslavich et al. 2010; Yidi & Sarmiento 2011. Morrocoy: Bitter et al. 2009. La Restinga,
Margarita: Grune et al. 2015).
Remarks. The systematics of the genus Haminoea is confusing due to the poor definition of the species
characters, most of the taxonomic work is shell-based and shells are very similar between species which makes
separation difficult. A review of the worldwide species of Haminoea is underway (Malaquias, in progress) and we
here tentatively identify these specimens as H. elegans due to the presence of spiral grooves in the shell.
20. Haminoea antillarum (d´Orbigny, 1841)
Material examined. SEL, 3 spcs, H = 9–12 mm, IVICCM000057. OLH, 3 spcs, H = 5 mm, IVICCM000171.
Ecology. Inside mangrove lagoons (1–2 m deep) with soft bottom and brown and green algae including
Halimeda spp.
Distribution. Bahamas, Bermuda, Brazil, Cayman Islands, Colombia, Costa Rica (?), Cuba, Guadaloupe,
Jamaica, Mexico, Panama, Puerto Rico, USA, Virgin Islands, Venezuela (Valdés et al. 2006; Rosenberg 2009;
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Miloslavich et al. 2010. Chacopata peninsula, Sucre: Macsotay & Campos 2001: as Haminoea antillarum
guadaloupensis. La Restinga, Margarita: Grune et al. 2015).
21. Haminoea succinea (Conrad, 1846)
Material examined. SPM, 1 spc., H = 10 mm, NRC.
Ecology. Uncommon. Shallow waters among algae and seaweed.
Distribution. Bermuda, Colombia, Guadeloupe, Mexico, Puerto Rico, St. Martin/St. Maarten, St. Barthelemy/
St. Bartholomew, USA, Venezuela (Valdés et al. 2006; Rosenberg 2009; Miloslavich et al. 2010; Yidi & Sarmiento
2011. Punta Arenas (Margarita Island) and Sucre: Macsotay & Campos 2001. Morrocoy: Bitter et al. 2009).
Remarks. Venezuela is the southern limit of the distribution of this species.
22. Haminoea petitii (d'Orbigny, 1841)
Material examined. OLH, 2 spcs, H = 7–8 mm, IVICCM000150.
Ecology. Inside shallow mangrove lagoons (1–2 m deep) with muddy bottom and abundant brown and green
algae (only empty shells).
Distribution. ABC Islands, Brazil, Cayman Islands, Colombia, Cuba, Guadeloupe, Mexico, Puerto Rico,
USA, Virgin Islands, Venezuela (Morrocoy: Bitter et al. 2009).
Genus Atys Montfort, 1810
23. Atys caribaeus (d´Orbigny, 1841)
Distribution. Brazil, Colombia, Costa Rica, Cuba, Guadeloupe, Jamaica, Mexico, Panama, Puerto Rico, United
States, Virgin Islands, Venezuela (Valdés et al. 2006; Miloslavich et al. 2010; Yidi & Sarmiento 2011).
Remarks. No material of this species was found. Valdés et al. (2006) considered Atys sharpi Vanatta, 1901 a
synonym of this species.
24. Atys guildingi Sowerby II, 1869
Distribution. Brazil, Colombia, Costa Rica, Cuba, Guadeloupe, Panama, Venezuela (Valdés et al. 2006; Yidi &
Sarmiento 2011).
Remarks. No material of this species was found. Venezuela is the southern limit of the distribution of this
species.
25. Atys riiseanus Mörch, 1875
Distribution. ABC Islands, Anguilla, Bahamas, Brazil, Colombia, Costa Rica, Cuba, Guadeloupe, Dominican
Republic, Guadeloupe, Mexico, Panama, Puerto Rico, St. Martin/St. Maarten, Trinidad & Tobago, USA, Virgin
Islands, Venezuela (Miloslavich et al. 2010).
Remarks. No material of this species was found.
26. Atys sharpi Vanatta, 1901
Distribution. Bahamas, Brazil, Colombia, Costa Rica, Cuba, Guadeloupe, Jamaica, Mexico, Panama, Puerto Rico,
USA, Virgin Islands, Venezuela (Yidi & Sarmiento 2011).
Remarks. No material of this species was found.
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Order Runcinacea Burn, 1963
Family Runcinidae H. Adams & A. Adams, 1854
Genus Lapinura Er. Marcus & Ev. Marcus, 1970
27*. Lapinura divae (Ev. Marcus & Er. Marcus, 1963)
(Figure 2 H)
Material examined. MR, 2 spcs, H = 2 mm, JOC.
Ecology. Found in shallow protected areas, at the base of the algae Caulerpa racemosa.
Distribution. Bahamas, Costa Rica, Guadeloupe, Jamaica, Venezuela.
Remarks. The genus Lapinura is characterized by an external shell protecting the gills, a unique character that
clearly separates it from other genera in the Family (Ortea et al. 2013). Venezuela represents the southern limit of
the distribution of this species.
Order Umbraculida Odhner, 1939
Family Umbraculidae Dall, 1889
Genus Umbraculum Schumacher, 1817
28. Umbraculum umbraculum (Lightfoot, 1786)
Distribution. Brazil, Colombia, Costa Rica, Cuba, Guadeloupe, Honduras, USA, Venezuela (Punta Moreno Bank,
Nueva Esparta: Macsotay & Campos, 2001). Circumtropical (Valdés et al. 2006).
Remarks. No material of this species was found. Venezuela is the southern limit of the distribution of this
species in the Caribbean.
Order Anaspidea Fischer, 1883
Family Aplysiidae Lamarck, 1809
Genus Aplysia Linnaeus, 1767
29. Aplysia dactylomela Rang, 1828
(Figure 2 I)
Material examined. WYL, 1 spc., H = 60 mm, IVICCM000111. YL, 3 spcs, H = 50–120 mm, ZMBN 84916,
ZMBN 84917. APV, 1 spc., H = 51 mm, IVICCM000078. CBM, 4 spcs, H = 20–101 mm, NRC. SPM, 3 spcs, H =
55–88 mm, NRC. CHC, 1 spc., PNC.
Ecology. Crawling on seagrass or rocky-muddy bottoms (1 m deep). This species release a purple fluid when
disturbed.
Distribution. ABC Islands, Anguilla, Bahamas, Barbados, Bermuda, Belize, Brazil, Cayman Islands,
Colombia, Costa Rica, Cuba, Guadeloupe, Jamaica, Martinique, Mexico, Panama, Puerto Rico, St. Kitts/St.
Christopher, St. Martin/St. Maarten, USA, Virgin Islands, Venezuela (Valdés et al. 2006; Rios 2009; Miloslavich et
al. 2010; Yidi & Sarmiento 2011. Punta Charagato, Cubagua island & La Galera, Margarita island: Rodriguez
1959. Isla Larga, South Island: Marcus & Marcus 1967a. Los Roques: Work 1969; Rosenberg 2009. Taguao, Las
Salinas (Vargas), Chirimena (Miranda), Morrocoy National Park (Falcon), Mochima Bay & Golfo de Cariaco
(Sucre), Margarita island (Nueva Esparta) & Los Roques: Rivero et al. 2003). Confined to the Atlantic Ocean,
where it is common (Valdés et al. 2013).
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Remarks. This is an Atlantic subtropical species (Valdés et al. 2013) characterized by a whitish-creamy colour
with distinct black rings.
30. Aplysia brasiliana Rang, 1828
(Figure 2 J)
Material examined. YL, 1 spc., H = 25 mm, ZMBN 84911, 1 spc., H = 45 mm, ZMBN 84910. 3 spcs, H = 20–60
mm, ZMBN 84914. CMM, >50 spcs, H = 70–106 mm, NRC. ECU, >50 spcs, H = 25–186 mm, MCC.
Ecology. On seagrass or rocky-sandy bottoms (1 m deep). This species release a purple fluid when disturbed.
Distribution. ABC Islands, Bermuda, Brazil, Colombia, Costa Rica, Guadeloupe, Mexico, USA, Venezuela
(Valdés et al. 2006; Rios 2009; Miloslavich et al. 2010; Rosenberg 2009: as A. fasciata; Yidi & Sarmiento 2011: as
A. fasciata. Puerto Viejo, Camuri Grande, Taguao, Las Salinas (Vargas), Mochima Bay, Isla Caribe (Sucre) &
Cubagua island (Nueva Esparta): Rivero et al. 2003. La Restinga, Margarita: Grune et al. 2015).
Remarks. Rosenberg (2009) and Yidi & Sarmiento (2011) considered A. brasiliana a junior synonym of
Aplysia fasciata Poiret, 1789. This is not surprising since there is much confusion surrounding the systematics of
the genus Aplysia with names being often used arbitrarily. Only an integrated taxonomic approach combining both
morphological, anatomical, and DNA characters across the entire geographical range of species could end the
uncertainties surrounding the systematics of Aplysia. Meanwhile we consider that there are characters enough to
distinguish both species, of which, only A. brasiliana can be found in the Caribbean.
31. Aplysia cervina (Dall & Simpson, 1901)
Ecology. On seaweed (T. testudinum or Zoostera marina), under rocks on sandy bottoms or among mangrove.
Distribution. ABC Islands, Brazil, Colombia, Mexico, Puerto Rico, USA, Venezuela (Miloslavich et al. 2010.
Taguao, Las Salinas (Vargas) & Bahia Cienaga (Ocumare de la Costa, Aragua): Rivero et al. 2003).
Remarks. No material of this species was found.
32. Aplysia juliana Quoy & Gaimard, 1832
(Figure 2 K)
Material examined. YL, 1 spc., H = 30 mm, MCC.
Ecology. Shallow waters. On the coast, on rocky shores with Ulva spp.
Distribution. ABC Islands, Barbados, Brazil, Cuba, St. Lucia, USA, Venezuela (Miloslavich et al. 2010. Los
Roques: Marcus 1972. Taguao, Las Salinas (Vargas), Playa Corrales (Miranda), El Peñon (Sucre) & Los Roques:
Rivero et al. 2003). Circumtropical (Valdés et al. 2006).
33. Aplysia morio (A. E. Verrill, 1901)
Ecology. Found in tide pools on the coast. Active swimmer.
Distribution. Bahamas, Bermuda, USA, Venezuela (Miloslavich et al. 2010. La Restinga, Margarita: Grune et
al. 2015).
Remarks. No material of this species was found. Venezuela is the southern limit of the distribution of this
species.
34. Aplysia parvula Guilding in Mørch, 1863
(Figure 5 J)
Material examined. FCM, 4 spcs, H = 20–30 mm, JOC. IA, 1 spc., PNC.
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Ecology. Intertidal. Rocky shores on algae or sandy bottoms on seaweed.
Distribution. ABC Islands, Anguilla, Bahamas, Bermuda, Brazil, Cayman Islands, Costa Rica, Cuba,
Guadeloupe, Jamaica, Mexico, Puerto Rico, St. Vincent & the Grenadines, USA, Virgin Islands, Venezuela
(Miloslavich et al. 2010. Las Salinas & Quebrada La Iguana (Arrecifes, Vargas): Rivero et al. 2003).
Circumtropical (Valdés et al. 2006).
Genus Bursatella Blainville, 1817
35. Bursatella leachii Blainville, 1817
(Plate 4 A)
Material examined. SLR, 4 spcs, H = 60–70 mm, NRC. EOL, 2 spcs, H = 50–65 mm, IVICCM000020. OLH, 2
spcs, H = 80–90 mm, IVICCM000136. CHC, 1 spc., PNC.
Ecology. On mangrove lagoons on T. testudinum (1–3 m deep). This species releases a purple fluid when
disturbed.
Distribution. ABC Islands, Bermuda, Brazil, Colombia, Costa Rica, Cuba, Guadeloupe, Panama, Puerto Rico,
Trinidad & Tobago, USA, Venezuela (Valdés et al. 2006; Miloslavich et al. 2010. La Restinga, Margarita: Grune et
al. 2015). Circumtropical (Valdés et al. 2006).
Genus Dolabrifera Gray, 1847
36. Dolabrifera dolabrifera (Rang, 1828)
(Figure 2 L)
Material examined. SLR, 4 spcs, H = 40–60 mm, NRC. SPM, 2 spcs, H = 30–50 mm, NRC. MG, 1spc., H = 35
mm, IVICCM000095. WYL, 3 spcs, H = 20-30 mm, IVICCM000113. DCH, 1 spc., H = 17 mm, IVICCM000180.
EYL, 4 spcs, H = 60 mm, ZMBN 84907. YL, 1spc., H = 70 mm, ZMBN 84918.
Ecology. Common under rocks on reef lagoons (0.5–1 m deep).
Distribution. ABC Islands, Antigua, Bahamas, Barbados, Bermuda, Brazil, Cayman Islands, Colombia, Costa
Rica, Cuba, French Guiana, Guadeloupe, Mexico, Nicaragua, Puerto Rico, Saba, St. Martin/St. Maarten, St.
Vincent & the Grenadines, USA, Virgin Islands, Venezuela (Valdés et al. 2006; Massemin et al. 2009; Rios 2009;
Yidi & Sarmiento 2011. Dos Mosquises, north side of Southern Island, Los Roques: Marcus & Marcus 1967a, Los
Roques: Work 1969. La Restinga, Margarita: Grune et al. 2015). Circumtropical (Valdés et al. 2006).
Genus Petalifera Gray, 1847
37. Petalifera petalifera (Rang, 1828)
Distribution. ABC Islands, Bahamas, Barbados, Brazil, Cuba, Guadeloupe, Martinique, Puerto Rico, Venezuela
(Miloslavich et al. 2010. La Restinga, Margarita: Grune et al. 2015). Circumtropical (Valdés et al. 2006).
Remarks. No material of this species was found.
38*. Petalifera ramosa Baba, 1959
(Figure 2 M)
Material examined. APV, 1 spc., H = 20 mm, IVICCM000080.
Ecology. Sheltered areas, near mangroves, on T. testudinum.
Distribution. Bahamas, Belize, Colombia, Costa Rica, Cuba, Guadeloupe, Honduras, Jamaica, Martinique,
Mexico, Puerto Rico, USA, Venezuela. Circumtropical (Valdés et al. 2006).
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Remarks. This is the first record of the species in Venezuela and the new southern limit of its geographical
distribution.
Genus Phyllaplysia Fischer, 1872
39*. Phyllaplysia engeli Er. Marcus, 1955
(Plate 3 A)
Material examined. OLM, 2 spcs, H = 4–5 mm, MCC.
Ecology. Sheltered areas, near mangroves, at the internal base of the leaves of T. testudinum.
Distribution. ABC Islands, Bahamas, Barbados, Brazil, Colombia, Costa Rica, Cuba, Guadeloupe, Mexico,
Puerto Rico, St. Martin/St. Maarten, USA, Venezuela.
Remarks. Abundant, usually small, cryptic with the leaves of T. testudinum.
Genus Syphonota H. Adams & A. Adams, 1854
40*. Syphonota geographica (Adams & Reeve, 1850)
(Plate 5 K)
Material examined. CHC, 1 spc., PNC.
Ecology. Shallow sandy-muddy bottoms with T. testudinum. Active swimmer.
Distribution. Dominica, Dry Tortugas, St. Lucia, USA, Venezuela. Also found in the Indo-Pacific (Valdés et
al. 2006).
Remarks. Venezuela is the southern limit of the distribution of this species in the Western Atlantic.
Genus Stylocheilus Gould, 1852
41. Stylocheilus striatus (Quoy &Gaimard, 1832)
(Plate 3 B)
Material examined. BGM, 1 spc., H = 21 mm, MCC. AR, 1 spc., PNC.
Ecology. Mangrove lagoons, on rocky-muddy areas or seagrass (0.5–3 m deep).
Distribution. ABC Islands, Bahamas, Barbados, Bermuda, Brazil, Cayman Islands, Colombia, Costa Rica,
Cuba, Guadeloupe, Jamaica, Mexico, Puerto Rico, USA, Venezuela (Valdés et al. 2006). Cosmopolitan (Valdés et
al. 2006).
42*. Stylocheilus longicauda (Quoy & Gaimard, 1825)
Material examined. MR, 5 spcs, H = 30–35 mm, JOC. CMM, 1 spc., H = 30 mm, NRC.
Ecology. In coralline areas, with live coral and coral rubble.
Distribution. Brazil, Puerto Rico, USA, Venezuela. Circumtropical (Valdés et al. 2006).
Remarks. This is the first record of the species in Venezuela.
Superorder Panpulmonata Jörger, Stöger, Kano, Fukuda, Knebelsberger & Schrödl, 2010
Order Sacoglossa Ihering, 1876
Family Oxynoidae Stoliczka, 1868
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Genus Lobiger Krohn, 1847
43. Lobiger souverbii Fischer, 1857
(Figure 3 D)
Material examined. FCM, 2 spcs, H = 20 mm, JOC. MR, 1 spc., H = 25 mm, JOC. BGM, 1 spc., H = 22 mm,
MCC. IA, 1 spc., PNC.
Ecology. In sheltered areas on fronds of Caulerpa racemosa. This species releases a white fluid when
disturbed.
Distribution. ABC Islands, Bahamas, Barbados, Brazil, Costa Rica, Cuba, Cayman Islands, Guadeloupe,
Jamaica, Mexico, Puerto Rico, USA, Venezuela (Valdés et al. 2006. Los Roques: Gavagnin et al. 2000).
Remarks. This species is cryptic with the algae Caulerpa racemosa.
Genus Oxynoe Rafinesque, 1814
44. Oxynoe antillarum Mörch, 1863
(Figure 3 E)
Material examined. MR, 3 spcs, H = 9-13 mm, JOC. DCH, 2 spcs, H = 11-14 mm, IVICCM000107. IA, 1 spc.,
PNC.
Ecology. In sheltered areas with Caulerpa racemosa. It releases a white fluid when disturbed.
Distribution. ABC Islands, Bahamas, Barbados, Bermuda, Brazil, Cayman Islands, Costa Rica, Cuba,
Guadeloupe, Jamaica, Mexico, Panama, Puerto Rico, USA, Virgin Islands, Venezuela (Los Roques: Gavagnin et al.
2000. La Restinga, Margarita: Grune et al. 2015).
Remarks. It was found together with Lobiger souverbii.
45*. Oxynoe aguayoi Jaume, 1945
Material examined. FCM, 4 spcs, H = 7–8 mm, JOC.
Ecology. Found on Caulerpa racemosa.
Distribution. Cuba, Venezuela.
Remarks. This is the first record of this species in Venezuela. Marcus & Marcus (1967) considered O. aguayoi
a synonym of O. antillarum, arguing that Oxynoe shells are variable within species. Jensen (1980) described
Oxynoe azuropunctata based on specimens from Florida and stated that the type of O. aguayoi was lost, however,
the type of the latter species is housed in the collections of the Institute of Ecology and Systematics, Cuba, together
with the additional studied material used by Jaume (1945), which also includes shells and complete specimens of
O. antillarum sent by Mörch. In addition, the type locality of O. aguayoi is Jaimanitas, west coast of Havana
(Cuba), where both O. antillarum and O. aguayoi co-occur on Caulerpa racemosa (own data). The width/length
ratio varies between 1.5 and 1.84 in O. aguayoi, while in O. antillarum is 1.33 to 1.67 (Ortea et al. 1999). Valdés et
al. (2006) accepted the synonymy of the latter two species but left open the possibility that both could be valid.
Venezuela is the southern limit of the distribution of this species.
Family Boselliidae Ev. Marcus, 1982
Genus Bosellia Trinchese, 1891
46*. Bosellia curasoae Er. Marcus & Ev. Marcus, 1970
Material examined. MR, 1 spc., H = 10 mm, JOC.
Ecology. In sheltered sandy areas, on Halimeda tuna.
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Distribution. ABC Islands, Puerto Rico, Barbados, Bermuda, Brazil, Colombia, Guadeloupe, USA,
Venezuela.
Remarks. Both eastern and western populations of Boselia have been often considered conspecific under the
species name B. mimetica (e.g. Valdés et al. 2006; Rosenberg 2009; Yidi & Sarmiento 2011). Anatomical research
performed by the first author revealed however substantial anatomical differences (unpublished data). Moreover, a
molecular phylogenetic comparison of specimens identified as B. mimetica from the Mediterranean and Caribbean
regions showed these specimens to belong to different species (Carmona et al. 2011). It seems very likely that the
diversity of Boselia in the Atlantic Ocean is higher than so far recognized.
Family Plakobranchidae Gray, 1840
Genus Elysia Risso, 1818
47. Elysia crispata Mörch, 1863
(Figure 3 F)
Material examined. TT, 1 spc., H = 23 mm, IVICCM000099. WYL, 1 spc., H = 23 mm, IVICCM000100. LMD,
1 spc., H = 30 mm, IVICCM000115. PPD, 1 spc., H = 7 mm, IVICCM000004. DCH, 3 spcs, H = 40–50 mm,
IVICCM000106. DMR, 2 spcs, H = 25–85 mm, IVICCM0000010. PAM, 3 spcs, H = 20–30 mm, IVICCM000160.
EYL, 2 spcs, H = 20–30 mm, ZMBN 84909. OLM, 6 spcs, ZMBN 84929. APV, 10 spcs, H = 15–26 mm,
IVICCM000071. CBM, 15 spcs, H = 7–28 mm, NRC. CMM, 8 spcs, H = 7–22 mm, NRC. SLR, 22 spcs, H = 2–29
mm, NRC. CSM, 50 spcs, H = 4–31 mm, NRC. NPM, 3 spcs, H = 15–18 mm, NRC. APV, 7 spcs, H = 11–22 mm,
NRC. PNM, 1 spc., PNC.
Ecology. Crawling on rocks on reef fringe and inside mangrove lagoons (0.5–1 m) on Thalassia spp. and
mangrove roots.
Distribution. ABC Islands, Bahamas, Barbados, Belize, Bermuda, Brazil, Cayman Islands, Colombia, Costa
Rica, Cuba, Guadeloupe, Haiti, Honduras, Jamaica, Mexico, Panama, St. Martin/St. Maarten, St. Vincent & the
Grenadines, Trinidad & Tobago, USA, Virgin Islands, Venezuela (Marcus 1957; Marcus & Marcus 1960; Marcus
& Marcus 1962; Marcus & Marcus 1970; Marcus & Hughes 1974; Marcus 1977; Thompson 1977; Gavagnin et al.
2000; Valdés et al. 2006. Dos Mosquises, Los Roques, South Island: Marcus & Marcus 1967a. Los Roques: Work
1969. Mochima: Gavagnin et al. 1996; 1997. Golfo de Cariaco, Sucre: Fernandez & Jimenez 2007).
Remarks. This species was abundant in all sampling sites both on reef and mangrove environments.
According to Valdés et al. (2006) feeds upon Caulerpa sertularioides, but its diet likely encompasses a broader
range of items since Caulerpa was not always present on the areas where the species was observed. Marcus &
Marcus (1967a) record this species on the hard coral Acropora. Gavagnin et al. (1996; 1997) refer the presence of
protective chemicals against sunlight in specimens from Mochima, Venezuela.
48. Elysia ornata (Swainson, 1840)
(Figure 3 G)
Material examined. MR, 1 spc., H = 15 mm, JOC. YL, 1 spc., H~10 mm, ZMBN 84949. AR, 1 spc., PNC.
Ecology. Found on the reef fringe (1 m deep) associated with seaweeds or under rocks.
Distribution. ABC Islands, Bahamas, Barbados, Bermuda, Brazil, Colombia, Costa Rica, Cuba, Guadeloupe,
Jamaica, Mexico, St. Vincent & the Grenadines, Trinidad & Tobago, USA, Venezuela (Valdés et al. 2006).
Circumtropical (Valdés et al. 2006).
49. Elysia papillosa Verril, 1901
(Figure 5 M)
Material examined. ICM, 2 spcs, H = 6–30, 10 mm, JOC. APV, 1 spc., H = 12 mm, IVICCM000081. IA, 1 spc.,
PNC.
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Ecology. Under rocks, (0.5–4 m deep); also found on the seagrass T. testudinum.
Distribution. ABC Islands, Bahamas, Belize, Bermuda, Cayman Islands, Costa Rica, Cuba, Grenada,
Guadeloupe, Honduras, Martinique, Mexico, Panama, Sta. Lucia, Trinidad & Tobago, USA, Venezuela (Valdés et
al. 2006).
Remarks. Venezuela is the southern limit of the distribution of this species.
50. Elysia flava Verril, 1901
(Figure 3 H)
Material examined. PPD, 1 spc., H = 7.5 mm, IVICCM000002.
Ecology. Intertidal rocky shores, under rocks.
Distribution. Brazil, Bermuda, Costa Rica, Cuba, Guadeloupe, Jamaica, Mexico, Panama, Puerto Rico,
Venezuela (Valdés et al. 2006). Amphiatalntic (Valdés et al. 2006; García & Bertsch 2009).
51. Elysia serca Verrill, 1901
Distribution. ABC Islands, Barbados, Brazil, Puerto Rico, St. Martin, USA, Venezuela (La Restinga, Margarita:
Grune et al. 2015).
Remarks. No material of this species was found.
52*. Elysia cauze Er. Marcus, 1957
(Figure 3 J)
Material examined. BLR, 3 spcs, H = 7–16 mm, MCC.
Ecology. Mangrove lagoons on Caulerpa racemosa.
Distribution. ABC Islands, Barbados, Brazil, Guadeloupe, Jamaica, Puerto Rico, St. Martin/St. Maarten,
USA, Venezuela (La Restinga, Margarita: Grune et al. 2015: as E. subornata).
Remarks. This species is often considered a synonym of Elysia subornata Verrill, 1901 (that was
misidentified with E. cauze by Grune et al. 2015). The original description of the latter species is brief and do not
mention the presence of black rings on the mantle associated with glandular openings, a feature considered to be
diagnostic of E. cauze.
53*. Elysia tuca Ev. Marcus & Er. Marcus, 1967
(Figure 3 I)
Material examined. MR, 1 spc., H = 9 mm, JOC. APV, 1 spc., H = 9 mm, IVICCM000089. LJM, 4 spcs, H = 6–10
mm, MCC.
Ecology. On algae, but it can also be found associated with T. testudinum.
Distribution. ABC Islands, Bahamas, Barbados, Bermuda, Brazil, Cayman Islands, Costa Rica, Guadeloupe,
Jamaica, Mexico, Panama, Puerto Rico, USA, Venezuela.
54*. Elysia nisbeti Thompson, 1977
Material examined. MC, 2 spcs, H = 6–8 mm, JOC.
Ecology. On Caulerpa taxifolia.
Distribution. Bermuda, Guadeloupe, Jamaica, Venezuela.
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Remarks. The specimens of E. nisbeti with unknown origin, studied by Gavagnin et al. (2000: 1564, 1572-
1574) came from Mochima and Los Mosquises (Venezuela) (Jesus Ortea, op. cit.). Venezuela is the southern limit
of the distribution of this species.
55*. Elysia zuleicae Ortea & Espinosa, 2002
Material examined. FCM, 2 spcs, H = 2–8 mm, JOC.
Ecology. On algae of the genus Udothea.
Distribution. Bahamas, Costa Rica, Cuba, Guadeloupe, Venezuela.
Remarks. This is the new southern limit of the distribution of this species.
Genus Checholysia Ortea, Caballer, Moro & Espinosa, 2005
Remarks. Checholysia has been considered a junior synonym of Elysia by Bouchet (2015), but the members of
this genus bear a stylet in their penis, thus, we consider it as valid until new studies are performed.
56*. Checholysia patina (Ev. Marcus, 1980)
Material examined. MR, 4 spcs, H = 4–15 mm, JOC.
Ecology. On algae.
Distribution. Bahamas, Costa Rica, Guadeloupe, Mexico, USA, Venezuela.
Remarks. Venezuela is the southern limit of the distribution of this species.
57*. Checholysia annedupontae Ortea, Espinosa & Caballer, 2005 (in Ortea et al. 2005)
Material examined. MC, 5 spcs, H = 3–8 mm, JOC. APV, 4 spcs, H = 10–17 mm, MCC. GFM, 2 spcs, H = 12
mm, MCC. CVM, 2 spcs, H = 3–9 mm, MCC. LJM, 1 spc., H = 13 mm, MCC. OLM, 1 spc., H = 8 mm, MCC.
GCM, 2 spcs, H = 5, 10 mm, ZMBN 84921.
Ecology. On algae of the genera Bryopsis, Batophora, and on Avrainvillea longicaulis between 0.5–1 m deep.
Distribution. Cuba, Venezuela.
Remarks. This species, previously considered endemic to Cuba, was found abundantlly on leaves of the algae
A. longicaulis together with egg-masses. Venezuela is the southern limit of the distribution of this species.
Genus Thuridilla Bergh, 1872
58*. Thuridilla malaquita Ortea & Buske, 2014
(Figure 5 N)
Material examined. CHC, 1 spc., H = 7 mm, GCC.
Ecology. On algae of the genera Bryopsis, Batophora, and A. longicaulis between 0.5–1 m deep.
Distribution. Colombia, Costa Rica, Grenada, USA (as Thuridilla sp.), Martinique, Venezuela.
Remarks. Venezuela is the southern limit of the distribution of this species. This is the second record of the
species after the original description.
Family Caliphyllidae Tiberi, 1881
Genus Cyerce Bergh, 1870
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59*. Cyerce habanensis Ortea & Templado, 1989
(Figure 3 K)
Material examined. MR, 2 spcs, H = 12 mm, JOC. OLM, 2 spcs, H = 11–14 mm, MCC. IA, 1 spc., PNC.
Ecology. On algae of the genus Halimeda.
Distribution. Bahamas, Costa Rica, Cuba, Venezuela.
Remarks. Val d és et al. (2006) synonymyzed this species with Cyerce antillensis and considered Cyerce
edmundsi a valid species. However, the original description and the study of the holotype of the latter species
(BMNH No. 19776W) revealed the lack of a transverse groove in the foot, which is a typical character of the genus
Cyerce. Additionaly, C. edmundsi has pearl-like glands in the papillae, a feature absent in C. habanensis, and a
radula resembling those of Mourgona germaineae. Moreover, Thompson (1988) on a taxonomic discussion of the
genus Cyerce did not consider C. edmundsi a valid species. Likely, C. edmundsi is a synonym of M. germaineae
and C. habanensis a valid species. Venezuela is the southern limit of the distribution of this species.
Family Limapontiidae Gray, 1847
Genus Ercolania Trinchese, 1872
60. Ercolania coerulea Trinchese, 1892
Distribution. ABC Islands, Brazil, Jamaica, Mexico, St. Kitts/St. Christopher, St. Lucia, USA, Virgin Islands,
Venezuela (Valdés et al. 2006; Rosenberg 2009).
Remarks. No material of this species was found.
61*. Ercolania fuscata (Gould, 1870)
(Figure 3 L)
Material examined. OLH, 2 spcs, H = 3.2–3.8 mm, MCC.
Ecology. On filamentous green algae in muddy bottoms associated with mangrove lagoons.
Distribution. ABC Islands, Brazil, Canada, Cuba, Jamaica, Puerto Rico, USA, Venezuela.
Genus Stiliger Ehrenberg, 1828
62. Stiliger cricetus Er. Marcus & Ev. Marcus, 1970
Material examined. MC, 1 spc., H = 5 mm, JOC.
Ecology. On algae of the genus Bryopsis or Batophora oerstedi (1 m deep).
Distribution. ABC islands, Bahamas, Venezuela (Miloslavich et al. 2010: as Ercolania cricetus. Mochima
Bay: Mollo et al. 1994).
Remarks. Mollo et al. (1994) redescribed and reinstated S. cricetus based on one specimen from Venezuela.
Later, Valdés et al. (2006) recorded Ercolania coerulea in Venezuela and considered S. cricetus to be a synonym
but no reasons were given to explain the synonymization. Due to the complex taxonomy of the Sacoglossa and the
tendency for cryptic speciation in the group (Carmona et al. 2011), we maintain S. cricetus as a valid species until
additional data is available. Venezuela is the southern geographical limit of the distribution of this species.
Genus Placida Trinchese, 1876
63*. Placida cremoniana (Trinchese, 1892)
(Figure 5 O–P)
Material examined. CHC, 1 spc., PNC.
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Ecology. Shallow bottoms, on Bryopsis spp.
Distribution. Venezuela. Eastern Atlantic: Spain, Macaronesic Islands, Mediterranean (Cervera et al., 2006).
Remarks. This is the first record for the species in the Caribbean. The Eastern Pacific and Indo-Pacific records
of this species in Baja California, Mexico, Guam (Hermosillo et al. 2006) and Australia (Burn 1998) need further
investigation.
64. Placida kingstoni T.E. Thompson, 1977
Distribution. Bermuda, Costa Rica, Jamaica, Martinica, USA, Venezuela (La Restinga, Margarita: Grune et al.
2015).
Remarks. No material of this species was found. Venezuela is the southern limit of the distribution of this
species.
65. Placida verticilata Ortea, 1982
(Figure 3 M)
Material examined. HMM, 12 spcs, H = 2–4 mm, MCC.
Ecology. Inside and among the branches of Codium spp. in rocky bottoms (intertidal to 5 m deep).
Distribution. ABC Islands, Colombia, Costa Rica, Cuba, Guadeloupe, Jamaica, USA, Venezuela (El Horcón,
Macanao, Margarita Island: Caballer 2007). Amphiatlantic (Valdés et al. 2006; García & Bertsch 2009).
Remarks. Placida dendritica is a supposedly cosmopolitan species, difficult to distinguish from P. verticilata,
however, previous studies based on material from various Caribbean locations (Caballer 2007), suggest that the
records of P. dendritica from the Caribbean actually belong to P. verticilata. Venezuela is the southern limit of the
distribution of this species.
Genus Costasiella Pruvot-Fol, 1951
66*. Costasiella nonatoi Ev. Marcus & Er. Marcus, 1960
(Figure 3 N)
Material examined. CVM, 1 spc., H = 3 mm, MCC.
Ecology. On sandy-muddy bottoms with filamentous algae and empty shells with patches of Acetabularia spp.
and Avrainvillea spp.
Distribution. Bahamas, Brazil, Cayman Islands, Costa Rica, Puerto Rico, USA, Venezuela.
Family Cylindrobullidae Thiele, 1931
Genus Cylindrobulla Fischer, 1857
67. Cylindrobulla beauii Fischer, 1857
(Figure 3 O)
Material examined. LJM, 1 spc., H = 9 mm, ZMBN 84930.
Ecology. Near roots of Halimeda and Thalassia, on the vicinity of mangrove habitats.
Distribution. ABC Islands, Bahamas, Barbados, Belize, Bermuda, Brazil, Colombia, Cuba, Guadeloupe,
Jamaica, Mexico, Puerto Rico, Turks and Caicos, USA, Virgin Islands, Venezuela (Valdés et al. 2006; Rios 2009;
Yidi & Sarmiento 2011. Los Roques: Rosenberg 2009).
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Family Volvatellidae Pilsbry, 1895
Genus Ascobulla Ev. Marcus, 1972
68. Ascobulla ulla (Er. Marcus & Ev. Marcus, 1970)
Distribution. ABC Islands, Bahamas, Belize, Bermuda, Brazil, Cayman Islands, Costa Rica, Guadeloupe, Mexico,
Turks and Caicos, USA, Virgin Islands, Venezuela (Valdés et al. 2006; Rios 2009; Miloslavich et al. 2010. Vargas:
Rosenberg 2009).
Remarks. No material of this species was found.
Clade Nudipleura Wägele & Willan, 2000
Order Pleurobranchomorpha Pelseneer, 1906
Family Pleurobranchidae Gray, 1827
Genus Berthellina Gardiner, 1936
69. Berthellina quadridens (Mörch, 1863)
Material examined. CHC, 1 spc., PNC.
Ecology. Under rocks in shallow water.
Distribution. Barbados, Belize, Brazil, Colombia, Costa Rica, Cuba, Guadeloupe, Haiti, Mexico, Panama,
Puerto Rico, USA, Virgin Islands, Venezuela (Thompson 1977; Valdés et al. 2006; Rios 2009; Yidi & Sarmiento
2011: as Gymnotoplax quadridens; Ortea et al. 2014. Isla Larga (west end of north side, Los Roques): Marcus &
Marcus 1967a. Los Roques: Work 1969).
Genus Berthella Blainville, 1824
70. Berthella stellata (Risso, 1826)
Material examined. MC, 1 spc., H = 8 mm, JOC.
Ecology. Under rocks in exposed areas (1 m deep).
Distribution. Bahamas, Brazil, Cayman Islands, Cuba, Grenada, Guadeloupe, Mexico, St. Vincent & the
Grenadines, Venezuela (Valdés et al. 2006). Circumtropical (Valdés et al. 2006).
71. Berthella agassizii (MacFarland, 1909)
(Figure 3 C)
Material examined. PPD, 1 spc., H = 6 mm, IVICCM000003.
Ecology. On exposed rocky shores, under rocks.
Distribution. ABC Islands, Bermuda, Brazil, Colombia, Venezuela (Rios 1985; 2009).
Genus Pleurobranchus Cuvier, 1804
72. Pleurobranchus areolatus Mörch, 1863
(Figure 5 L)
Material examined. PCM, 3 spcs, H = 15 mm, JOC. IA, 1 spc., PNC.
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Ecology. Reef lagoons under coral rubble.
Distribution. ABC Islands, Bahamas, Barbados, Bermuda, Brazil, Cayman Islands, Colombia, Costa Rica,
Cuba, Guadeloupe, Jamaica, Mexico, Panama, Puerto Rico, St. Martin/St. Maarten, St. Thomas, USA, Virgin
Islands, Venezuela (Thompson 1977; Valdés et al. 2006; Rios 2009; Yidi & Sarmiento 2011. Isla de Aves: Marcus
& Marcus 1963. Isla de Aves and Los Roques, Isla Larga, west end, north side: Marcus & Marcus 1967a. Los
Roques: Work 1969; Marcus & Marcus 1970).
Genus Pleurobranchaea Leue, 1813
73. Pleurobranchaea inconspicua Bergh, 1897
Distribution. Argentina, Brazil, Colombia, Mexico, Surinam, USA, Venezuela (Valdés et al. 2006).
Remarks. No material of this species was found. West Africa (Valdés et al. 2006).
Order Nudibranchia Cuvier, 1817
Family Aegiridae P. Fischer, 1883
Genus Aegires Lovén, 1844
74*. Aegires sublaevis Odhner, 1932
Material examined. MR, 1 spc., H = 8 mm, JOC.
Ecology. Shallow rocky bottoms.
Distribution. Bahamas, Bermuda, Costa Rica, Guadeloupe, Honduras, Panama, USA, Virgin Islands,
Venezuela. Eastern Atlantic (Valdés et al. 2006).
Remarks. The occurrence of this species in Venezuela represents its southern limit.
75*. Aegires cf. ortizi Templado, Luque & Ortea, 1987
Material examined. OLH, 1 spc., H = 7 mm, IVICCM000132.
Ecology. On old or dead stalks of Halimeda spp.
Distribution. Bahamas, Cayman Islands, Cuba, Guadeloupe, USA, Venezuela.
Remarks. Venezuela constitutes the southern geographical limit of this species.
Family Goniodorididae H. Adams & A. Adams, 1854
Genus Okenia Menke, 1830
76. Okenia evelinae Er. Marcus, 1957
Distribution. Bahamas, Barbados, Brazil, Cuba, USA, Venezuela (Chimana Grande, Sucre: Valdés et al. 2006.
Mochima bay (Sucre): Ortea et al. 2009).
Remarks. No material of this species was found.
77. Okenia zoobotryon (Smallwood, 1910)
(Figure 5 Q)
Material examined. FCM, 1 spc., H = 9 mm, JOC. PTH, 1 spc., H = 5 mm, MCC. TA, 2 spcs, H = 8–10 mm,
IVICCM000006. AR, 2 spcs, H = 8-9 mm, GCC. AR, 1 spc., PNC.
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Ecology. On mangrove roots or submerged wood posts near surface, associated with the bryozoans Zoobotryon
spp. The spawn is a coiled translucent cord with 8–10 white eggs inside.
Distribution. Bermuda, Cuba, Mexico, Belice, Bahamas, Jamaica, Martinique, Puerto Rico, USA, Venezuela
(La Restinga, Margarita: Grune et al. 2015).
Remarks. An illustration of O. zoobotryon can be seen in Valdés et al. (2006) as Okenia sp. 2, while the
species named as O. zoobotryon is B. polycerelloides (Ortea et al. 2009). Pola (2014) designated a neotype from the
type locality, discussed the distribution of the species, and included B. polycerelloides in the list of synonymies of
O. zoobotryon without further discussion. The occurrence of this species in Venezuela constitutes the southern
geographical limit.
Genus Bermudella Odhner, 1941
78*. Bermudella polycerelloides Ortea & Bouchet, 1983
(Figure 4 C)
Material examined. BLR, 2 spcs, H = 4–5 mm, IVICCM000029.
Ecology. The spawn is a translucent flat mass found attached to the bottom with 1.8 mm diameter containing
large eggs located far apart from each other.
Distribution. Bermuda, Venezuela (Valdés et al. 2006: as Okenia zoobotryon). Eastern Atlantic: Canary
Islands (Ortea & Bouchet 1983).
Remarks. Bermudella polycerelloides Ortea & Bouchet, 1983 was considered a junior synonym of O.
zoobotryon by Valdés et al. (2006) and Pola (2014), but Ortea et al. (2009) separated both species and redefined the
characters that distinguish both taxa. The type locality of B. polycerelloides is the Canary Islands in the eastern
Atlantic and we do not discard the possibility that Caribbean specimens can belong to a distinct cryptic species of
Bermudella. Only future comparative anatomical and molecular phylogenetic work, will shead light of this
hypothesis. Venezuela is the southern limit of its distribution.
Family Polyceridae Alder & Hancock, 1845
Genus Polycera Cuvier, 1817
79. Polycera chilluna Er. Marcus, 1961
Distribution. USA, Venezuela (Valdés et al. 2006).
Remarks. This species is only known from two localities in the western Atlantic, Venezuela is the southern
limit of its distribution. No material of this species was found.
80. Polycera odhneri Er. Marcus, 1955
(Figure 4 E)
Material examined. ECU, 1 spc., H = 10 mm, MCC. BLR, 8 spcs, H = 1–17.5 mm, MCC. TA, 25 spcs, H = 8–25
mm, IVICCM000008.
Ecology. Crawling on briozoans on mangrove roots. The spawn is a coiled ribbon with small eggs that can
reach up to 33 mm in length.
Distribution. ABC Islands, Bahamas, Barbados, Bermuda, Brazil, Cuba, Guadeloupe, USA, Venezuela
(Valdés et al. 2006. La Restinga, Margarita: Grune et al. 2015).
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Genus Polycerella Verril, 1880
81. Polycerella emertoni Verrill, 1880
Material examined. FCM, 1 spc., H = 9 mm, JOC.
Ecology. On submerged wood posts, near surface.
Distribution. Bahamas, Brazil, Cuba, USA, Venezuela (Marcus & Marcus 1960; 1964; 1970; Rios 1985;
Va ld é s et al. 2006; Rios 2009; Miloslavich et al. 2010. Anzoategui: Rosenberg 2009).
Genus Tambja Burn, 1962
82. Tambja cf. stegosauriformis Pola, Cervera & Gosliner, 2005
Distribution. Brazil, Venezuela (Valdés et al. 2006).
Remarks. Val d és et al. (2006) reported this species for Venezuela pointing out that their specimens had a
different colour pattern than the animals from Brazil. Venezuela is the northern limit of the distribution of this
species. No material of this species was found.
Family Hexabranchidae Bergh, 1891
Genus Caribranchus Ortea, Caballer & Moro, 2003
83. Caribranchus morsomus (Ev. Marcus & Er. Marcus, 1962)
(Figure 4 F–G)
Material examined. CSM, 1 spc., H = 60 mm, MCC.
Ecology. Rocky shores (5–20 m deep), feeding on red sponge. Due to his colour and swimming behaviour, has
been called “Spanish dancer”.
Distribution. ABC Islands, Cayman Islands, Colombia, Costa Rica, Guadeloupe, Panama, Puerto Rico, USA,
Virgin Islands, Venezuela (Valdés et al. 2006).
Remarks. Val d és et al. (2006) considered the (Caribbean) genus Caribranchus a synonym of (the Indo-
Pacific) Hexabranchus arguing the lack of phylogenetic evidences to support the former genus. However,
Caribranchus is characterized by presence of tubercles on the dorsum, smooth labial cuticle, and radula with
medial tooth, whereas Hexabranchus have smooth dorsum, armed labial cuticles with sticks, and radula lacking
medial tooth. We interprete such differences are sufficient to continue with both genera, even if both are
monotypic. Venezuela is the southern limit of the distribution of this species.
Family Dorididae Rafinesque, 1815
Genus Aphelodoris Bergh, 1879
84. Aphelodoris antillensis Bergh, 1879
(Figure 6 A)
Material examined. PCM, 1 spc., H = 20 mm, JOC. IA, 1 spc., PNC.
Ecology. Under slabs of coral reef down to 10 m deep.
Distribution. ABC Islands, Antigua, Barbados, Cayman Islands, Costa Rica, Dominican Republic, Grenada,
Guadeloupe, Jamaica, Martinique, Mexico, Panama, Puerto Rico, St. Lucia, St. Vincent & the Grenadines, USA,
Virgin Islands, Venezuela (Valdés et al. 2006; Miloslavich et al. 2010. Margarita Island: Rosenberg 2009).
Remarks. Venezuela is the southern limit of the distribution of this species.
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Genus Doris Linnaeus, 1758
85. Doris verrucosa Linnaeus, 1758
Distribution. Brazil, Costa Rica, USA Venezuela (La Restinga, Margarita: Grune et al. 2015).
Remarks. No material of this species was found.
86. Doris bovena Er. Marcus, 1955
(Figure 4 D)
Material examined. PG, 2 spcs, H = 14 mm, JOC. FCM, 2 spcs, H = 12.5–15 mm, JOC. TA, 1 spcs, H = 20 mm,
IVICCM000009.
Ecology. Associated to roots of red mangroves inside coastal lagoons (0.5–1 m deep).
Distribution. ABC Islands, Brazil, Guadeloupe, Mexico, USA, Venezuela (Valdés et al. 2006).
87. Doris kyolis (Ev. Marcus & Er. Marcus, 1967a)
Material examined. FCM, 1 spc., H = 15 mm, JOC. FCM, 7 spcs, H = 12–25 mm, JOC. MR, 1 spc., H = 14 mm,
JOC.
Ecology. Shallow waters. Under rocks, feeding on blue sponges.
Distribution. ABC Islands, Bahamas, Barbados, Brazil, Colombia, Grenada, Guadeloupe, Puerto Rico, St.
Lucia, St. Vincent & the Grenadines, USA, Virgin Islands, Venezuela (Valdés et al. 2006. La Restinga, Margarita:
Grune et al. 2015).
Remarks. Venezuela is the southern limit of the distribution of this species.
Family Discodorididae Bergh, 1891
Genus Discodoris Bergh, 1877
88. Discodoris branneri MacFarland, 1909
Material examined. BSM, 1 spc., H = 27 mm, NRC. CHC, 1 spc., PNC.
Ecology. Under rocks, on the intertidal. It autotomizes parts of the mantle when disturbed.
Distribution. Bahamas, Barbados, Brazil, Cayman Islands, Colombia, Costa Rica, Guadeloupe, Honduras,
Jamaica, Martinique, Panama, Puerto Rico, St. Lucia, St. Vincent & the Grenadines, USA, Venezuela (Valdés et al.
2006).
Remarks. Discodoris evelinae Er. Marcus, 1955, one of the most cited species in the Caribbean, was included
in the list of synonyms of D. branneri by Dayrat (2010).
Genus Jorunna Bergh, 1876
89*. Jorunna spazzola (Er. Marcus, 1955)
Material examined. MR, 2 spcs, H = 12–20 mm, JOC.
Ecology. Under rocks, in shallow waters. It changes color when captured.
Distribucion. ABC Islands, Barbados, Brazil, Costa Rica, Cuba, Guadeloupe, Honduras, Mexico, Turks and
Caicos, USA, Virgin Islands, Venezuela.
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Genus Geitodoris Bergh, 1891
90. Geitodoris immunda Bergh, 1894
Distribution. Brazil, Costa Rica, USA, Venezuela (Valdés et al. 2006).
Remarks. No material of this species was found.
Genus Platydoris Bergh, 1877
91*. Platydoris angustipes (Mörch, 1863)
(Figure 6 B)
Material examined. MR, 5 spcs, H = 40–60 mm, JOC. CHC, 1 spc., PNC.
Ecology. Rocky bottoms, 2 m deep, under rocks.
Distribution. ABC Islands, Antigua, Bahamas, Barbuda, Brazil, Cayman Islands, Costa Rica, Cuba, Grenada,
Guadeloupe, Honduras, Jamaica, Martinique, Mexico, Panama, St. Barthelemy/St. Bartholomew, St. Martin/St.
Maarten, St. Lucia, St. Vincent & the Grenadines, Turks and Caicos, Puerto Rico, USA, Virgin Islands, Venezuela.
Genus Sclerodoris Eliot, 1904
92*. Sclerodoris prea (Ev. Marcus & Er. Marcus, 1967)
(Figure 6 C)
Material examined. MR, 1 spc., H = 17 mm, JOC. IA, 1 spc., PNC.
Ecology. Shallow waters, collected at 1 m deep.
Distribution. Bahamas, Barbados, Guadeloupe, Jamaica, USA, Venezuela.
Remarks. Venezuela represents the southern limit of the distribution of this species.
Familiy Dendrodorididae O'Donoghue, 1924
Genus Dendrodoris Ehrenberg, 1831
93. Dendrodoris warta Ev. Marcus & Gallagher, 1976
(Figure 4 I)
Material examined. AM, 1 spc., H = 62 mm, MCFUDENA
Ecology. Shallow water under rocks.
Distribution. USA, Venezuela (Valdés et al. 2006. La Restinga, Margarita: Grune et al. 2015).
Remarks. This species is only known from two countries in the western Atlantic, Venezuela is the southern
limit of its distribution.
94. Dendrodoris krebsii (Mörch, 1863)
(Figure 4 H)
Material examined. MC, 1 spc., H = 30 mm, JOC. PCM, 1 spc., H = 30 mm, JOC. NPM, 2 spcs, H = 22–25 mm,
NRC. ECU, 1 spc., H = 23 mm, MCC. APV, 3 spcs, H = 20–35 mm, MCC. DCH, 2 spcs, H = 40 mm,
IVICCM000108. PNM, 1 spc., H = 35 mm, GCC. PNM, 1 spc., PNC.
Ecology. Shallow waters until 10 m deep.
Distribution. ABC Islands, Antigua, Bahamas, Barbados, Belize, Brazil, Colombia, Cayman Islands,
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Colombia, Costa Rica, Cuba, Dominican Republic, Grenada, Guadeloupe, Honduras, Jamaica, Martinique,
Mexico, Panama, St. Kitts/St. Christopher, St. Lucia, St. Martin/St. Maarten, St. Vincent & the Grenadines, USA,
Virgin Islands, Venezuela (Valdés et al. 2006; Rios 2009. Los Roques, Dos Mosquises, south end of Southern
Island: Marcus & Marcus 1967a. Los Roques: Work 1969. La Restinga, Margarita: Grune et al. 2015).
Remarks. This species shows high chromatic variability with specimens depicting tones of red, black, white or
a combination of colours.
Family Cadlinidae Bergh, 1891
Genus Cadlina Bergh, 1878
95*. Cadlina rumia Er. Marcus, 1955
(Figure 4 B)
Material examined. PTH, 1 spc., H = 7 mm, MCC.
Ecology. On rocky bottoms with patches of Thalassia and sponges.
Distribution. ABC Islands, Bahamas, Brazil, Cuba, Guadeloupe, Jamaica, Panama, St. Martin/St. Maarten,
USA, Venezuela.
Family Chromodorididae Bergh, 1891
Genus Tyrinna Bergh, 1898
96. Tyrinna evelinae (Er. Marcus, 1958)
(Figure 4 L)
Material examined. NPM, 1 spc., H = 10 mm, NRC.
Ecology. Shallow waters around 3 m deep.
Distribution. Brazil, Costa Rica, Dominican Republic, Jamaica, Puerto Rico, Venezuela (National Park of
Morrocoy, Falcon: Ortea et al. 2003a; Rios 2009). In the Pacific: from Gulf of California, to northern Perú (Schrödl
& Millen 2001; Uribe et al. 2013).
Genus Felimida Ev. Marcus, 1971b
97*. Felimida binza (Ev. Marcus & Er. Marcus, 1963)
(Figure 4 J)
Material examined. NPM, 2 spcs, H = 8–14 mm, NRC. MC, 1 spc., H = 15 mm, JOC. OLM, 1 spc., H = 6 mm,
MCC.
Ecology. On Caulerpa racemosa near mangroves around 3 m deep.
Distribution. ABC Islands, Brazil, Colombia, Costa Rica, Cuba, Guadeloupe, USA, Virgin Islands,
Venezuela.
98*. Felimida clenchi (Russel, 1935)
(Figure 4 K)
Material examined. CSM, 1 spc., H = 10 mm. ZMBN 84946. CSM, 2 spcs, H = 8–9 mm, MCC. CHC, 1 spc.,
PNC.
Ecology. On coral reefs and under rocks; around 4 m deep.
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Distribution. USA, ABC Islands, Bahamas, Barbados, Bermuda, Brazil, Cayman Islands, Colombia, Costa
Rica, Guadeloupe, Jamaica, Panama, St. Lucia, Virgin Islands, Venezuela.
99. Felimida grahami Thompson, 1980
Distribution. Jamaica, Panama, Puerto Rico, St. Lucia, St. Martin/St. Maarten, St. Vincent & the Grenadines,
Venezuela (Valdés et al. 2006. Dependencias Federales: Rosenberg 2009).
Remarks. No material of this species was found. Venezuela is the southern limit of the distribution of this
species.
Genus Felimare (Ev. Marcus & Er. Marcus, 1963)
100. Felimare acriba (Ev. Marcus & Er. Marcus, 1967)
(Figure 6 D)
Material examined. CNM, 1 spc., PNC.
Ecology. On Acropora palmata.
Distribution. Costa Rica, Guadeloupe, Mexico, Puerto Rico, St. Lucia, St. Martin/St. Maarten, Venezuela
(Archipielago de Los Roques: Marcus & Marcus 1967a; Work 1969; Ortea et al. 1996).
Remarks. Specimens from Dos Mosquises, Los Roques (Venezuela) were used in the original description of
this species, where the type locality was not specified. Venezuela is the southern limit of the distribution of this
species.
101*. Felimare fregona (Ortea & Caballer in Ortea, Espinosa, Buske & Caballer, 2013)
(Figure 6 E–F)
Material examined. AR, 1 spc., H = 22 mm, GCC. AR, 1 spc., PNC.
Ecology. On rocky bottoms feeding on blue sponges.
Distribution. Curaçao, Guadeloupe, Puerto Rico, Virgin Islands, Venezuela (Valdés et al. 2006: as
Hypselodoris sp. 3).
Remarks. Venezuela is the southern limit of the distribution of the species.
102*. Felimare kempfi (Ev. Marcus, 1971a)
(Figure 6 G)
Material examined. CHC, 1 spc., PNC.
Ecology. On rocky bottoms.
Distribution. Brazil, Costa Rica, Mexico, Panama, Puerto Rico, USA, Venezuela.
103. Felimare marci (Ev. Marcus, 1971c)
Distribution. Belize, Brazil, Jamaica, Mexico, Venezuela (Ortea et al. 1996; Valdés et al. 2006; Rios 2009. Los
Roques Archipelago: Marcus 1970).
Remarks. Originally described based on specimens from Brazil and Venezuela that seem to belong to two
different species. Dominguez (2006) and García et al. (2008) redescribed this species based on Brazilian
specimens. The taxonomic status of Venezuelan specimens remains to be clarified. The record of this species in
Mexico (Ortea et al. 1996) corresponds in fact to Hypselodoris olgae Ortea & Bacallado, 2007. No material of this
species was found.
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104. Felimare molloi (Ortea & Valdés in Ortea, Valdés, & García-Gómez, 1996)
Distribution. Mexico, Venezuela (Miloslavich et al. 2010. Isla Picuda, Chimanas, Mochima: Ortea et al. 1996).
Remarks. This species was considered endemic to Venezuela but Ortigosa (2009: p. 42, Fig. 50) recorded it as
Mexichromis sp. in Yucatan, Mexico, including data on the internal anatomy. Venezuela is the southern limit of the
distribution of the species and the type locality (Isla Picuda, Chimanas, Mochima). No material of this species was
found.
105*. Felimare picta (Schultz in Philippi, 1836)
Material examined. SLR, 3 spcs, H = 10–30 mm, NRC.
Ecology. On Thalassia digitum near mangroves, also on dead coral; around 1 m deep.
Distribution. Brazil, Puerto Rico, USA (Ortea et al. 1996), Venezuela. Widespread in both sides of the
Atlantic (Valdés et al. 2006; García & Bertsch 2009).
106. Felimare ruthae (Ev. Marcus & Hughes, 1974)
(Figure 4 M)
Material examined. SPM, 2 spcs, H = 10–12 mm, NRC. CPM, 1 spc., H = 13 mm, ZMBN 84943. CSM, 3 spcs, H
= 14 mm, ZMBN 84945. CHC, 1 spc., H = 12 mm, GCC. CHC, 1 spc., PNC.
Ecology. On coral reefs, under rocks, and on coral rubble, down to 5 m deep.
Distribution. Bahamas, Barbados, Costa Rica, Cuba, Guadeloupe, Jamaica, Mexico, Panama, Venezuela
(Valdés et al. 2006).
Remarks. Common in coral habitats in the National Park of Morrocoy. Venezuela is the southern limit of the
distribution of this species.
107. Felimare samueli (Caballer & Ortea, 2012)
(Figure 4 N)
Material examined. OLM, 3 spcs, H = 10–85 mm, MCC. IA, 1 spc., PNC.
Ecology. On sand with T. testudinum and Halimeda spp., near mangroves, 1–2 m deep.
Distribution. Venezuela (Caballer & Ortea, 2012).
Family Arminidae Iredale & O'Donoghue, 1923
Genus Armina Rafinesque, 1814
108. Armina muelleri (Ihering, 1886)
Distribution. Brazil, Colombia, Dominica, Mexico, Surinam, USA, Venezuela (Valdés et al. 2006; Rosenberg
2009).
Remarks. No material of this species was found.
Family Flabellinidae Bergh in Carus, 1889
Genus Flabellina Gray in Griffith & Pidgeon, 1833
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109*. Flabellina dushia (Ev. Marcus & Er. Marcus, 1963)
(Figure 6 H)
Material examined. CHC, 1 spc., PNC. IA, 1 spc., PNC.
Ecology. On algae with hydroids.
Distribution. Bahamas, Brazil, Costa Rica, Curaçao, Guadeloupe, Jamaica, Martinique, USA, Venezuela.
110. Flabellina engeli engeli Ev. Marcus & Er. Marcus, 1968
(Figure 6 I)
Material examined. CHC, 1 spc., H = 18 mm, GCC. CNM, 1 spc., PNC.
Ecology. Crawling on bottoms with Halimeda spp. and rubble.
Distribution. Barbados, Colombia, Costa Rica, Cuba, Curaçao, Grenada, Martinique, Puerto Rico, St Lucia,
USA, Venezuela (Valdés et al. 2006: as F. engeli).
Remarks. This name probably refers to a complex of species, or at least subspecies (DaCosta et al. 2007).
Venezuela is the southern limit of the distribution of F. engeli engeli, the records to F. e n g e l i from Brazil (Padula &
Santos 2006; Padula et al. 2012) are owed to F. engeli lucianae (DaCosta et al. 2007).
111. Flabellina verta (Ev. Marcus, 1970)
(Figure 6 J)
Material examined. CHC, 1 spc., PNC. CHC, 1 spc., PNC.
Ecology. Shallow waters. On hydroids of the genus Eudendrium.
Distribution. Brazil, Virgin Islands, USA, Venezuela (Los Testigos Island: Millen & Hamann 2006).
112. Flabellina marcusorum Gosliner & Kuzirian, 1990
Material examined. CHC, 1 spc., H = 18 mm, GCC.
Ecology. Shallow barrier reefs near sandy beaches or mangrove areas.
Distribution. ABC Islands, Brazil, Costa Rica, Venezuela (Valdés et al. 2006; Rios 2009; Yidi & Sarmiento
2011). Tropical Eastern Pacific (Valdés et al. 2006).
Family Facelinidae Bergh in Carus, 1889
Genus Austraeolis Burn, 1962
113*. Austraeolis catina Ev. Marcus & Er. Marcus, 1967
(Figure 6 K)
Material examined. CHC, 1 spc., PNC.
Ecology. Rocky bottoms with hydroids.
Distribution. Bahamas, Belize, Puerto Rico, St. Lucia, USA, Venezuela.
Remarks. Venezuela is the southern limit of the distribution of this species.
Genus Dondice Er. Marcus, 1958
114. Dondice occidentalis Engel, 1925
(Figure 5 A)
Material examined. LJM, 1 spc., H = 34 mm, MCC.
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Ecology. On algae with hydroids attached.
Distribution. ABC Islands, Bahamas, Belize, Bermuda, Brazil, Cayman Islands, Colombia, Costa Rica, Cuba,
Grenada, Guadeloupe, Jamaica, Martinique, Mexico, Panama, St. Martin/St. Maarten, St. Vincent & the
Grenadines, Trinidad & Tobago, Turks and Caicos, USA, Venezuela (Marcus & Marcus 1970; Marcus 1976; 1977;
Thompson 1980; Rios 1985; Valdés et al. 2006; Rios 2009; Miloslavich et al. 2010. Barcelona, Anzoategui:
Marcus & Marcus 1964. Guanta, Anzoategui: Marcus & Marcus 1967c. La Restinga, Margarita: Grune et al.
2015).
115. Dondice parguerensis Brandon & Cutress, 1985
Material examined. EOL, 1 spc., H = 34 mm, IVICCM000023. ML, 1 spc., H = 19 mm, MCC.
Ecology. This is a rare species occuring associated with the jellyfish Cassiopea xamachana. The spaw is a
white egg cord found around the arms of the jellyfish.
Distribution. Bermuda, Guadeloupe, Panama, Puerto Rico, Venezuela (Higuerote, Miranda: Mariño et al.
2011. La Restinga, Margarita: Grune et al. 2015).
Remarks. Venezuela is the southern limit of the distribution of this species. It is likely that an adequate
sampling on Cassiopea jellyfish across the Caribbean would confirm the presence of this species on other
localities.
Genus Godiva Macnae, 1954
116*. Godiva rubrolineata Edmunds, 1964
(Figure 5 B)
Material examined. BLR, 2 spcs, H = 6–6.5 mm, MCC. CNM, 1 spc., H = 7 mm, GCC. CHC, 1 spc., PNC.
Ecology. On algae with hydroids attached.
Distribution. ABC Islands, Brazil, Cuba, Guadeloupe, Jamaica, Puerto Rico, USA, Venezuela.
Genus Favorinus Gray, 1850
117. Favorinus auritulus Er. Marcus, 1955
(Figure 5 C)
Material examined. OLH, 6 spcs, H = 5–10 mm, MCC.
Ecology. Coastal lagoons surrounded by mangroves; found on spaws of Bulla occidentalis, upon which they
seem to feed on.
Distribution. ABC Islands, Antigua, Bahamas, Barbados, Bermuda, Brazil, Cuba, Guadeloupe, Jamaica,
Martinique, Puerto Rico, Trinidad & Tobago, USA, Venezuela (Laguna de la Restinga: Villaba & Crescini 2013. La
Restinga, Margarita: Grune et al. 2015).
Genus Nanuca Er. Marcus, 1957
118*. Nanuca sebastiani Er. Marcus, 1957
(Figure 6 L)
Material examined. IA, 1 spc., PNC.
Ecology. Shallow waters. On algae with hydroids attached.
Distribution. Barbados, Bermuda, Bonaire, Brazil, Cayman Islands, Costa Rica, Cuba, Curaçao, Guadeloupe,
Martinique, Mexico, USA, Virgin Islands, Venezuela.
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Genus Phidiana Gray, 1850
119. Phidiana lynceus Bergh, 1867
(Figure 5 D)
Material examined. MC, 5 spcs, H = 15–29 mm, JOC. MR, 1 spc., H = 19 mm, JOC. BLR, 4 spcs, H = 14–32 mm,
MCC. PPD, 1 spc., H = 16 mm, MMC. EYL, 1 spc., H = 17 mm, MMC. TA, 3 spcs, H = 17–32 mm,
IVICCM000005. TA, 3 spcs, H = 11–22 mm, IVICCM000012. PPD, 2 spcs, H = 10–17 mm, IVICCM000001. PC,
1 spc., H = 20 mm, IVICCM000161. TA, 20 spcs, H = 8–32 mm, IVICCM000172. BLR, 3 spcs, H = 12–40 mm,
IVICCM000028. WYL, 1 spc., H = 6 mm, IVICCM000112. CHC, 1 spc., PNC.
Ecology. Very common on shallow water, on algae, mangrove roots and reefs.
Distribution. ABC Islands, Bahamas, Barbados, Brazil, Colombia, Costa Rica, Guadeloupe, Jamaica,
Martinique, Mexico, St. Lucia St. Martin/St. Maarten, St. Vincent & the Grenadines, USA, Virgin Islands,
Venezuela (Valdés et al. 2006; Miloslavich et al. 2010. La Restinga, Margarita: Grune et al. 2015).
Genus Learchis Bergh, 1896
120*. Learchis poica Ev. Marcus & Er. Marcus, 1960
Material examined. CHC, 1 spc., PNC.
Ecology. On hydroids.
Distribution. Aruba, Bahamas, Barbados, Cayman Islands, Costa Rica, Curaçao, Grenada, Jamaica,
Martinique, Mexico, Puerto Rico, St. Lucia, St. Vincent & the Grenadines, Virgin Islands, USA, Venezuela.
Remarks. Venezuela is the southern limit of the distribution of this species.
121. Learchis? ignis Crescini, De Sisto & Villalba, 2013a
Distribution. Venezuela (Bay of Charagato, Cubagua Island: Crescini et al. 2013a).
Remarks. No material of this species was found. Endemic to Venezuela. This species was placed in the genus
Learchis, nevertheless it has a radula with lateral teeth which excludes this species from the family Facelinidae.
Family Aeolidiidae Gray, 1827
Genus Spurilla Bergh, 1864
122. Spurilla braziliana MacFarland, 1909
Material examined. MC, 1 spc., H = 15 mm, JOC.
Ecology. Shallow water, on anemones under rocks.
Distribution. ABC Islands, Barbados, Belize, Bermuda, Brazil, Colombia, Colombia, Costa Rica, Cuba,
Guadeloupe, Honduras, Jamaica, Mexico, Puerto Rico, St. Vincent & the Grenadines, USA, Venezuela (Valdés et
al. 2006. La Restinga, Margarita: Grune et al. 2015: as S. neapolitana). In the Pacific: Australia, Costa Rica,
Hawaii, Japan & Peru (Carmona et al. 2014).
Remarks. Carmona et al. (2014) recently revised the species of the Spurilla neapolitana (Delle Chiaje, 1841)
complex, they established that all the records to this species in the Caribbean were owed to S. braziliana, but
omitted several records (recovered here), and assumed as their own the record of the species in Cuba, which is
indeed owed to Espinosa et al. (2006).
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123*. Spurilla sargassicola Bergh, 1871
(Figure 5 E)
Material examined. BGM, 1 spc., H = 15 mm, MCC.
Ecology. On Caulerpa spp.
Distribution. Bahamas, Mexico, Sargasso Sea, Venezuela.
Remarks. Venezuela is the southern limit of the distribution of this species.
Genus Berghia Trinchese, 1877
124. Berghia creutzbergi Er. Marcus & Ev. Marcus, 1970
(Figure 6 M)
Material examined. ICM, 1 spc., H = 15 mm, JOC. CHC, 1 spc., PNC.
Ecology. Under rocks; around 1 m deep.
Distribution. ABC Islands, Bahamas, Barbados, Brazil, Cayman Islands, Costa Rica, Guadeloupe, USA,
Venezuela (Valdés et al. 2006. Dependencias Federales: Rosenberg 2009).
125. Berghia rissodominguezi Muniain & Ortea, 1999
Distribution. ABC Islands, Argentina, Brazil, Jamaica, Mexico, USA, Venezuela (La Restinga, Margarita: Grune
et al. 2015).
Remarks. No material of this species was found.
Genus Anteaeolidiella Miller, 2001
126. Anteaeolidiella lurana (Ev. Marcus & Er. Marcus, 1967d)
Distribution. ABC Islands, Bermuda, Brazil, Mexico, Puerto Rico, Venezuela (Valdés et al. 2006. Dependencias
Federales: Rosenberg 2009). Eastern Atlantic: Italy, Spain. Indo-Pacific: Australia (Carmona et al. 2014b).
Remarks. No material of this species was found.
Family Eubranchidae Odhner, 1934
Genus Eubranchus Forbes, 1838
127*. Eubranchus conicla (Er. Marcus, 1958)
(Figure 5 F)
Material examined. ECU, 3 spcs, H = 2–3 mm, MCC.
Ecology. On bryozoans growing on wooden poles near the surface.
Distribution. Bahamas, Barbados, Brazil, Costa Rica (?), Cuba, Honduras, Jamaica, Trinidad and Tobago,
USA, Venezuela.
Remarks. The reference of this species in Costa Rica is based on Valdés et al. (2006) who considered E.
convenientis (described based on specimens from Costa Rica) to be a synonym of E. conicla.
Family Tergipedidae Bergh, 1889
Genus Cuthona Alder & Hancock, 1855
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128*. Cuthona iris Edmunds & Just, 1983
(Figure 6 N)
Material examined. CHC, 1 spc., PNC.
Ecology. Shallow water, crawling under rocks.
Distribution. Barbados, Costa Rica, St. Vincent & the Grenadines, Venezuela.
Remarks. Venezuela is the southern limit of the distribution of this species.
Family Tritoniidae Lamarck, 1809
Genus Marionia Vayssière, 1877
129. Marionia tedi Ev. Marcus, 1983
Distribution. Mexico, USA, Yucatan Strait, Venezuela (Valdés et al. 2006; Miloslavich et al. 2010. Isla Margarita:
Rosenberg 2009).
Remarks. No material of this species was found. Venezuela is the southern limit of the distribution of this
species.
Genus Tritoniopsis Eliot, 1905
130*. Tritoniopsis frydis Er. Marcus & Ev. Marcus, 1970
(Figure 6 O)
Material examined. CHC, 1 spc., PNC.
Ecology. On sea fans.
Distribution. Bahamas, Belize, Bermuda, Cayman Islands, Curaçao, Guadeloupe, Honduras, USA, Venezuela.
Remarks. Venezuela is the southern limit of the distribution of this species.
Family Hancockiidae MacFarland, 1923
Genus Hancockia Gosse, 1877
131. Hancockia ryrca Er. Marcus, 1957
Distribution. ABC Islands, Bahamas, Barbados, Brazil, Venezuela (Valdés et al. 2006. Dependencias Federales:
Rosenberg 2009).
Remarks. No material of this species was found.
Family Bornellidae Bergh, 1874
Genus Bornella Gray, 1850
132*. Bornella calcarata Mörch, 1863
Material examined. SLR, 2 spcs, H = 40–50 mm, NRC.
Ecology. Inhabits mainly in reefs, but it can be found in rocky shelfs, seagrass beds, coral rubble, sandy
bottoms and algae, because this species is able to swim.
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FIGURE 2. A, Migaya felis, living specimen and spawn. B, Navanax gemmatus. C, Chelidonura hirundinina. D, Philine
buchensis. E, Bulla occidentalis, juvenile. F, Bulla occidentalis. G, Haminoea cf. elegans. H, Lapinura divae. I, Aplysia
dactylomela. J, Aplysia brasiliana. K, Aplysia juliana. L, Dolabrifera dolabrifera. M, Petalifera ramosa.
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FIGURE 3. A, Phyllaplysia engeli. B, Stylocheilus striatus. C, Berthella agassizii. D, Lobiger souverbii. E, Oxynoe antillarum.
F, Elysia crispata. G, Elysia ornata. H, Elysia flava. I, Elysia tuca. J, Elysia cauze. K, Cyerce habanensis. L, Ercolania fuscata.
M, Placida verticilata. N, Costasiella nonatoi. O, Cylindrobulla beauii.
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FIGURE 4. A, Bursatella leachii. B, Cadlina rumia. C, Bermudella polycerelloides. D, Doris bovena. E, Polycera odhneri. F,
Caribranchus morsomus, ventral view. G, Caribranchus morsomus, dorsal view. H, Dendrodoris krebsii. I, Dendrodoris warta
(fixed). J, Felimida binza. K, Felimida clenchi. L, Tyrinna evelinae. M, Felimare ruthae. N, Felimare samueli.
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FIGURE 5. A, Dondice occidentalis. B, Godiva rubrolineata. C, Favorinus auritulus. D, Phidiana lynceus. E, Spurilla
sargassicola. F, Eubranchus conicla. G, Scyllaea pelagica. H, Doto casandra. I, Chelidonura cubana. J, Aplysia parvula. K,
Syphonota geographica. L, Pleurobranchus areolatus. M, Elysia papillosa. N, Thuridilla malaquita. O–P, Placida cremoniana.
Q, Okenia zoobotryon. I–Q, Sea slugs in their natural environment (Gaby Carias).
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FIGURE 6. A, Aphelodoris antillensis. B, Platydoris angustipes. C, Sclerodoris prea. D, Felimare acriba. E–F, Felimare
fregona. G, Felimare kempfi. H, Coryphella dushia. I, Flabellina engeli. J, Flabellina verta. K, Austraeolis catina. L Nanuca
sebastiani. M, Berghia creutzbergi. N, Cuthona iris. O, Tritoniopsis frydis. Sea slugs in their natural environment (Gaby
Carias).
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Distribution. Bahamas, Belize, Brazil, Cayman Islands, Colombia, Costa Rica, French Guiana, Guadeloupe,
Jamaica, Mexico, Puerto Rico, St. Lucia, St. Vincent & the Grenadines, Surinam, Trinidad & Tobago, USA, Virgin
Islands, Venezuela.
Family Scyllaeidae Alder & Hancock, 1855
Genus Scyllaea Linnaeus, 1758
133*. Scyllaea pelagica Linnaeus, 1758
(Figure 5 G)
Material examined. BSM, 2 spcs, H = 50–60 mm, NRC. LJM, 1 spc., H = 61 mm, MCC.
Ecology. Found on benthic Sargassum spp. in shallow water areas with strong currents, however, it is typically
found in seaweed floating offshore.
Distribucion. ABC Islands, Bahamas, Bermuda, Brazil, Costa Rica, Mexico, Sargasso Sea, USA, Venezuela.
Cosmopolitan in temperate warm waters (Valdés et al. 2006).
Family Dotidae Gray, 1853
Genus Doto Oken, 1815
134*. Doto casandra Ortea, 2013
(Figure 5 H)
Material examined. BLR, 1 spc., H = 5 mm, IVICCM000031.
Ecology. On hydrozoans growing on mangrove roots.
Distribution. Bahamas, Costa Rica, Cuba, Curaçao, México, Puerto Rico, Venezuela (Laguna de La Restinga,
Margarita Island: Crescini et al. 2013b; Grune et al. 2015: as D. chica).
Remarks. Ortea (2013) pointed out the similarity between this species and Doto chica, the latter apparently
endemic to Florida and surrounded by taxonomic uncertainty. Venezuela is the southern limit of the distribution of
this species.
Final remarks
The opisthobranchs from the Caribbean province received considerable attention in recent times and its total
diversity was summarized by Valdés et al. (2006) and García & Bertsch (2009), who referred to the occurrence of
308 and 329 species, respectively. To date, the best synthesis of the opisthobranch species known in Venezuela was
due to Valdés et al. (2006), who mentioned the occurrence of 57 species, plus seven determined only to genus level.
In this study we report the occurrence of 134 species, which more than doubles the number of previous known
diversity. Forty-six species are reported for the first time to the country; one of them is a new record for the
Caribbean Sea.
Out of the 134 species recorded in Venezuela, 26 (19.4 %) belong to the Cephalaspidea, 1 (~0.8 %) to the
Runcinacea, 1 (~0.8 %) to the Umbraculida, 14 (~10.5 %) to the Anaspidea, 5 (3.6 %) to the Pleurobranchomorpha,
26 (19.4 %) to the Sacoglossa and 61 (45.5%) to the Nudibranchia. These percentages are similar to those reported
by García & Bertsch (2009) for the entire Caribbean region, with disparities observed in the Cephalaspidea (~9 %
lower in Venezuela) and the Anaspidea (~6 % higher in Venezuela).
The new records for the opisthobranch fauna of Venezuela are: Migaya felis, Chelidonura hirundinina,
Chelidonura cubana, Lapinura divae, Petalifera ramosa, Phyllaplysia engeli, Syphonota geographica,
Stylocheilus longicauda, Oxynoe aguayoi, Bosellia curasoae, Elysia cauze, Elysia tuca, Elysia nisbeti, Elysia
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zuleicae, Checholysia patina, Checholysia annedupontae, Thuridilla malaquita, Cyerce habanensis, Ercolania
fuscata, Costasiella nonatoi, Aegires sublaevis, Aegires cf. ortizi, Bermudella polycerelloides, Jorunna spazzola,
Platydoris angustipes, Sclerodoris prea, Cadlina rumia, Felimida binza, Felimida clenchi, Felimare fregona,
Felimare kempfi, Felimare picta, Flabellina dushia, Austraeolis catina, Godiva rubrolineata, Nanuca sebastiani,
Learchis poica, Spurilla sargassicola, Eubranchus conicla, Cuthona iris, Tritoniopsis frydis, Bornella calcarata,
Scyllaea pelagica, Doto casandra, and Placida cremoniana; the latter also a new record for the Caribbean fauna.
Only one species has Venezuela as its northern limit of distribution, namely Tambja cf. stegosauriformis. Six
species have their type locality in the country: Felimare acriba, Felimare marci, Felimare molloi, Felimare
samueli, Learchis? ignis and Philine buchensis.
Venezuela is the southern limit of the geographical distribution of 49 of the species found in the country (37 %
of the total): Migaya felis, Chelidonura hirundinina, Chelidonura cubana, Cylichna krebsii, Retusa sulcata,
Volvulella minuta, Volvulella texasiana, Haminoea succinea, Atys guildingi, Lapinura divae, Umbraculum
umbraculum, Aplysia morio, Petalifera ramosa, Syphonota geographica, Oxynoe aguayoi, Elysia papillosa, Elysia
nisbeti, Elysia zuleicae, Checholysia patina, Checholysia annedupontae, Thuridilla malaquita, Cyerce habanensis,
Stiliger cricetus, Placida kingstoni, Placida verticilata, Aegires sublaevis, Aegires cf. ortizi, Okenia zoobotryon,
Bermudella polycerelloides, Polycera chilluna, Caribranchus morsomus, Aphelodoris antillensis, Doris kyolis,
Sclerodoris prea, Dendrodoris warta, Felimare grahami, Felimare acriba, Felimare ruthae, Felimare fregona,
Felimare molloi, Flabellina engeli engeli, Austraeolis catina, Dondice parguerensis, Learchis poica, Spurilla
sargasicola, Cuthona iris, Marionia tedi, Tritoniopsis frydis and Doto casandra.
García & Bertsch (2009) found out that ~48 % of the opisthobranch species in the Caribbean are endemic to
the Province, which means that 52 % (at the most) are amphiatlantic (26%), or are shared with the adjacent
Provinces (Brazilian or Carolinian: sensu Spalding et al. 2007; Briggs & Bowen, 2012). On the other hand,
Venezuela harbours alone about 40 % of the entire diversity of the opisthobranchs known in the Caribbean, but ~62
% (84) of these species are also present in Brazil; this is at least 10 % above the calculated maximum (García &
Bertsch, 2009). Its geographical position on the southern edge of the Caribbean, close to the Amazon-Orinoco
barrier, seems to be the most likely explanation for this imbalance of Brazilian fauna; therefore, Venezuela may be
considered a transition area between the Caribbean and Brazil. This is consistent with the conclusions of Diaz
(1995), who studied the biogeography of molluscs in Southern Caribbean and stated that “…the Leeward Islands
(including the islands of Venezuela) should be regarded as a gradual transition area to the Antillean or West Indian
Province, Aruba being the first step”.
Galvão Filho et al. (2015) studied the opisthobranch fauna from Ceara State, in Northern Brazil, on the “other
side” of the Amazon-Orinoco barrier, and listed 57 species, of which 7 % were endemic to the Brazilian Province.
This is the expected proportion according to García & Bertsch (2009). Interestingly, ~90 % of Ceara
opisthobranchs are shared with the Caribbean. Galvão Filho et al. (2015) argued that this situation could be due to
an underestimated diversity, since opisthobranchs from Northern Brazil are still poorly studied. But the available
data (~90 % of common fauna with the Caribbean) suggests that the northern Brazilian coast may also be a
transitional zone between biogeographic Provinces. While a “accumulation effect” may be responsible for the high
diversity observed in Venezuela, this has yet to be confirmed by future findings in Ceara.
Finally, there is a growing body of evidences based on integrative taxonomic studies, pointing out to the
existence of cryptic species complexes among the “opisthobranchs” inhabiting the Western Atlantic (Ornelas-
Gatdula et al. 2011; Ornelas-Gatdula & Valdés 2012; Ornelas-Gatdula et al. 2011; Camacho-García et al. 2014),
this may imply that in the near future some of the identifications given in this paper will have to be revised.
Acknowledgements
Our thanks go to TN Freddy Armada, Commander of the Scientific-Military Base Simon Bolivar in Aves Island,
and to his entire crew. To the Commanders of the ships that transported us to Aves Island and their crews: Oceanic
Surveillance Patrol (PC-23 Yekuana): CF Jesus Martin, and Tugboat (RA 11): CF Antonio Mejia. To Ruben
Portillo, Juan Bolaños, Jesus Enrique Hernandez and Regulo Lopez (UDO) for their help and hospitality during the
fieldtrips to Margarita Island. To Moncho and the rest of the fishermen, for his help and hospitality during the
fieldtrips to La Tortuga Island. To Mr. Pedro Mata and his family for their support with the boat and the rescues in
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Higuerote. To Angel Viloria (IVIC) for his support in the campaign in Venezuela in March 2010. Some of these
results have been obtained thanks to the project IVIC 915 “Biodiversidad marina en Venezuela y su relación con la
dinámica costera. Los moluscos como grupo focal y fuente de nuevas moléculas”. We thank Philippe Bouchet, P.
Maestrati, V. Héros and B. Buge (Biodiversity Exploration Unit, Muséum national d’Histoire naturelle, Paris), for
their kindness and help during the internership as Maître de Conferences of the first author at the MNHN. We are
grateful to Paula Mikkelsen (Paleontological Research Institution, New York) for her help with the identification of
the Cylindrobullidae and to Michael Schödl for his valuable comments and input to improve this manuscript. Two
anonymous referees provided helpful comments on an earlier manuscript version.
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... Aunque la mayoría tiene hábitos bentónicos, hay grupos de vida pelágica. Se distribuyen desde aguas someras hasta grandes profundidades, ocupando hábitats como arrecifes de coral, sustratos rocosos, fondos blandos, praderas de fanerógamas y algas (Caballer et al., 2015;Googheart et al., 2016). Según García & Bertsch (2009), el número de especies de heterobranquios en el Caribe es de 329, siendo este una de las áreas con mayor riqueza de todo el Atlántico. ...
... detalles morfológicos del ejemplar recolectado; c) detalles morfológicos del ejemplar no recolectado co: cola; pa: parapodios; ri: rinóforos. Longitud: a-b) 2,3 cm, c) 1 cm Comentarios: en el Caribe colombiano no hay reportes de este género, del cual solo se conocen cuatro especies registradas en el Atlántico y el mar Caribe: O. antillarum en Brasil (Padula, 2008), Venezuela (Caballer et al., 2015), Curazao y Panamá (Krug et al., 2018), Honduras (Caviedes et (Krug et al., 2018). En estas localidades las especies han sido reportadas en sustratos rocosos y algas del género Caulerpa, siendo esta última su nicho natural. ...
... En este estudio se presenta el primer registro en el Caribe colombiano del género Oxynoe; el espécimen encontrado presenta características morfológicas externas que se asemejan a la especie O. antillarum, la única que cuenta con una distribución cercana a Colombia, en el área de Centroamérica (Panamá) y Suramérica (Venezuela y Brasil) (Padula, 2008;Caballer et al., 2015;Krug et al., 2018). Los miembros de este género presentan una gran plasticidad fenotípica entre individuos de una misma especie, lo cual hace necesarias las descripciones de sus estructuras internas, como la rádula y el aparato reproductor e, incluso, análisis moleculares (Krug et al., 2018). ...
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The subclass Heterobranchia is one of the most diverse among gastropod mollusks; it is present in marine, freshwater, and terrestrial environments and includes interstitial slugs, nudibranchs, intertidal limpets, and land snails and slugs. There is no official list of the diversity of marine or terrestrial heterobranchs for the department of Atlántico. The main objective of the study was to contribute to the first inventory of this gastropod group in this marine-coastal area in the Colombian Caribbean characterized by conditions of high sedimentation, turbidity, and resuspension generated by its proximity to the Magdalena River mouth. The information presented in this study was collected during 38 non-regular field trips in several localities between 2017 and 2021 and complemented by an exhaustive literature and database review. We registered a richness of 17 species distributed in nine Department localities. Heterobranchs were recorded only in 11 of the field trips. The results highlight the first record of the genus Oxynoe for the Colombian Caribbean and the second of the genus Carminodoris in Colombia, of which only one report was known from the department of Bolívar. We registered five species for the first time in this area: three members of the family Aplysiidae and one each of the Discodorididae and Oxynoidae families. This information contributes to the knowledge and understanding of an important gastropod group distributed in this coastal Colombian area.
... At present, eight species of Haminoea are recognized as valid in the Eastern Atlantic (EA) between the southern shores of the British Isles and Angola in West Africa, namely H. hydatis (Linnaeus, 1758; type locality Mediterranean Sea), H. navicula (Da Costa, 1778; type locality Weymouth, Dorset, England), H. orbignyana (Férussac, 1822; type locality near La Rochelle, Bay of Biscay, France), H. elegans (Gray, 1825; type locality south of British Isles and Mediterranean Sea), H. orteai Talavera, Murillo & Templado, 1987 (type locality Salinas del Rasall, Murcia, Spain), H. templadoi García, Pérez-Hurtado & García-Gómez, 1991 (type locality Huelva, Spain), H. exigua (Schaefer, 1992; type locality Adriatic Sea, Italy/ Croatia) and H. fusari (Álvarez, García & Villani, 1993; type locality Lake Fusaro, Italy) (Malaquias & Cervera, 2006;Martínez & Ortea, 1997;Rolán & Ryall, 1999). In the Western Atlantic (WA) four species are often accepted as valid in current literature occurring between Florida, USA and Rio Grande do Sul in Brazil, namely H. elegans, H. antillarum (d'Orbigny, 1841; type locality Saint Thomas, U.S. Virgin Islands), H. petitii (d 'Orbigny, 1841; type locality Cuba) and H. succinea (Conrad, 1846; type locality Tampa Bay, Florida, USA) (Caballer et al., 2015;García et al., 2008;Rios, 2009;Valdés et al., 2006). In the Eastern Pacific (EP) three species are commonly recognized as valid between Alaska and Panama, namely H. ovalis Pease, 1868 (type locality Tahiti, French Polynesia), H. virescens (Sowerby, 1833; type locality Pitcarin Island or California, USA; see Valdés, 2019) and H. vesicula (Gould, 1855; type locality San Diego, California, USA) (Behrens & Hermosillo, 2005;Hermosillo et al., 2006;Valdés & Camacho-Garcia, 2004). ...
... Another case is the species name H. elegans introduced by Gray (1825) based on shells from the British Isles and the Mediterranean Sea, yet, the name is commonly attributed to one of the tropical western Atlantic species (e.g. Caballer et al., 2015;Malaquias, 2014;Marcus, 1976;Marcus & Marcus, 1967;Redfern, 2001;Valdés et al., 2006) and also to spiralled shells occurring in tropical West Africa (Gabon, Republic of the Congo, São Tomé and | 3 TURANI et al. Príncipe, Angola;Bernard, 1984;Martínez & Ortea, 1997;Rolán & Ryall, 1999). ...
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Haminoea are herbivorous, coastal snails occurring in temperate and tropical waters of the Atlantic and Eastern Pacific oceans, with one species present in temperate South Africa (Indian Ocean). The genus is taxonomically difficult as several available nominal species were introduced based on shell descriptions alone, or described based on subtle differences in morpho‐anatomical features, without a phylogenetic molecular framework. Fifteen species are currently accepted as valid in recent scientific literature and field guides (eight Eastern Atlantic, one temperate Indian Ocean, four Western Atlantic and three Eastern Pacific). Here we generate the first complete phylogeny (Bayesian and Maximum Likelihood) of this genus based on multilocus molecular data (COI, 12S rRNA, 16S rRNA, 28S rRNA) using a taxon set accumulated over a period of 15 years, coupled with species delimitation analyses methods (ABGD, ASAP, bPTP) and morpho‐anatomical studies. The goal of this study is to provide insights into the taxonomy, phylogenetic relationships and geographical distributions of species while generating a framework for future systematic reviews of the genus, as well as to study speciation and historical biogeography. Our results rendered four possible hypotheses of species diversity: with 14, 15, 19 and 20 candidate species and point to the fact that several taxa presently regarded as valid might be conspecific (e.g. H. orteai – H. templadoi – H. exigua ; and H. alfredensis – H. antillarum – H. orbignyana ), while highlighting the existence of a complex of four or five species often identified as H. elegans . Pervasive nomenclatural problems in the genus, including with the type species H. hydatis , are highlighted and discussed.
... These authors proposed the genus Caribranchus Ortea et al. (2003) to accommodate H. morsomus. Nevertheless, with rare exceptions (Debelius & Kuiter, 2007;Gutiérrez et al., 2015 Ortea & Buske, 2018;Ortea et al., 2012), this name has been ignored in the literature. regarded the genus Caribranchus as a synonym of Hexabranchus due to the lack of phylogenetic evidence. ...
... Distribution Caribbean: British Virgin Islands(Marcus & Marcus, ), Puerto Rico(Marcus & Marcus, 1968), Guadeloupe(Ortea et al., 2012), Venezuela(Gutiérrez et al., 2015), Panama(Collin et al., 2005), Costa Rica, Honduras, Aruba, Puerto Rico, St. Marteen/St Martin, St. Lucia, Martinique, Antigua, Grenada, St. Vincent and the Grenadines, Trinidad & Tobago (Valdés et al., 2006). Material examined One specimen. ...
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Color ontogeny and variations associated with discrete morphological differences may generate taxonomical challenges, which requires multiple data types and in-depth historical review. The nudibranch known as the Spanish dancer, Hexabranchus sanguineus , is a classic example with over 200 years of taxonomic confusion. Currently, H. sanguineus is accepted by most authors as a single species from the Indo-Pacific Ocean with Hexabranchus morsomus as a valid species from the Atlantic Ocean. Yet, despite these species being highly studied, their systematic status remains debatable. Over 30 synonyms have been proposed for H. sanguineus and even a distinct genus for H. morsomus . Here we provide, for the first time, a comprehensive review of all proposed names and an integrative taxonomic revision of the genus including morphological and molecular data. Our results reveal that H. sanguineus is a complex of five species: four previously described and an undescribed species, one of the largest nudibranchs in the world: Hexabranchus giganteus sp. nov. The genus Caribranchus is considered a junior synonym of Hexabranchus Ehrenberg, 1828 and the ontogeny of color pattern is discussed.
... In fact, the Peruvian sea is recognized as one of the impoverished regions worldwide in terms of nudibranch diversity (Schrödl 1997(Schrödl , 2002(Schrödl , 2003Schrödl and Hooker 2014). In comparison, other regions such as the Caribbean Sea, the Tropical Eastern Pacific, the Indian Ocean, the Mediterranean Sea (Sachidhanandam et al. 2000;Valdés et al. 2006;Chavanich et al. 2013;Ah Shee Tee et al. 2019;Furfaro et al. 2020;Londoño-Cruz 2021), as well as other South American countries including Brazil, Chile, and Venezuela (Fischer and Cervera 2005b;Ardila et al. 2007;Aldea et al. 2011;Padula et al. 2011;Alvim and Pimenta 2013;Gutiérrez et al. 2015;Araya and Valdés 2016; Londoño-Cruz 2021) exhibit considerably higher richness of nudibranch species. ...
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Nudibranchs, as a group, have received limited attention in terms of scientific study along the coastline of Peru. Here, an updated and comprehensive list of nudibranch species found in the Peruvian sea is presented, compiled through an extensive review of relevant literature. This compilation encompasses a total of 31 species, classified into two suborders, 10 superfamilies, 20 families, and 28 genera. With respect to the biogeographic provinces along the Peruvian coast, 23 species inhabit the Warm Temperate Southeastern Pacific province, 18 species occur in the Tropical Eastern Pacific province, and 10 species are found in both provinces, crossing the transitional zone between them. In terms of distribution patterns, two species exhibit a cosmopolitan distribution ( Glaucus atlanticus and Fiona pinnata ), while two species display a circumtropical distribution ( Cephalopyge trematoides and Phylliroe bucephala ). One species exhibits a bipolar distribution in the Eastern Pacific and possesses an amphi-South American distribution ( Rostanga pulchra ). Additionally, six species exhibit an amphi-South American distribution ( Rostanga pulchra , Diaulula punctuolata , Doto uva , Tyrinna evelinae , Tyrinna delicata , and Doris fontainii ), and two species are endemic to Peru ( Corambe mancorensis and Felimare sechurana ). This study provides comprehensive information on biogeographical aspects, geographical distributions, and taxonomic updates within the nudibranch species documented in Peru. Furthermore, we discuss the status of species listed in previous literature that have not been confirmed by collections, referring to them as potentially occurring species.
... In fact, the Peruvian sea is considered one of the poorest regions in the world in terms of nudibranch species (Schrödl 1997(Schrödl , 2002(Schrödl , 2003Schrödl and Hooker 2014). Comparatively, other regions such as the Caribbean Sea, the Tropical Eastern Pacific, the Indian Ocean, the Mediterranean Sea (Sachidhanandam et al. 2000;Valdés 2006;Chavanich et al. 2013;Ah Shee Tee et al. 2019;Furfaro et al. 2020;Londoño-Cruz 2021) and other South American countries such as Brazil, Chile, and Venezuela (Fischer and Cervera 2005b;Ardila et al. 2007;Aldea et al. 2011;Padula et al. 2011;Alvim and Pimenta 2013;Gutiérrez et al. 2015;Araya and Valdés 2016; Londoño-Cruz 2021) exhibit a greater richness of nudibranchs. ...