The asteroid species of Lamarck (Echinodermata: Asteroidea)

Directeur De la publication / Publication director : Bruno David Président du Muséum national d’Histoire naturelle réDactrice en chef / editor-in-chief : Laure Desutter-Grandcolas assistante De réDaction / assistant editor : Anne Mabille ([email protected]) Mise en page / Page layout : Anne Mabille coMité scientifique / scientific board : Nesrine Akkari (Naturhistorisches Museum, Vienne, Autriche) Maria Marta Cigliano (Museo de La Plata, La Plata, Argentine) Serge Gofas (Universidad de Málaga, Málaga, Espagne) Sylvain Hugel (CNRS, Université de Strasbourg, France) Marco Isaia (Università degli Studi di Torino, Turin, Italie) Rafael Marquez (CSIC, Madrid, Espagne) Jose Christopher E. Mendoza (Lee Kong Chian Natural History Museum, Singapour) Annemarie Ohler (MNHN, Paris, France) Jean-Yves Rasplus (INRA, Montferrier-sur-Lez, France) Wanda M. Weiner (Polish Academy of Sciences, Cracovie, Pologne) couverture / cover : Réalisée à partir de différentes planches de Lesueur reproduites dans cet article (avec l’aimable autorisation du Muséum d’Histoire naturelle du Havre)/Made from various plates by Lesueur reproduced in this article (with the kind permission of the Natural History Museum of Le Havre). 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Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / https://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2021 ISSN (imprimé / print) : 1280-9551/ ISSN (électronique / electronic) : 1638-9387 213 ZOOSYSTEMA • 2021 • 43 (13) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com urn:lsid:zoobank.org:pub:47EAA3BE-82DB-4363-BA57-43F4CF508AC6 Jangoux M. 2021. — The asteroid species of Lamarck (Echinodermata: Asteroidea). Zoosystema 43 (13): 213-252. https://doi.org/10.5252/zoosystema2021v43a13. http://zoosystema.com/43/13 ABSTRACT Zoologist Péron and artist Lesueur, both members of the scientific staff of the Baudin expedition to the Southern Lands (September 1800-March 1804), collected during their voyage 36 different species of asteroids. This is what wrote Lamarck in a report made in June 1804. This number was clearly reduced by Lamarck himself who, in his 1816 publication, listed only 15 species from the South Seas in the Paris Museum collection. However, the different asteroids collected during the expedition were drawn by Lesueur (water colours and pencil drawings) who thus realised a real pictorial register. Lesueur’s drawings are housed in the Le Havre Museum. Due to their realism and precision, the drawings make it easy to identify the species. Confrontation of Lamarck’s report (1804) and publication (1816) with Lesueur’s drawings (done between 1802 and 1804) gives a new, more precise idea of the importance of the collection of South Seas asteroids brought back to France and allows to reliably count the number of new species that it contained. Also, this makes it possible to complete the often too brief descriptions of some Lamarckian species and to clarify their status. Eleven taxonomic changes are thus proposed here: Asterias calcar var. quinqueangula is synonymized with Parvulastra exigua (Lamarck, 1816), Asterias calcitrapa var. 1 with Bollonaster pectinatus (Sladen, 1883), Asterias calcitrapa var. 2 with Astropecten vappa Müller& Troschel, 1843, Asterias nodosa var. 3 with Protoreaster lincki (Blainville, 1830), Asterias pentagonula with Tosia australis Gray, 1840, Asterias pleyadella with Protoreaster sp., Asterias punctata with Asteropsis carinifera (Lamarck, 1816), Asterias rosacea var. lobis senis with Anseropoda sp., Asteriscus setaceus with Paranepanthia grandis (H.L. Clark, 1928), Astrogonium lamarckii Müller& Troschel, 1842 with Goniaster tessellatus (Lamarck, 1816) and Asterias cuspidata is moved to the genus Mediaster Stimpson, 1857 as Mediaster cuspidatus (Lamarck, 1816) n.comb. RÉSUMÉ Les espèces d’astéries de Lamarck (Echinodermata: Asteroidea). Le zoologiste Péron et le dessinateur Lesueur, membres de l’équipe scientifique de l’expédition Baudin aux Terres australes (septembre 1800-mars 1804), y ont récolté, selon un rapport de Lamarck daté de juin 1804, 36 espèces différentes d’astéries. Ce nombre fut nettement réduit, en 1816, par Lamarck lui-même qui ne répertoria que 15 espèces d’origine australe dans les collections du muséum de Paris. Or, les différentes astéries récoltées pendant l’expédition furent dessinées par Lesueur (aquarelles et crayonnés) qui en réalisa un véritable registre pictural. Conservés au muséum du Havre, les dessins de Lesueur, par leur réalisme et leur précision, permettent aisément d’identifier les espèces. La confrontation du rapport de Lamarck (1804) et de sa publication (1816) avec les dessins de Lesueur (réalisés entre 1802 et 1804) donne une image précise et nouvelle de l’importance de la collection d’astéries ramenée à l’époque en France et permet de déterminer Michel JANGOUX Laboratoire de Biologie marine, Université Libre de Bruxelles, 50 avenue F. D. Roosevelt, B-1050 Bruxelles (Belgium) and Laboratoire de Biologie marine, Université de Mons, 20 Place du Parc, B-7000 Mons (Belgium) Submitted on 24 February 2020 | Accepted on 11 May 2020 | Published on 11 May 2021 The asteroid species of Lamarck (Echinodermata: Asteroidea) KEY WORDS Lesueur, Asteroidea, Echinodermeta, sea stars, Australia, Tasmania, Baudin expedition, new synonymies, new combination. 214 ZOOSYSTEMA • 2021 • 43 (13) Jangoux M. fiablement le nombre de nouvelles espèces australes recueillies alors. Une confrontation qui a également permis de compléter les descriptions, souvent très brèves, de certaines des espèces d’astéries établies par Lamarck et d’éclaircir leur statut. Onze changements taxonomiques sont ainsi proposés ici: Asterias calcar var. quinqueangula est synonyme de Parvulastra exigua (Lamarck, 1816), Asterias calcitrapa var. 1 de Bollonaster pectinatus (Sladen, 1883), Asterias calcitrapa var. 2 de Astropecten vappa Müller & Troschel, 1843, Asterias nodosa var. 3 de Protoreaster lincki (Blainville, 1830), Asterias pentagonula de Tosia australis Gray, 1840, Asterias pleyadella de Protoreaster sp., Asterias punctata de Asteropsis carinifera (Lamarck, 1816), Asterias rosacea var. lobis senis de Anseropoda sp., Asteriscus setaceus de Paranepanthia grandis (H.L. Clark, 1928), Astrogonium lamarckii Müller& Troschel, 1842 de Goniaster tessellatus (Lamarck, 1816) et Asterias cuspidata est transférée dans le genre Mediaster Stimpson, 1857 en tant que Mediaster cuspidatus (Lamarck, 1816) n. comb. INTRODUCTION The part devoted to asteroids in the second volume of Lamarck’s Histoire naturelle des Animaux sans Vertèbres (1816) reports on 44species and 23varieties. Some of the new species Lamarck then described originated from European seas and the Atlantic and Indian coasts of Africa though most came from Australian coasts. Indeed, the Paris Museum had received the very large collection of sea stars made during the Voyage de Découvertes aux Terres australes, the so-called Baudin expedition, in the early nineteen century (1800-1804). The main collectors of this collection were the zoologist François Péron and his friend the artist Charles-Alexandre Lesueur, both members of the expedition scientific staff sailing on the vessel Le Géographe. The collections of the Baudin expedition reached the Muséum national d’Histoire naturelle (MNHN) at the end of April 1804. Two months later Lamarck presented several reports on invertebrates to his colleagues at the Muséum; one of which dealing with echinoderms (Jangoux in press). The collection of asteroid contains 264individuals representing, Lamarck wrote, 36different species of Asterias, many of which were, he said, new. What he did not know at the time was that the asteroids collected were drawn (watercolours and pencil drawings) by Lesueur, the expedition artist. Fortunately, Lesueur’s drawings have been preserved and kept in good condition, being housed since the last quarter of the 19th century in the Muséum d’Histoire naturelle du Havre (MHNH) (the complicated story of these drawings and their progressive acquisition by the MHNH is detailed in Bonnemains [1995]). There are 53groups of asteroids drawings in the Lesueur’s collection. They are very realistic and most of them allow identification of species. Moreover, they are so precise that some show the arrangement of skeletal plates and details on their spine armament. Based on these drawings, the number of species collected was assessed at about 50 (Jangoux 1984), more than the 36species reported by Lamarck in 1804, and still much more than the 15species collected by Péron and Lesueur that Lamarck (1816) said he found in the MNHN. The number of species collected during Baudin’s voyage differs distinctly according to the sources (50, 36or 15) without knowing the exact number of new species collected by Péron and Lesueur, except that it should be more than 15! Lesueur’s drawings of asteroids only concern the individuals of the Baudin expedition. He thus made a pictorial register representing almost all the asteroids that reached the Muséum in April 1804. Thus, two groups of new species from the South Seas should be considered: those described by Lamarck in 1816, and those not reported by Lamarck in 1816, although the specimens have been seen, counted and sorted by him when the collected material reached the Muséum in 1804. The Baudin expedition is well documented and, while Lamarck rarely cites the collecting localities (in most cases he writes “originated from the South Seas” or even more allusively “from the voyage of Péron and Lesueur”) they can be found in various archives. Collections were made in West Australia (Geographe Bay, Swan River, Barren Islands [Bernier and Dorre Islands], Shark Bay, Depuch Island, King George Sound [Albany]), Tasmania (d’Entrecasteaux Channel, Maria Island, King Island), New South Wales (Port Jackson), South Australia (Kangaroo Island, Saint Peter and Franklin Islands) (Table1). Other non-Australian collecting localities were Timor (Kupang Bay), Mauritius, and the Cape of Good Hope. The taxonomic status of every Lamarckian species, whatever its origin, is presented or briefly discussed hereafter. As for the species collected during the Baudin expedition, their drawings are reproduced (Lesueur’s watercolours). Among them are those that, although sorted by Lamarck in 1804 and drawn by Lesueur, were not reported in Lamarck’s (1816) treatise. The latter were described by subsequent authors and are presented under their original names. The species are presented according to the alphabetical order of the specific names used in the original descriptions. Information provided consists of: the species name, the original bibliographic reference, the species current status (possibly, revised), the examined material, the species distribution, any additional relevant information (under the heading ‘Remark’). MATERIAL AND METHODS Two lists of asteroids have been drawn up. The first consists in all the new species established by Lamarck in his 1816 publication, whatever their geographical origin. The second brings together the species collected during the Baudin expedition (1800-1804), mainly in Australia and Tasmania. These were all drawn by Lesueur, and deposited in the Muséum where Lamarck received them in June 1804. The identifications were made from photographs of Lesueur’s original drawings. MOTS CLÉS Lesueur, Asteroidea, Echinodermeta, étoiles de mer, Australie, Tasmanie, expédition Baudin, synonymies nouvelles, combinaison nouvelle. 215 The asteroid of Lamarck ZOOSYSTEMA • 2021 • 43 (13) Whether or not the species are still present in the MNHN, the two lists make it possible to reconstruct the Parisian asteroid collection and its singularity at the beginning of the 19th century. AbbreviAtions MHNH Muséum d’Histoire naturelle du Havre, Le Havre; MNHN Muséum national d’Histoire naturelle, Paris. LIST OF ASTEROID SPECIES africanus Müller& Troschel, 1842, Asteracanthion (Fig. 1) Asteracanthion africanus Müller& Troschel, 1842: 15. Marthasterias africana – Wright etal. 2016: 446. Current stAtus.— Marthasterias africana (Müller& Troschel, 1842). M AteriAl exAMined .— MHNH; Lesueur drawings collection; ref.74026; Baudin expedition; Péron and Lesueur leg. distribution.— Southernmost part of Africa. reMArk Now recognised as a valid species, M.africana, since its description in 1842, was considered a variety or form of the common European asteroid Marthasterias glacialis (Linnaeus, 1758) (e.g. Mortensen 1933; Clark& Courtman-Stock 1976). Péron and Lesueur collected individuals of this common species when staying in the Cape of Good Hope area on their way back to France. Péron and Lesueur’s material of A.africanus could not be found in the MNHN. angulatus Müller& Troschel, 1842, Archaster (Fig. 2) Archaster angulatus Müller& Troschel, 1842: 66. — Sukarno& Jangoux 1977: 830. — Clark 1993: 238. Current stAtus.— Archaster angulatus Müller& Troschel, 1842. M AteriAl exAMined .— MHNH; Lesueur drawings collection; ref. 74044 and 74060; Baudin expedition; Péron and Lesueur legs. d istribution .— The species commonly occurs along the west and north coasts of Australia; it is also reported from South China Sea and the Philippines. reMArk Péron and Lesueur’s material of A.angulatus could not be found in the MNHN. australiae Döderlein, 1915, Anthenea (Fig. 3) Anthenea australiae Döderlein, 1915: 52. Current stAtus.— Anthenea australiae Döderlein, 1915. MAteriAl exAMined.— MNHN-IE-2014-542; 1specimen; Baudin expedition (1800-1804); Péron and Lesueur leg. [formerly identified as syntype of Asterias pentagonula] • MNHN-IE-2014-543; 1speci - men; same origin as precedent [formerly identified as syntype of Asterias pentagonula] • MHNH; Lesueur drawings collection; ref. 74052; Baudin expedition. distribution.— West and north-west Australian coasts. reMArk The MNHN houses two individuals of Döderlein’s Anthenea australiae under the name Anthenea pentagonula (Lamarck, 1816), a misidentification resulting from an earlier confusion about the status of Lamarck’s Asterias pentagonula (further explanation under that species). calcar var. octogona Lamarck, 1816, Asterias (Fig. 4) Asterias calcar var. octogona Lamarck, 1816: 557. Asteriscus australis (part) Müller& Troschel, 1842: 43. Meridiastra calcar – Clark 1993: 224. — O’Loughlin & Waters 2004: 3. Patiriella calcar – Clark in Clark & Mah 2001: 252. Current stAtus.— Meridiastra calcar (Lamarck, 1816). MAteriAl exAMined.— Australia • MNHN-IE-2014-248; holotype; King George Sound, Albany; Baudin expedition (1800-1804); Péron and Lesueur leg. • MHNH; Lesueur drawings collection ref. 74039; Baudin expedition. distribution.— South and east coasts of Australia, Tasmania. reMArks Three varieties of Asterias calcar were briefly described by Lamarck –viz. hexagona, octogona and quinqueangula– whose individuals were collected in Australian waters (King George Sound [Albany], West Australia) and are still in the MNHN. Table 1. — Baudin expedition: releases and moorings. Areas of collecting Periods of collecting Geographe Bay 31.V-9.VI.1801 Shark Bay, Barren Islands (1st stay) 29.VI-11.VII.1801 Depuch Island 17-18.VII.1801 Kupang Bay, Timor (1st stay) 22. VIII -13.XI.1801 D’Entrecasteaux Channel, Tasmania 20.I-17.II.1802 Maria Island, Tasmania 17-22.II.1802 Port Jackson (Sydney) 17.VI-18.XI.1802 King Island, Tasmania 7-13.XII.1802 Kangaroo Island 6.I-1.II.1803 Saint Peter and Franklin Islands 7-11.II.1803 King George Sound 18.II-1.III.1803 Shark Bay (2nd stay) 19-23.III.1803 Kupang Bay, Timor (2nd stay) 6.V-3.VI.1803 Mauritius 7.VIII-16.X.1803 Cape of Good Hope 5-26.I.1804 216 ZOOSYSTEMA • 2021 • 43 (13) Jangoux M. Lamarck reported their colour in life from a Lesueur drawing and without distinguishing the varieties (“Cette astérie est rouge, violette, brillante de couleur, et ressemble à une fleur lorsqu’elle est vivante [this sea star is red, violet, of brilliant colours and looks like a flower when alive]”). Lesueur’s drawings indicate these colours should at least characterize individuals of the variety octogona. The type specimen of Asterias calcar var. octogona was not recognised by Müller& Troschel (1842) who made it one of the types of their new species, Asteriscus australis. calcar var. hexagona Lamarck, 1816, Asterias (Fig. 5) Asterias calcar var. hexagona Lamarck, 1816: 557. Asteriscus australis (part.) Müller& Troschel, 1842: 43. Meridiastra gunni – Clark 1993: 225. — O’Loughlin & Waters 2004: 3. Current stAtus.— Meridiastra gunni (Gray, 1840). Fig. 1. — Asteracanthion africanus Müller & Troschel, 1842: A, B, abactinal (A) and actinal (B) surfaces; C, D, arm cross sections with gonad structure (D); E, arm actinal surface showing adambulacral spines; F, abactinal spines surrounded by rings of crossed pedicellariae (Lesueur collection 74026, details, courtesy of MHNH). A CD B EF 217 The asteroid of Lamarck ZOOSYSTEMA • 2021 • 43 (13) MAteriAl exAMined.— MNHN-IE-2014-508; holotype; King George Sound, Albany; Baudin expedition (1800-1804); Péron and Lesueur leg. • MHNH; Lesueur drawings collection; ref. 74003; Baudin expedition. distribution.— SW to SE Australia, Tasmania. reMArk The type specimen of Asterias calcar var. hexagona was not recognised by Müller& Troschel (1842) who made it one of the types of their new species, Asteriscus australis. calcar var. quinqueangula Lamarck, 1816, Asterias (Fig. 15C) Asterias calcar var. quinqueangula Lamarck, 1816: 557. Patiriella regularis – Clark 1993: 226. Current stAtus.— Asterias calcar var. quinqueangula is newly synonymized here with Parvulastra exigua (Lamarck, 1816). MAteriAl exAMined.— Australia • MNHN-IE-2014-30: Asterias calcar var. quinqueangula; 4syntypes; Australia (King George Sound, Albany); Baudin expedition (1800-1804); Péron and Lesueur leg. reMArk Since the publication of Perrier (1875: 299), Lamarck quinqueangula variety was considered a synonym of the common New Zealand asterinid, Patiriella regularis (Verrill, 1867). Yet this should be reconsidered. Indeed, although the presence of P.regularis in Australian waters, mostly Tasmanian, is proven, this presence is however recent. In fact P.regularis, a species endemic to New Zealand, is said to have invaded Australian waters in the 20th century after 1930 (Dartnall 1969). This makes it rather unlikely that it was collected by Péron and Lesueur in South Australia or Tasmania in the early 19th century. The status of quinqueangula variety is discussed further under the species Asterias exigua. Fig. 2 . — Archaster angulatus Müller & Troschel, 1842: A, B, abactinal surface of two individuals; C, arrangement and details of abactinal paxillae; D, arm cross-section showing the arrangement of arm plates; E, adambulacral armament and inferomarginal plates (Lesueur collection 74044 and 74060 details, courtesy of MHNH). AB CD E 218 ZOOSYSTEMA • 2021 • 43 (13) Jangoux M. Fig. 3. — Anthenea australiae Döderlein, 1915: A, B, abactinal and actinal surfaces; C, interradial arc and adambulacral spines and plates; D, madreporite and abactinal plates; E, specimen of A. australiae from the Baudin expedition [syntype of Asterias pentagonula] (A-D, Lesueur collection 74052, details, courtesy of MHNH; E, MNHN-IE-2014-543). Scale bar: D, 50 mm. A B C E D 219 The asteroid of Lamarck ZOOSYSTEMA • 2021 • 43 (13) Fig. 4. — Asterias calcar var. octogona Lamarck, 1816: A, B, abactinal (A) and actinal (B) surfaces; C, profile view; D, armament of abactinal plates; E, actinolateral plates and adambulacral spines (text: “dessous” [underside]); F, holotype of Meridiastra calcar (Lamarck, 1816) (A-E, Lesueur collection ref. 74039, details, courtesy of MHNH; F, MNHN-IE-2014-248). Scale bar: F, 20 mm. AB C D F E 226 ZOOSYSTEMA • 2021 • 43 (13) Jangoux M. Fig. 11. — Asterias cylindrica Lamarck, 1816: A, B, abactinal (A) and actinal (B) surfaces; C, adambulacral armament; D, holotype of the species (A-C, MHNH, Lesueur collection 74014, details, courtesy of MHNH; D, MNHN-IE-2014-471). The accompanying labels do not relate to the pictured specimen. The holotype of O. purpureus Perrier, 1869 is shown in Figure 32. Scale bar: 50 mm. AB C D 227 The asteroid of Lamarck ZOOSYSTEMA • 2021 • 43 (13) Fig. 12. — Echinaster decanus Müller & Troschel, 1843: A, B, abactinal (A) and actinal (B) surfaces (MHNH, Lesueur collection 74053, details, courtesy of MHNH). Fig. 13. — Asterias discoidea Lamarck, 1816: A, B, abactinal (A) and actinal (B) surfaces; C, adambulacral spines (above), interradial arcs (below) (MHNH, Lesueur collection 74006, details, courtesy of MHNH). Fig. 14. — Asterias echinophora Lamarck, 1816: abactinal and actinal surfaces of a syntype of the species (MNHN-IE-2014-72). Scale bar: 20 mm. AB AB C 228 ZOOSYSTEMA • 2021 • 43 (13) Jangoux M. exigua Lamarck, 1816, Asterias (Fig. 15) Asterias exigua Lamarck, 1816:554. Asterias calcar var. quinqueangula Lamarck, 1816: 557. Parvulastra exigua – O’Loughlin & Waters 2004: 225. Current stAtus.— Parvulastra exigua (Lamarck, 1816). M AteriAl exAMined .— South Africa • MNHN-IE-2014-526; Asterias exigua; 6syntypes; Cape of Good, Hope Baudin expedition (1800-1804); Péron and Lesueur leg. Australia • MNHN-IE-2014-30; Asterias calcar var. quinqueangula; 4 syntypes; Baudin expedition (1800-1804); King George Sound, Albany, Péron and Lesueur leg.• MHNH; Lesueur drawings collection, ref. 74004; Baudin expedition. distribution.— From the southernmost part of Africa to the south of Australia and Tasmania. Fig. 15. — Asterias exigua Lamarck, 1816: A, drawings of Bruguière (1791); B, type specimens of Asterias exigua. C, type specimens of Asterias calcar var. quinqueangula, D-F, Lesueur’s drawings of Asterias exigua: abactinal (D) and actinal (E) surfaces, and profile view (F) (B, MNHN-IE-2014-526; C, MNHN-IE-2014-30; D-F, MHNH, Lesueur collection 74004, details, courtesy of MHNH). A BC DE F 229 The asteroid of Lamarck ZOOSYSTEMA • 2021 • 43 (13) reMArks According to Lamarck (1816) Asterias exigua came from American seas and was the smallest known species of asteroid. As far as size is concerned, he is almost correct. Yet A.exigua does not occur in the Atlantic but in the south of Africa and Australia as well as in Tasmania. Indeed the type specimens of A.exigua were collected in South Africa by Péron and Lesueur in 1804 (Fig.15B). Individuals of a most likely similar species had been obtained early in 1803 by the same collectors in SE Australia (King George Sound, Albany). The latter were identified Asterina calcar var. quinqueangula by Lamarck, who added on a label that they should be considered juvenile individuals (“specimen junius”) (Fig.15C-F). Comparing the type specimens of the two Lamarckian species (and to the drawings of Bruguière [1791] to which Lamarck refered, Figure 15A), it is highly likely they are conspecific. Due to incorrect information (types of Asterias exigua presumed loss), Dartnall (1971) designated a neotype for the species. The latter originated from False Bay (South Africa) and is in the Tasmanian Museum, Hobart (ref. TM H508). granifera Lamarck, 1816, Asterias (Fig. 16) Asterias granifera Lamarck, 1816: 560. Fig. 16. — Asterias granifera Lamarck, 1816: A, B, abactinal (A) and actinal (B) surfaces; C, arm showing tube feet and adambulacral spines; D, holotype of the species (A-C, MHNH, Lesueur collection 74046 and 74047, details, courtesy of MHNH; D, MNHN-IE-2014-235). AB D C 230 ZOOSYSTEMA • 2021 • 43 (13) Jangoux M. Uniophora granifera – Clark in Clark & Mah 2001: 305. Current stAtus.— Uniophora granifera (Lamarck, 1816). M AteriAl exAMined .— South Seas • MNHN-IE-2014-235; holotype; Baudin expedition (1800-1804); Péron and Lesueur leg. • MHNH; Lesueur drawings collection; ref. 74046 and 74047; Baudin expedition. distribution.— South coast of Australia, Tasmania. helianthus Lamarck, 1816, Asterias (Fig. 17) Asterias helianthus Lamarck, 1816: 558. Heliaster helianthus – Clark in Clark& Mah 2001: 307. Current stAtus.— Heliaster helianthus (Lamarck, 1816). reMArk Lamarck only refered to Bruguière’s (1791: pl.108-109) drawings when describing the species. He gave no localities though one could expect the drawn individual came from Chilean coast. Bruguière’s material could not be found in the MNHN. hesperus Müller& Troschel, 1840, Archaster (Fig. 18) Archaster hesperus Müller& Troschel, 1840: 104. Craspidaster hesperus – Clark 1989: 274. C urrent stAtus .— Craspidaster hesperus (Müller& Troschel, 1840). AB Fig. 17. — Heliaster helianthus Lamarck, 1816: Bruguière’s (1791) drawings of the abactinal (A) and actinal (B) surfaces of the holotype. Fig. 18. — Archaster hesperus Müller & Troschel, 1840: A, B, abactinal (A) and actinal (B) surfaces (MHNH, Lesueur collection 74010, details, courtesy of MHNH). AB 231 The asteroid of Lamarck ZOOSYSTEMA • 2021 • 43 (13) MAteriAl exAMined.— Timor • MNHN-IE-2017-560; 2individuals; Baudin expedition (1800-1804); supposedly from Timor; Péron and Lesueur leg. • MHNH: Lesueur drawings collection, ref. 74010; Baudin expedition. d istribution .— Indonasia, Bay of Bengal, Singapore, Philippines, South China Sea. magnificum Müller& Troschel, 1842, Astrogonium (Fig. 19) Astrogonium magnificum Müller& Troschel, 1842: 53. Tosia magnifica – Clark 1993: 289. Current stAtus.— Tosia magnifica (Müller& Troschel, 1842). MAteriAl exAMined.— MHNH; Lesueur drawings collection; ref. 74056; Baudin expedition, Péron and Lesueur leg. distribution.— South of Australia and Tasmania. reMArk Péron and Lesueur’s material could not be found in the MNHN. milleporella Lamarck, 1816, Asterias (Fig. 20) Asterias milleporella Lamarck, 1816: 564. Fromia milleporella – Clark 1993: 332. Current stAtus.— Fromia milleporella (Lamarck, 1816). MAteriAl exAMined.— South Seas • MNHN-IE-2014-57, MNHNIE-2013-4376: 2 syntypes, Baudin expedition (1800-1804); Péron and Lesueur leg. • MHNH; Lesueur drawings collection; ref. 74016; Baudin expedition (1800-1804). distribution.— The species commonly occurs in the tropical Indo-West Pacific. It is not present in European seas contrary to Lamarck’s assumption. Fig. 19. — Astrogonium magnificum Müller & Troschel, 1842: A, B, abactinal (A) and actinal (B) surfaces; C, profile view; D, superomarginal and abactinal plates; E, madreporite and disc abactinal plates (MHNH, Lesueur collection 74056, details, courtesy of MHNH). AB C D E 232 ZOOSYSTEMA • 2021 • 43 (13) Jangoux M. Fig. 20. — Asterias milleporella Lamarck, 1816: A, B, abactinal and actinal surfaces; C, furrow spines; D, madreporite and disc abactinal plates; E, syntype of the species (A-D, MHNH, Lesueur collection 74016, details, courtesy of MHNH; E, MNHN-IE-2014-57). Scale bar: 20 mm. Fig. 21. — Asterias multifora Lamarck, 1816: A, abactinal surface; B, arm cross-section showing the arrangement of skeletal plates; C, adambulacral and actinolateral granules and plates (MHNH, Lesueur collection 74019, details, courtesy of MHNH). AB C D E ABC 233 The asteroid of Lamarck ZOOSYSTEMA • 2021 • 43 (13) multifora Lamarck, 1816, Asterias (Fig. 21) Asterias multifora Lamarck, 1816: 565. Linckia multifora – Clark 1993: 339. Current stAtus.— Linckia multifora (Lamarck, 1816). MAteriAl exAMined.— MHNH; Lesueur drawings collection; ref. 74019; Baudin expedition; Péron and Lesueur leg. d istribution .— The species commonly occurs in the whole Indo-West Pacific. It does not occur in European seas contrary to Lamarck’s assumption. reMArk Lamarck’s type material could not be found in MNHN. muricata Verrill, 1867, Coscinasterias (Fig. 22) Coscinasterias muricata Verrill, 1867: 249. — Clark in Clark & Mah 2001: 270. Current stAtus.— Coscinasterias muricata Verrill, 1867. MAteriAl exAMined.— MHNH; Lesueur drawings collection; ref. 74036; Baudin expedition (1800-1804); Péron and Lesueur leg. distribution.— Temperate Australia, Tasmania, New Zealand. reMArk Péron and Lesueur’s material could not be found in the MNHN. Fig. 22. — Coscinaterias muricata Verrill, 1867: A, B, abactinal (A) and actinal (B) surfaces; C, arm cross-section showing enlarged abactinal spines surronded by rings of crossed pedicellariae, below: upper view of such a spine and ring; C, actinal side of an arm section showing tube feet and adambulacral and inferomarginal spines (MHNH, Lesueur collection 74036, details, courtesy of MHNH). AB CD 234 ZOOSYSTEMA • 2021 • 43 (13) Jangoux M. nodosa Lamarck, 1816, Asterias (Fig. 22) Asterias nodosa Lamarck, 1816: 557. reMArk Considering the MNHN collection and the works of Linck (1733) and Bruguière (1791), Lamarck cited three varieties of Asterias nodosa which in fact correspond to two different species. nodosa var. 1 & 2 Lamarck, 1816, Asterias (Fig. 23) Asterias nodosa Lamarck, 1816: 557. Asterias nodosa Linnaeus, 1758: 661. Protoreaster nodosus – Clark 1993: 318. Current stAtus. — Protoreaster nodosus (Linnaeus, 1758). M AteriAl exAMined .— MHNH; Lesueur drawings collection; ref.74055; Baudin expedition (1800-1804) ; Péron and Lesueur leg. distribution.— Tropical East Indian ocean and tropical West Pacific. nodosa var. 3 Lamarck, 1816, Asterias (Fig. 23E-F) Asterias nodosa Lamarck, 1816:557 r evised stAtus . — Asterias nodosa var. 3 is newly synonymized here with Protoreaster lincki (Blainville, 1830). MAteriAl exAMined. — Mauritius • MNHN-IE-2017-837; specimen from the Baudin expedition (1800-1804) ; Péron and Lesueur leg. distribution. — Tropical West Indian ocean. reMArk P. lincki differs markedly from P. nodosus in having some of the distal supero-marginal plates bearing laterally-projecting spines. Also red and brown-beige are the dominant colours of abactinal and supero-marginal plates in P. lincki while they are of a varied brown in P. nodosus. Note also that P. lincki is very common in the west part of the Indian Ocean while P. nodosus is in the West Pacific. obtusangula Lamarck, 1816, Asterias (Fig. 24) Asterias obtusangula Lamarck, 1816: 556. Fig. 23. — Asterias nodosa Lamarck, 1816, A-D, varieties 1 & 2: A, B, abactinal (A) and actinal (B) surfaces; C, enlarged abactinal tubercule, D, interradial arc (MHNH, Lesueur’s collection 74055, details, courtesy of MHNH); E, F, variety 3: abactinal (E) and actinal (F) surfaces (MNHN-IE-2017-837). D E F B AC 235 The asteroid of Lamarck ZOOSYSTEMA • 2021 • 43 (13) Pseudoreaster obtusangulus – Clark 1993: 318. Current stAtus.— Pseudoreaster obtusangulus (Lamarck, 1816). MAteriAl exAMined.— South Seas • MNHN-IE-2014-92; holotype (fragments); Baudin expedition (1800-1804); Péron and Lesueur leg. • MHNH; Lesueur drawings collection; ref. 74051; Baudin expedition. distribution.— North-west Australia. ocellifera Lamarck, 1816, Asterias (Fig. 25) reMArk To accommodate the Lamarckian species Asterias ocellifera, Gray (1840) established the genus Nectria. Yet Gray had not seen Lamarck’s type specimens of A. ocellifera. When working on asteroids of the genus Nectria housed in the British Museum, Perrier (1875) realised that the species N. ocellifera sensu Gray was different from the species described by Lamarck while sharing some similarities with the second Paris specimen also identified as A. ocellifera by Lamarck. He consequently proposed the new name Nectria ocellata for Gray’s specimens and gave a precise description of one of the Gray’s individuals that will be considered later as the lectotype of the species (Zeidler & Rowe 1986). As a consequence, the second specimen of Lamarck, while collected by Péron and Lesueur and being representative of the Perrier’s species, is not a type of Nectria ocellata Perrier, 1875. Asterias ocellifera Lamarck, 1816: 553 (part). Nectria ocellifera – Clark 1993: 305. Current stAtus.— Nectria ocellifera (Lamarck, 1816). MAteriAl exAMined.— South Seas • MNHN-IE-2014-24: holotype; Baudin expedition (1800-1804); Péron and Lesueur leg. • MHNH; Lesueur drawings collection; ref. 74020 and 74050; Baudin expedition. distribution.— N.ocellifera is restricted to the south part of West Australia. ocellata Perrier, 1875, Nectria (Fig. 26) Asterias ocellifera (part) Lamarck, 1816: 553. Nectria ocellata Perrier, 1875: 188. — Clark 1993: 305. Fig. 24. — Asterias obtusangula Lamarck, 1816: A, B, abactinal (A) and actinal (B) surfaces; C, adambulacral spines and a sessile pedicellaria; D, remains of the holotype (A-C, MHNH, Lesueur collection 74051, details, courtesy of MHNH; D, MNHN-IE-2014-92 ). AB C D 242 ZOOSYSTEMA • 2021 • 43 (13) Jangoux M. rosacea var. lobis quindenis Lamarck, 1816, Asterias (Fig. 33) Asterias rosacea Lamarck, 1816: 558. Anseropoda rosacea – Clark 1993: 206. Current stAtus.— Anseropoda rosacea (Lamarck, 1816). MAteriAl exAMined.— MNHN-IE-2014-62; holotype; Baudin expedition (1800-1804); Péron and Lesueur leg. • MHNH; Lesueur drawings collection; ref. 7435; Baudin expedition. distribution.— West, north and north-east Australia, Indonesia, tropical West Pacific. reMArk Lamarck (1816) gave no type locality for the species. rosacea var. lobis senis Lamarck, 1816, Asterias (Fig. 34) Asterias rosacea Lamarck, 1816: 558. Anseropoda rosacea – Clark 1993: 206. Current stAtus.— Anseropoda sp. MAteriAl exAMined.— MHNH; Lesueur drawings collection; ref. 74001; Baudin expedition; Péron and Lesueur leg. reMArk Lamarck’s variety lobis senis corresponds to a still unidentified six-armed species of Anseropoda Nardo, 1834. Lamarck gave no locality for the species. Lamarck’s material of Asterias rosacea var. lobis senis could not be found in the MNHN. Fig. 32. — Ophidiaster purpureus Perrier, 1869 abactinal and actinal surfaces of the holotype (MNHN-IE-2014-404). 243 The asteroid of Lamarck ZOOSYSTEMA • 2021 • 43 (13) senegalensis Lamarck, 1816, Asterias (Fig. 35) Asterias senegalensis Lamarck, 1816: 567. Luidia senegalensis – Clark & Downey 1992: 21. Current stAtus.— Luidia senegalensis (Lamarck, 1816). MAteriAl exAMined.— Senegal • MNHN-IE-2014-160; 2 syntypes; Adanson leg. distribution.— Florida to south of Brazil. reMArk Except for its two syntypes, the species Luidia senegalensis was never reported from African coasts. The type locality could be wrong, as it is the case for several localities given by Adanson (see Madsen 1950: 205) and the syntypes most probably come from the West Indies. In fact, L.senegalensis is a shallow water species from western Atlantic. Note that the MNHN has a third specimen of L.senegalensis, also collected by Adanson. It belonged to the Michelin collection and was presumably not seen by Lamarck. Fig. 33. — Asterias rosacea var. lobis quindenis Lamarck, 1816: A, B, abactinal (A) and actinal (B) surfaces; C, holotype of A. rosacea (actinal view); D, interradial arc and adambulacral spines (text: “from 4 to 5 spikes”); (A, B, D, MHNH, Lesueur collection 74035, details, courtesy of MHNH; C, MNHN-IE-2014-62). AB CD 244 ZOOSYSTEMA • 2021 • 43 (13) Jangoux M. Fig. 34. — Asterias rosacea var. lobis senis Lamarck, 1816: A, B, abactinal (A) and actinal (B) surfaces; C, interradial arc; D, arrangement of abactinal spines (MHNH, Lesueur collection 74001, details, courtesy of MHNH). A B Fig. 35. — Asterias senegalensis Lamarck, 1816: abactinal (A) and actinal (B) surfaces of a syntype (MNHN-IE-2014-160). AB C D 245 The asteroid of Lamarck ZOOSYSTEMA • 2021 • 43 (13) setaceus Müller& Troschel, 1842, Asteriscus (Fig. 36) Asteriscus setaceus Müller& Troschel, 1842: 43. Asterina setacea – Clark 1993: 213. Current stAtus.— Asteriscus setaceus is newly synonymized here with Paranepanthia grandis (H.L. Clark, 1928). MAteriAl exAMined.— MNHN-IE-2014-638; holotype; Baudin expedition (1800-1804); Péron and Lesueur leg. • MHNH; Lesueur drawing collection 74024; Baudin expedition. distribution.— South coast of Australia, Tasmania reMArk According to a note by Lesueur, the edges of abactinal surface of the individual are whitish. The holotype is altered and the drawings lack details. The species is tentatively identified here Paranepanthia grandis due to its shape with flat edge and to the oblique orientation of the actinal plates and armamement (see also the website Atlas of living Australia https://www.ala.org.au/, species Paranepanthia grandis (H.L. Clark, 1928)). striata Lamarck, 1816,Asterias (Fig. 37) Asterias striata Lamarck, 1816: 564. Valvaster striatus – Clark 1993: 322. Current stAtus.— Valvaster striatus (Lamarck, 1816). MAteriAl exAMined.— Mauritius • MNHN-IE-2014-36; holotype; Mathieu leg.; 1812. subulata Lamarck, 1816, Asterias (Fig. 38) Asterias subulata Lamarck, 1816: 568. Chaetaster longipes – Clark 1993: 189. Current stAtus.— Chaetaster longipes (Bruzelius, 1805). distribution.— Mediterranea and East Atlantic (from Portugal to Liberia). reMArk Lamarck gave no information on the origin of Asterias subulata specimen though its description strongly supports the synonymy with the species Chaetaster longipes. Lamarck’s type material of Asterias subulata could not be found in the MNHN. subulata Gray, 1840, Metrodira (Fig. 39) Metrodira subulata Gray, 1840: 280. — Clark 1996: 243. Current stAtus.— Metrodira subulata Gray, 1840. MAteriAl exAMined.— MHNH; Lesueur drawings collection; ref. 74017; Baudin expedition; Péron and Lesueur leg. distribution.— North part of Australia, Gulf of Bengal, Indonesia, Philippines. reMArk Péron and Lesueur’s material of the species could not be found in the MNHN. AB Fig. 36. — Asteriscus setaceus Müller & Troschel, 1842: A, B, abactinal (A) and (B) actinal surfaces (MHNH, Lesueur collection 74024, details, courtesy of MHNH). 246 ZOOSYSTEMA • 2021 • 43 (13) Jangoux M. A B Fig. 37. — Asterias striata Lamarck, 1816: abactinal (A) and actinal (B) surfaces of the holotype (MNHN-IE-2014-36). Fig. 38. — Chaetaster longipes Lamarck, 1816: abactinal and actinal surfaces of a specimen of the MNHN (MNHN-IE-2014-541). 247 The asteroid of Lamarck ZOOSYSTEMA • 2021 • 43 (13) tenuispina Lamarck, 1816, Asterias (Fig. 40) Asterias tenuispina Lamarck, 1816: 561. Coscinasterias tenuispina – Clark & Downey 1992: 427. Current stAtus.— Coscinasterias tenuispina (Lamarck, 1816). distribution.— Mediterranea; East Atlantic from Bay of Biscay to Cape Verde Islands; West Atlantic (North Carolina, Cuba, Bahamas, eastern Brazil). reMArk Lamarck’s type material could not be found in the MNHN. tessellata var. C and D Lamarck, 1816, Asterias (Fig. 41) Asterias tessellata var. C and D Lamarck, 1816: 552. Astrogonium lamarckii Müller& Troschel, 1842: 56, n. syn. Goniaster tessellatus – Clark & Downey 1992: 245. Current stAtus.— Goniaster tessellatus (Lamarck, 1816). distribution.— North Carolina to northern Brazil and Morocco to Gabon. Contrarily to what is sometimes reported, Goniaster tessellatus does not occur in the Indo-Pacific ocean. reMArk Lamarck’s type material could not be found in the MNHN. Yet, Müller& Troschel (1842), while giving no information on Lamarck’s Asterias tessellata, described a new species (Astrogonium lamarckii) that they found in the Paris collection. Undoubtedly, Astrogonium lamarckii is the first junior synonym of Lamarck’s Asterias tessellata. The holotype of the former is still kept in the MNHN (Fig. 41). It is to be assumed that the type specimen of Astrogonium lamarckii of Müller& Troschel’s would also be that of Lamarck’s Asterias tessellata. Fig. 39. — Metrodira subulata Gray, 1840: A, B, abactinal (A) and actinal (B) surfaces; C, abactinal structure of an arm section showing the minute granulation of abactinal plates and the arrangements of pairs or triplets of spines (text: ‘above’); D, actinal structure of an arm section showing the triangular inner part of adambulacral plates bearing six furrow spines and projecting above the ambulacral groove; note also the occurrence of a single broad erected subambulacral spine (MHNH, Lesueur collection 74017, details, courtesy of MHNH). AB CD 248 ZOOSYSTEMA • 2021 • 43 (13) Jangoux M. Fig. 40. — Asterias tenuispina Lamarck, 1816: abactinal and actinal surfaces of a specimen of the MNHN (MNHN-IE-2014-532). Fig. 41 . — Astrogonium lamarcki Müller & Troschel, 1842 (holotype MNHN-IE-2014-65), a junior synonym and putative type specimen of Asterias tessellata Lamarck, 1816: abactinal and actinal views. 249 The asteroid of Lamarck ZOOSYSTEMA • 2021 • 43 (13) tyloplax H.L. Clark, 1914, Linckia (Fig. 42) Linckia tyloplax H.L. Clark, 1914: 147. Hacelia tyloplax – Rowe & Gates 1995: 85. Current stAtus.— Hacelia tyloplax (H.L. Clark, 1914) (see Rowe & Gates 1995: 85). MAteriAl exAMined.— MHNH; Lesueur drawings collection; ref. 74018; Baudin expedition; Péron and Lesueur leg. distribution.— West Australia. reMArk Péron and Lesueur’s material of the species could not be found in the MNHN. variolata Lamarck, 1816, Asterias (Fig. 43) Asterias variolata Lamarck, 1816: 565. Nardoa variolata – Clark 1993: 343. Current stAtus.— Nardoa variolata (Bruzelius, 1805). MAteriAl exAMined.— MHNH; Lesueur drawings collection; ref. 74037; Baudin expedition; Péron and Lesueur leg. distribution.— East coast of Africa. reMArk Lamarck’s material could not be found in the MNHN. vernicina Lamarck, 1816, Asterias (Fig. 44) Asterias vernicina Lamarck, 1816: 554. Petricia vernicina – Clark 1993: 321. Current stAtus.— Petricia vernicina (Lamarck, 1816). MAteriAl exAMined.— South Seas • MNHN-IE-2014-28; 2syntypes; Baudin expedition (1800-1804); Péron and Lesueur leg. • MHNH; Lesueur drawings collection; ref. 74049; Baudin expedition. distribution.— South coast of Australia to New South Wales, Tasmania, temperate South-West Pacific. Fig. 42 . — Linckia tyloplax H. L. Clark, 1914: A, B, abactinal (A) and actinal (B) surfaces; C, adambulacral spines (MHNH, Lesueur collection 74018, details, courtesy of MHNH). ABC D Fig. 43. — Asterias variolata Lamarck, 1816: A, B, abactinal (A) and actinal (B) surfaces; C, furrow spines; D, abactinal plates and granules (MHNH, Lesueur collection 74037, details, courtesy of MHNH). AB C 250 ZOOSYSTEMA • 2021 • 43 (13) Jangoux M. DISCUSSION Lamarck’s (1816) revision of the asteroid collection of the Paris Muséum should have been more extensive. Indeed, part of the asteroid material collected in the South Seas by Péron and Lesueur during the Baudin expedition was not included in his revision although having been received and sorted by him in 1804 (Jangoux 2021). Fortunately, Lesueur drew with great precision each asteroid species collected during the expedition, thus creating a pictorial register allowing them to be identified. Comparative studies of Lamarck’s description, Lamarck’s type specimens and Lesueur drawings have allowed to reconsider the status of some Lamarckian species. Current and revised status are summarised in Table2. Table3 brings together the 47species of asteroids presented previously: 31 species described by Lamarck in his 1816 publication (respectively, 21 species from the South Seas, four South Seas varieties erected into species, and seven species from elsewhere) and 16 additional species from the South Seas which reached the Museum in 1804 but were described by post Lamarckian authors. Péron and Lesueur were by far the main collectors as they brought back to Paris 39 different species, all from the South Seas, among which 36 were new to Science. By chance, this number corresponds to the number Fig. 44. — Asterias vernicina Lamarck, 1816: A, B, abactinal (A) and actinal (B) surfaces; C, adambulacral armament; D, syntype of Asterias vernicina (A-C, MHNH, Lesueur collection, 74049, details, courtesy of MHNH; D, MNHN-IE-2014-28). A D BC 251 The asteroid of Lamarck ZOOSYSTEMA • 2021 • 43 (13) Table 2. — Recapitulation of the taxonomic changes proposed in the present paper. See text and Table 3 for detail. Original name Older status Current status calcar var. quinqueangula, Asterias n. syn. Patiriella regularis (Verrill, 1867) Parvulastra exigua (Lamarck, 1816) calcitrapa var. 1, Asterias n. syn. Nomen dubium Bollonaster pectinatus (Sladen, 1883) calcitrapa var. 2, Asterias n. syn. Nomen dubium Astropecten vappa Müller & Troschel, 1843 cuspidata, Asterias Goniaster tessellatus (Lamarck, 1816) Mediaster cuspidatus (Lamarck, 1816) n. comb. lamarckii, Astrogonium n. syn. Pentagonaster lamarcki (Müller & Trochel, 1842) Goniaster tessellatus Lamarck, 1816 nodosa var. 3, Asterias n. syn. Protoreaster nodosus (Linnaeus, 1758) Protoreaster lincki (Blainville, 1830) pentagonula, Asterias n. syn. Anthenea chinensis Gray, 1840 Tosia australis (Müller & Troschel, 1842) pleyadella, Asterias n. syn. Goniodiscaster pleyadella (Lamarck, 1816) Protoreaster sp. punctata, Asterias n. syn. Petricia vernicina (Lamarck, 1816) Asteropsis carinifera (Lamarck, 1816) rosacea var. lobis senis, Asterias n. syn. Anseropoda rosacea (Lamarck, 1816) Anseropoda sp. setaceus, Asteriscus n. syn. Asterina setacea (Müller & Troschel, 1842) Paranepanthia grandis (H.L. Clark, 1928) Table 3. — The Lamarckian Asteroids. Abbreviations and symbol: Död.: Döderlein; Lam.: Lamarck; M.Tr.: Müller & Troschel; WORMS: World Register of Marine Species; * synonym; 1, collections by Péron and Lesueur and drawings by Lesueur were done between 1801 and 1804; 2, Lesueur’s drawings do not represent type specimen. Original name Status Origin Collector(s)1Drawings1 Photos (type) Photos (other) 1. africanus, Asteracanthion M.Tr. Marthasterias africana (M.Tr.) South Africa Péron & LesueurLesueur – – 2. angulatus, Archaster M.Tr. Archaster angulatus M.Tr. South Seas Péron & LesueurLesueur – – 3. australiae, Anthenea Död. Anthenea australiae Död. W Australia Péron & LesueurLesueur MNHN – 4. calcar hexagona, Asterias Lam. Meridiastra gunni (Gray)* SW Australia Péron & LesueurLesueur MNHN – 5. calcar octogona, Asterias Lam. Meridiastra calcar (Lam.) SW Australia Péron & LesueurLesueur MNHN – 6. calcar quinqueangula, Asterias Lam. n. syn. Parvulastra exigua (Lam.) SW Australia Péron & LesueurLesueur MNHN – 7. calcitrapa var. 1, Asterias Lam. n. syn. Bollonaster pectinatus (Sladen) South Seas Péron & LesueurLesueur – – 8. calcitrapa var. 2, Asterias Lam. n. syn .Astropecten vappa (M.Tr.) South Seas Péron & Lesueur Lesueur – – 9. carinifera, Asterias Lam. Asteropsis carinifera (Lam.) South Seas Péron & LesueurLesueur MNHN – 10. clavigera, Asterias Lam. Mithrodia clavigera (Lam.) Mauritius Péron & Lesueur – MNHN – 11. cuspidata, Asterias Lam. Mediaster cuspidatus (Lam.) n. comb. South Seas Péron & LesueurLesueur MNHN 12. cylindrica, Asterias (part) Lam. Dactylosaster cylindricus (Lam.) Mauritius Péron & LesueurLesueur MNHN – 13. cylindrica, Asterias (part) Lam. Ophidiaster purpuratus Perrier * Mauritius Péron & LesueurLesueur MNHN – 14. decanus, Echinaster M.Tr. Plectaster decanus (M.Tr.) South Seas Péron & LesueurLesueur – – 15. discoidea, Asterias Lam. Culcita schmideliana (Bruzelius)* South Seas Péron & LesueurLesueur – – 16. echinophora, Asterias Lam. Echinaster echinophorus (Lam.) Brazil Unknown – MNHN – 17. exigua, Asterias Lam. Parvulastra exigua (Lam.) SW Australia Péron & LesueurLesueur MNHN – 18. granifera, Asterias Lam. Uniophora granifera (Lam.) South Seas Péron & LesueurLesueur MNHN – 19. helianthus, Asterias Lam. Heliaster helianthus (Lam.) ?Chile Unknown Bruguière – – 20. hesperus, Archaster M.Tr.Craspidaster hesperus (M.Tr.) South Seas Péron & Lesueur Lesueur2– – 21. lamarckii, Astrogonium M.Tr. n. syn. Goniaster tessellatus (Lam.) Atlantic Unknown – MNHN – 22. magnificum, Astrologum M.Tr. Tosia magnifica (M.Tr.) South Seas Péron & LesueurLesueur – – 23. milleporella, Asterias Lam. Fromia milleporella (Lam.) South Seas Péron & Lesueur Lesueur MNHN – 24. multifora, Asterias Lam. Linckia multifora (Lam.) South Seas Péron & LesueurLesueur – – 25. muricata, Coscinasterias Verrill Coscinasterias muricata Verrill South Seas Péron & LesueurLesueur – – 26. nodosa var. 1 & 2, Asterias Lam. Protoreaster nodosus (Linnaeus)* South Seas Péron & LesueurLesueur – – 27. nodosa var. 3, Asterias Lam n. syn. Protoreaster lincki (Blainville) Mauritius Péron & Lesueur – – MNHN 28. obtusangula, Asterias Lam. Pseudoreaster obtusangula (Lam.) South Seas Péron & LesueurLesueur MNHN – 29. ocellifera, Asterias (part) Lam. Nectria ocellifera (Lam.) South Seas Péron & LesueurLesueur MNHN – 30. ocellifera, Asterias (part) Lam.Nectria ocellata Perrier South Seas Péron & Lesueur – – MNHN 31. ophidiana, Asterias Lam. Ophidiaster ophidianus (Lam.) Mediterranea Unknown – – WORMS 32. penicillaris, Asterias Lam. Asterinopsis penicillaris (Lam.) South Seas Péron & LesueurLesueur – – 33. pentagonula, Asterias Lam. n. syn. Tosia australis Gray South Seas Péron & LesueurLesueur – – 34. pleyadella, Asterias Lam. n. syn. Protoreaster sp. South Seas Péron & LesueurLesueur MNHN – 35. punctata, Asterias Lam. n. syn. Asteropsis carinifera (Lam.) South Seas Péron & LesueurLesueur MNHN – 36. purpureus, Ophidiaster Perrier Ophidiaster hemprichi M.Tr.*Mauritius Péron & Lesueur Lesueur – – 37. rosacea lobis quindenis, Asterias Lam. Anseropoda rosacea (Lam.) South Seas Péron & LesueurLesueur MNHN – 38. rosacea lobis senis, Asterias Lam. n. syn. Anseropoda sp. South Seas Péron & LesueurLesueur – – 39. senegalensis, Asterias Lam. Luidia senegalensis (Lam.) West Indies Adanson – MNHN – 40. setaceus, Asteriscus M.Tr. n. syn. Paranepanthia grandis H. L. Clark South Seas Péron & LesueurLesueur – – 41. striata, Asterias Lam. Valvaster striatus (Lam.) Mauritius Mathieu – MNHN – 42. subulata, Asterias Lam. Chaetaster longipes (Bruzelius)* Mediterranea Unknown – – MNHN 43. subulata, Metrodira Gray Metrodira subulata Gray South Seas Péron & Lesueur Lesueur – – 44. tenuispina, Asterias Lam. Coscinasterias tenuispina (Lam.) Mediterranea Unknown – – MNHN 45. tessellata var. C & D, Asterias Lam. Goniaster tessellatus (Lam.) Atlantic Unknown – MNHN – 46. tyloplax, Lincka H. L. Clark Hacelia tyloplax (H. L. Clark) South Seas Péron & LesueurLesueur – – 47. variolata, Asterias Lam. Nardoa variolata (Bruzelius) Mauritius Unknown Lesueur2– – 48. vernicina, Asterias Lam. Petricia vernicina (Lam.) South Seas Péron & LesueurLesueur MNHN –