The asteroid species of Lamarck (Echinodermata: Asteroidea)

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.


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.
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.   (Gray, 1840).         reMArk Though the species was collected by Péron and Lesueur during the Baudin expedition, no drawings of this species could be found in the Lesueur collection at Le Havre Museum.   reMArks Both Lesueur's drawings and the original holotype label attest that Lamarck's Asterias cuspidata originates from the South Seas. Surprisingly A. cuspidata was considered a synonym of the Atlantic Asterias tessellata Lamarck, 1816, which is more than unlikely both geographically and morphologically. Lamarck himself reported that A. cuspidata "differs from Asterias tessellata because its angles continue with long spikes resembling straight horns or rays [On l'en distingue par ses angles prolongés en longues pointes comme des cornes droites ou des rayons]". In fact, the cuspidata-tessellata problem results from the confusion between Asterias cuspidata Lamarck and Goniaster cuspidatus Gray, 1840 the latter being indeed synonym of Lamarck's Asterias tessellata. From its overall morphological features, Asterias cuspidata belongs to the Goniasteridae, and its closest genus is Mediaster Stimpson, 1857. Indeed A. cuspidata is clearly related to the Australian species Mediaster australiensis H. L. Clark, 1916(Clark 1916. 9), a species known from Bass Strait and around Tasmania. Some differences occur, however, such as the general appearance (the disc area of A. cuspidata is narrower giving it a more stellate form) or the occurrence in A. cuspidata of a row of enlarged carinal plates. Therefore, awaiting additional information of M. australiensis variability, it sounds better to keep the two species separate.      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).

AbbreviAtions
granifera Lamarck, 1816, Asterias (Fig. 16) Asterias granifera Lamarck, 1816: 560.    (Müller & Troschel, 1840).      reMArk Péron and Lesueur's material could not be found in the MNHN.         reMArks Asterias pentagonula was often said to belong to the genus Anthenea which includes asteroids with an enlarged disc and relatively short though well-individualised arms. However, Lamarck's description emphasises the peculiar shape of the species as "Aplati, presque discoïde, avec des angles courts [flat, almost disc-shaped, with shallow angles]". Such an appearance does not fit with Anthenea and rather suggests that the species belongs to the genus Tosia Gray, 1840, a suggestion that is consistent with both its origin (South Seas) and collectors (Péron and Lesueur). Several authors consider that Asterias pentagonula is a Chinese species. This is unlikely and could result from the synonymy of A. pentagonula with Anthenea chinensis Gray, 1840in Müller & Troschel (1842. This is a rather strange proposal as Gray's species originated from the China-Japan area while Asterias pentagula came from the South Seas. Liao & Clark (1995: 99) consider that these species are two different taxa. The MNHN specimens identified Anthenea pentagonula belong to the species Anthenea australiae Döderlein, 1915; see previously under that species. reMArk When discussing Asterias pleyadella status, Perrier (1875: 275) suggested that its representatives are juveniles closer to oreasterids than to goniasterids. He consequently related A. pleyadella to the genus Pentaceros Gray, 1840. This makes sense, indeed, considering the overall shape of the individuals, their carinate arms and, particularly, the five radially positioned tubercles surrounding the central part of the disc.
I agree with Perrier's analysis but consider Asterias pleyadella a juvenile Protoreaster, the common oreasterid genus of the Indian Ocean. Lesueur drew (pencil drawings) the best preserved individual of A. pleyadella.

reMArks
There is a strange confusion between Lamarck's Asterias punctata and Lamarck's Asterias vernicina, the two species being erroneously considered synonymous. This confusion presumably arose from Perrier's stay in the British Museum in the early 1870s.
After having compared London and Paris asteroid specimens, he proposed, with reason, to synonymyse Lamarck's Asterias vernicina and Gray's Petricia punctata, which he confirmed in his 1875 revision, combining the two species under the name Petricia vernicina. Of course, this did not imply that Lamarck's Asterias punctata was also a synonym of Lamarck's Asterias vernicina, a synonymy which was, however, tacitly accepted by subsequent authors. The situation became still more confused as neither Müller & Troschel (1842) nor Perrier (1875) discussed Lamarck's Asterias punctata. They found no trace of A. punctata individuals in the MNHN which would mean the specimens have been lost for quite a long time. Lamarck's Latin diagnosis of Asterias punctata is as follows: "pentagona, inermis, utrinque tessellata ; tesselis dorsi sinuatoangulatis, punctatis ; margine articulato [pentagonal, spineless [and] checkered on both sides; upper side perforate with rounded tiles; articulated arm". Nota: "Tiles" refer to abactinal plates and 'perforate' to papulae.] Both this diagnosis and the MNHN pictures of Asteropsis carinifera syntypes are clearly reminiscent of the shape of juveniles Asteropsis carinifera already illustrated by Loriol (1885: 67, pl 20 figs 7-10). One may therefore conclude that A. punctata and A. carinifera are synonyms, the former corresponding to the juvenile form of the species.  (Müller & Troschel, 1842)). Yet Perrier did not consider the second Lamarck specimen of Asterias cylindrica to designate the holotype of O. purpureus. He assigned as holotype an individual collected by Rousseau in the Seychelles in 1842.  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.  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.   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.

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