NOTES ICHTYOLOGIQUES

ON THE AUTHORSHIP, IDENTITY AND be determined by re-examination of original TAXONOMIC POSITION OF PLEURONEC- published literature, whereas the question of TES COMMERSONNII LACEPÈDE, 1802 identity of this species can only be resolved by (PLEURONECTIFORMES, ). referring to the unpublished illustration of this Thomas A. MUNROE, National Marine Fishe- species prepared for Commerson, which provides ries Service National Systematics Laboratory, new information that permits the correct taxo- National Museum of Natural History, Washington, nomic placement for this nominal species. DC 20560-0153, USA. [[email protected]. Date of authorship, and authorship itself, edu] & Martine DESOUTTER, Laboratoire for Lacepède’s nominal species, Pleuronectes d’Ichtyologie générale et appliquée, Muséum commersonnii, have been the subject of alternative national d’Histoire naturelle, 43 rue Cuvier, 75231 views by subsequent investigators. Contributing to Paris cedex 05, FRANCE. [[email protected]] confusion surrounding the authorship and taxo- nomic placement of this nominal species are the RÉSUMÉ.!-!Sur la paternité, l’identité et la lack of type specimens and inconsistencies be- position taxinomique de Pleuronectes commer- tween the original description and the published sonnii Lacepède, 1802 (Pleuronectiformes, Solei- illustration of the species. This confusion is dae). largely due to the fact that the illustration of this Les auteurs discutent et démontrent la va- species published one year prior (Lacepède, 1801) lidité taxinomique de l’espèce décrite par Lace- to the description (Lacepède, 1802) was labelled pède, en 1802, sous le nom de Pleuronectes with a vernacular name, which was incorrectly commersonnii. Cette espèce appartient bien à la interpreted as a scientific binomial by some famille des Soleidae et est maintenant placée dans subsequent authors (e.g., Cantor, 1849; Günther, le genre . 1862). Cuvier (1817) attributed authorship of the species to Lacepède (1801), but then in the same Key!words.!-!Soleidae - Pleuronectes commerson- paper questioned whether this name should be nii - . considered a nomen nudum. Swainson (1839) attributed authorship of this species to Russell (as commersonni) without Lacepède figured (1801: plate 12, fig.!2) further comment. Cantor (1849) and Günther and described (1802) a nominal species of flatfish, (1862) both attributed authorship to Lacepède, but Pleuronectes commersonnii, based entirely on considered the correct scientific name of the information originally contained in an unpublished species to be based on the name “Le Pleuronecte illustration and manuscript of Commerson (MS Commersonnien” accompanying the illustration of 528, XXV.181; “Bibliothèque centrale, Muséum this nominal species published in 1801, rather than national d’Histoire naturelle, Paris”). Since the Pleuronectes commersonnii applied in the text publication of Lacepède’s work, considerable description of 1802. In his revision of the soleid confusion has ensued concerning the authorship Synaptura, Chabanaud (1938) also doubted and date of authorship of this nominal species, as that Lacepède actually provided a binomial for well as its identity and taxonomic placement. The this nominal species and attributed authorship of name proposed by Lacepède has most frequently the species to Swainson (1839). Subsequent works been applied to a commonly occurring species of (Menon and Joglekar, 1978) disagreed with soleid flatfish (usually Synaptura commersonnii) Chabanaud (1938), and instead, referred of some commercial importance in the Indian Lacepède’s species either to the soleid genera Ocean (Menon and Joklekar, 1978). No type Brachirus, or more often, Synaptura. Fricke specimen of this nominal species has been found (1999) recently concluded that two species were (the specimen illustrated is assumed lost), nor described by Lacepède (1801, 1802). have we any reason to believe that one is still Establishing the appropriate date and extant. Given the long history of association of authorship for this nominal species requires Lacepède’s name with this commercially impor- evaluating the name associated with the illustra- tant flatfish, the uncertainty associated with this tion published in Lacepède (1801). The illustration name needs to be resolved in order to stabilize (originally prepared for Commerson) was accom- nomenclature for this species. Authorship and date panied in Lacepède (1801) only by the vernacular of authorship of Pleuronectes commersonnii can name “Le Pleuronecte Commersonnien.” The

Cybium 2001, 25(3): 273-277. 274 MUNROE & DESOUTTER name “Le Pleuronecte Commersonnien” in in the original description has been the major Lacepède (1801) was incorrectly interpreted as a factor contributing to the confusion regarding the latinized binominal for the species by some subse- identity of this nominal species. The figure pub- quent authors (e.g., Cantor, 1849; Günther, 1862). lished in Lacepède (1801) depicts a sinistral Rather, it is a vernacular name constructed in a flatfish with soleid features (Fig.!1a). Lacepède fashion consistent with other vernacular names for (1802) placed this species in his all-encompassing their respective species. Thus, “Le Pleuronecte family (sensu Linnaeus) among the Commersonnien” does not constitute a scientific sinistral pleuronectiform species. The written name for this nominal species. This conclusion account (Lacepède, 1802) also describes a sinistral agrees with those of Chabanaud (1938) and Esch- species (as did the account in Commerson’s meyer (1998) that the date of authorship for unpublished manuscript), but based on the infor- Pleuronectes commersonnii Lacepède does not mation contained therein, the taxonomic affinity derive from the illustration published in 1801. of this species among sinistral can not be Formal description of this nominal spe- ascertained (see also Chabanaud, 1937). cies occurs in Lacepède (1802: 656-657), which Cuvier (1817) first recognised the pro- includes a written description and latinized bino- blem of discrepancies between the written descrip- mial (Pleuronectes commersonnii) for the nominal tion and the fish figured in Lacepède (1802: 656- species represented in the drawing published in 657 and 1801, pl.!12, fig.!2, respectively). He 1801 as evidenced by cross-referencing of the suggested that perhaps at least two species were vernacular name “Le Pleuronecte Commerson- included in the original description. Nevertheless, nien” between the two publications. This finding after noting these discrepancies, Cuvier, perhaps is contrary to the synonymy presented in Cha- more influenced by the features of the illustrated banaud (1938), which did not consider Lacepède fish than by the written description, placed Pleu- (1802) as the author of Pleuronectes commerson- ronectes commersonnii Lacepède into his subdivi- nii. sion of flatfishes containing the soles. Swainson The identity and taxonomic placement of (1839) and Cantor (1849), also placed this nomi- Lacepède’s nominal species has long been the nal species among soleid fishes (dextral flatfishes, subject of differing opinions. While a majority of e.g., with eyes on the right side of the body). Since previous authors beginning with Cuvier (1817) Cantor (1849), there has been long-standing considered Pleuronectes commersonnii as a valid assignment of this species in the soleid genus soleid flatfish species (flatfish species with dextral Synaptura (see synonymy in Chababaud, 1938). orientation), other authors have questioned the Günther (1862), although expressing some uncer- exact identity and taxonomic placement of tainty about the authorship of the species, placed it Lacepède’s species (Chabanaud, 1937; Esch- in the soleid genus Synaptura. Chabanaud (1937), meyer, 1998; Fricke, 1999). Re-examination of after lengthy discussion of the confusion surroun- Lacepède (1801, 1802) and the unpublished ding the name Pleuronectes commersonnii, was Commerson manuscript reveals that Lacepède unable to reconcile placing this nominal species, described this nominal species based entirely on based on a sinistral specimen, into the Soleidae. information originally contained in the unpu- After careful consideration and elimination of the blished illustration and manuscript of Commerson possibility that the specimen Lacepède described (MS 528, XXV.181). No evidence exists to indi- may have been a reversed soleid, and after consi- cate that Lacepède either actually examined the dering other alternative taxonomic placements, specimen illustrated for Commerson, or any other Chabanaud (1937) concluded that identification specimen of this nominal species while preparing and correct taxonomic placement of Lacepède’s his description of Pleuronectes commersonnii. species was unresolvable until further information Furthermore, no type specimens were designated became available. Nonetheless, in the next year, by Lacepède and no potential types are known to Chabanaud (1938) considered this species as a exist for this nominal species (Chabanaud, 1937), member of the soleid genus Synaptura (although a situation that precludes determination of the attributing authorship to Swainson). Although identity and taxonomic placement of this species detailed, Chabanaud’s (1937, 1938) discussions by direct examination of specimens. essentially repeated arguments presented by The inconsistency between the informa- Cuvier (1817) regarding discrepancies between tion available in the illustration and that provided the illustration and written description appearing On the authorship, identity and taxonomy of Pleuronectes commersonnii 275

Fig.!1.!-!Illustrations of Pleuronectes commersonnii. a: Illustration originally published as figure!2 of plate 12 in Lacepède (1801). b: Unpublished drawing (MS 528, XXV.181; Bibliothèque centrale du Muséum national d’Histoire naturelle, Paris) prepared by Commerson’s artist, Jossigny. in Lacepède (1801, 1802) and did not involve any other hand, consulted the unpublished manuscript examination of Commerson’s original drawing. and drawings of Commerson. However, after Thus, his conclusions were made without benefit referring to this work and that of Lacepède, he of this resource. Menon and Joglekar (1978), in arrived at several erroneous conclusions. Fricke their systematic study of some soleid genera (1999: 570, 573) recognised that the illustration occurring in Indian waters, considered Lacepède’s and written description of Lacepède were pub- species to be a member of Synaptura within the lished in different years (1801, 1802) and then Soleidae without consulting the original descrip- concluded that two distinct species were treated in tion or figures of Commerson. Lacepède’s works. Fricke (1999: 573) treated More recently, Eschmeyer (1998) and Pleuronecte Commersonnien Lacepède, 1801, as a Fricke (1999), although accepting Lacepède as the junior synonym of the soleid marmo- original describer, suggested quite different taxo- ratus with an authorship of Commerson and nomic placements (Pleuronectidae and , Lacepède in Lacepède (1802) based on the fish respectively) for this nominal species. Eschmeyer illustrated in figure!2 of plate 12 of Lacepède (1998) did not provide justification for consider- (1801). Although this figure is a soleid, the spe- ing Lacepède’s nominal species as a member of cies illustrated with the dorsal and anal fins con- the dextral Pleuronectidae. Fricke (1999), on the joined with the caudal fin is definitely not Parda- 276 MUNROE & DESOUTTER chirus marmoratus, which has dorsal and anal fins conjoined dorsal, anal and caudal fins) all strongly separate from the caudal fin. Apparently, Fricke indicate that the illustrated fish was a soleid (1999) confused figures!2 and 3 on plate 12 of flatfish. Lacepède (1801). He erroneously concluded that If Pleuronectes commersonnii is a soleid figure!2 was based on a large, bleached specimen flatfish, then why would it be depicted in the of , when in fact figure!3 published illustration in a sinistral orientation illustrates Pardachirus marmoratus. It is this mix since soleids, except for rare cases of asymmetry up, we believe, which prompted Fricke into reversal (see Gudger, 1935), typically have a erroneously placing Pleuronecte Commersonnien dextral orientation!? Consulting the unpublished in the synonymy of Pardachirus marmoratus. drawing of this fish prepared by Commerson’s Furthermore, the name appearing on plate 12 artist, Jossigny, reveals the answer to this ques- published in Lacepède (1801) is a vernacular tion. The original drawing of the species subse- name, not a scientific binomial. The fact that quently described by Lacepède (1802) as Pleu- Fricke did not italicise this name in the synonymy ronectes commersonnii clearly shows this illustra- of Pardachirus marmoratus suggests that he tion is based on a dextral species of flatfish recognised it as a vernacular name, but then erred (Fig.!1b) and the original unpublished drawing in treating this as an available name for a species prepared by Jossigny renders an assignment of this different than that appearing in the written descrip- species to the Soleidae unambiguous, which is an tion. The name Pleuronectes commersonnii pro- important conclusion given that no type specimens posed in Lacepède (1802) is, in contrast, the only are known to exist. For some inexplicable reason, scientific name used in the original description of perhaps to match the written description appearing this species. That only one species was intended is in Commerson’s unpublished manuscript, the evidenced by the cross-referencing between the original drawing of a dextral soleid appears as a illustration and text description within Lacepède’s sinistral flatfish (see Fig.!1a) when published in works (1801, 1802). Lacepède (1801). Although not commonly occur- The second name Fricke treated (1999: ring, figures of flatfishes appear even in recently 570) was that of Pleuronectes commersonnii published works with unintentional reversal in (authorship listed as Commerson and Lacepède in symmetry due to publisher’s errors when arrang- Lacepède, 1802) based on the text description ing photographs of these fishes (a recent example (pp.!599-600, 652-657). He concluded that this is the unintentionally reversed specimen of nominal species was referrable to the bothid Aseraggodes haackeanus in figure!361, p.!860, of flatfish, Bothus mancus (Broussonet, 1782). Gomon et al., 1994). Because Lacepède’s description is somewhat In conclusion, Pleuronectes commerson- vague, we can not rule out the possibility that nii Lacepède (1802) is a valid name for a soleid more than one species was included in the original flatfish. This determination is based on character- description, or that the written description was istics of the fish illustrated in Lacepède (1801), in based on a species other than a soleid. However, combination with the dextral orientation of the we completely disagree with Fricke’s assignment fish in the unpublished drawing of this species of Pleuronectes commersonnii in the synonymy of originally prepared for Commerson. Given the Bothus mancus within the Bothidae because the long history (beginning with Cuvier, 1817) of illustration (Fig.!1a) of “Le Pleuronecte Commer- recognition of Pleuronectes commersonnii as an sonnien” clearly does not represent a member of available name for a species of soleid flatfish and the family Bothidae (see discussion in Chabanaud, the information contained in the unpublished 1937). Also, the overall body shape, a terminal drawings of this species, it is in the best interest of mouth without overhanging snout, presence of a nomenclatural stability to continue to recognise pectoral fin, the single lateral line and caudal fin Lacepède’s species as a member of the Soleidae. shape, definitely distinguishes the figured speci- Features of the illustrated fish indicate that this men from any member of the Cynoglossidae. species is assignable to the genus Synaptura However, given the sinistral orientation of the fish (sensu Chabanaud, 1938; Desoutter et al., 2001). illustrated, it is difficult to confidently place this Therefore, the appropriate current nomenclature species in the Soleidae, although certain features for this species is Synaptura commersonnii of the illustrated fish (e.g., the small mouth, the (Lacepède, 1802). preopercle concealed beneath the skin, and the On the authorship, identity and taxonomy of Pleuronectes commersonnii 277

Acknowledgments.!-!R. Vari, United States ESCHMEYER W.N., 1998.!-!Catalog of fishes. National Museum (USNM), provided comments Spec. Publ. n°!1. 3 vols, 2905 p. California on an earlier draft. We thank D. Smith (USNM) Academy of Sciences. for assistance with interpretations of the Interna- FRICKE R., 1999.!-!Fishes of the Mascarene tional Code of Zoological Nomenclature. Funding islands (Réunion, Mauritius, Rodriguez). An for TM to work in the Laboratoire d’Ichtyologie annotated checklist with descriptions of new générale et appliquée was provided by the visiting species. Theses Zoologicae 31: 759!p. Koe- professorship program at the Muséum national nigstein Koeltz Scientific Books. d’Histoire naturelle. Figures were prepared from GOMON M.F., GLOVER J.C.M. & R.H. slides provided by the “Bibliothèque centrale du KUITER, 1994.!-!The fishes of Australia’s Muséum national d’Histoire naturelle”. south coast. 992 p. Adelaide: State Print. GUDGER E.W., 1935.!-!Abnormalities in flat- fishes (Heterosomata). I. Reversal of sides. A REFERENCES comparative study of the known data. J. Mor- phol., 58(1): 1-39. CANTOR T.E., 1849.!-!Catalogue of Malayan GÜNTHER A., 1862.!-!Catalogue of the fishes in Fishes. Asiat. Soc. Bengal J., 18(2): 981- the British Museum. Catalogue of the Acan- 1443. thopterygii Pharyngognathi and Anacanthini CHABANAUD P., 1928.!-!Remarques sur quel- in the collection of the British Museum. ques genres de la famille des Soleidae. Bull. Vol.!4: 534 p. Soc. Zool. Fr., 53: 272-279. LACEPÈDE B.G.E., 1801.!-!Histoire naturelle des CHABANAUD P., 1937.!-!Qu’est ce que le poissons, 3: i-lxxi!+!558 p. “Pleuronectes commersonien” de Lacepède? LACEPÈDE B.G.E., 1802.!-!Histoire naturelle des Bull. Mus. natn. Hist. nat., 9(3): 193-198. poissons, 4: i-xliv!+!728 p. CHABANAUD P., 1938.!-!Contribution à la MENON A.G.K. & A. JOGLEKAR, 1978.! morphologie et à la systématique des Téléos- -!Taxonomic status of the genera Synaptura téens assymétriques. 1re partie. Révision du Cantor, 1550 (sic) and Euryglossa Kaup, genre Synaptura. Arch. Mus. Hist. Nat., (6)15: 1858 with descriptions of species referable to 61-108. these occurring in seas of India. J. Mar. Biol. CUVIER G., 1817.!-!Le Règne distribué Assoc. India, 20(1-2): 10-21. d’après son Organisation pour Servir de Base SWAINSON W., 1839.!-!The natural history and à l’Histoire naturelle des Animaux et classification of fishes, amphibians, and repti- d’Introduction à l’Anatomie comparée. Les les, or monocardian . London. Vol. 1: Reptiles, les Poissons, les Mollusques et les i-vi!+!448 p. Annélides. Edition 1, 2: i-xviii!+!532 p. DESOUTTER M., MUNROE T.A. & F. CHAPLEAU, 2001.!-!Nomenclatural status of Brachirus Swainson, Synaptura Cantor and Euryglossa Kaup (Soleidae, Pleuronectifor- Reçu le 12.12.2000. mes). Ichthyol. Res., 48: in press. Accepté pour publication le 04.07.2001.