Zootaxa 1893: 65–68 (2008) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Correspondence ZOOTAXA Copyright © 2008 · Magnolia Press ISSN1175-5334(online edition) Comments on “Chersine Merrem, 1820 and Chersina Gray, 1831: a nomenclatural survey by Bour & Ohler, Zootaxa, 1752: 66–68” UWE FRITZ1 & OTTO KRAUS2 1Museum of Zoology (Museum für Tierkunde), Natural History State Collections Dresden, A. B. Meyer Building, D-01109 Dresden, Germany. E-mail: [email protected] 2Zoologisches Institut und Zoologisches Museum, Universität Hamburg, Martin-Luther-King-Platz 3, D-20146 Hamburg, Germany. E-mail: [email protected] In a recent correspondence, Bour & Ohler (2008) readdress the question of whether or not Testudo hermanni Gmelin, 1789 is the type species of the genus-group names Chersine Merrem, 1820 and Medaestia Wussow, 1916 (Reptilia: Testudines: Testudinidae). The senior author, Roger Bour, co-authored the earlier description of a new genus-group name for T. hermanni (Eurotestudo Lapparent de Broin, Bour, Parham & Perälä, 2006), a name considered as an objective jun- ior synonym of both Chersine and Medaestia by Fritz & Bininda-Emonds (2007) and Fritz & Havaš (2007), who argued that all three names are based on the same type species, T. hermanni. Bour & Ohler (2008) disputed this conclusion. In the present paper, we wish to clarify nomenclatural issues raised by Bour & Ohler (2008). Which is the type species of Chersine Merrem, 1820? Bour & Ohler (2008) claim that Chersine is a replacement name (nomen novum) for Testudo Linnaeus, 1758 (type species: Testudo graeca Linnaeus, 1758), and consequently has the same type species, because Merrem (1820: p. 29) ‘only included Testudo as synonym of Chersine.’ However, Merrem (1820: p. 17) listed Testudo as synonym of his order ‘Schild-Pholidoten (Testudinata)’ as well, indicating a different concept of Testudo compared to its current use. When the nameTestudo was introduced, Linnaeus (1758) applied it to all eleven chelonian species known to him (Table 1), includ- ing sea turtles (Cheloniidae), New World pond turtles (Emydidae), tortoises (Testudinidae), and snapping turtles (Chely- dridae). A type species for Testudo was not fixed until Fitzinger (1843: p. 29) designated Testudo graeca Linnaeus, 1758, implying that when Chersine was erected, Testudo was still without type species and covering a wide range of highly dis- tinct chelonian species. Hence, the argument by Bour & Ohler (2008) is invalid. TABLE 1. Chelonian species described in the 10th Edition of the Systema Naturae by Linnaeus (1758) and their current taxonomic allocation according to Fritz & Havaš (2007). Original name Current name Family Testudo caretta Caretta caretta Cheloniidae Testudo carinata Terrapene carolina Emydidae Testudo carolina Terrapene carolina Emydidae Testudo geometrica Psammobates geometricus Testudinidae Testudo graeca Testudo graeca Testudinidae Testudo lutaria Emys orbicularis Emydidae Testudo mydas Chelonia mydas Cheloniidae Testudo orbicularis Emys orbicularis Emydidae Testudo pusilla Testudo graeca Testudinidae Testudo scabra unclear unclear Testudo serpentina Chelydra serpentina Chelydridae Accepted by S. Carranza: 18 Aug. 2008; published: 6 Oct. 2008 65 Chersine, for which Merrem (1820: p. 29) gave a clear bilingual diagnosis in German and Latin, is not a replacement name of Testudo. It was created to comprise only ‘Landschildkröten’ (= tortoises; Merrem 1820: footnote p. 29) but, in contrast to Linnaeus (1758), not sea turtles (placed by Merrem into Caretta and Sphargis) and not most freshwater turtles (placed by Merrem into Emys,Matamata, Terrapene, and Trionyx). This is also evident from the fact that Merrem included in the synonymy of Chersine, besides Testudo, Pliny’s Testudines terrestres. Parenthetically it may be noted, however, that Merrem’s ‘tortoises’ included quite a variety of chelonians of distinct families, not only true tortoises of the family Testudinidae (Table 2). TABLE 2. Chelonian species placed by Merrem (1820) in Chersine and their current taxonomic allocation according to Fritz & Havaš (2007). Merrem’s name Current name Family Chersine areolata Homopus areolatus (Thunberg, 1787) Testudinidae Chersine calcarata Geochelone sulcata (Miller, 1779) Testudinidae Chersine denticulata Chelonoidis denticulata (Linnaeus, 1766) Testudinidae Chersine elegans Geochelone elegans (Schoepff, 1795) Testudinidae Chersine fasciata Homopus areolatus (Thunberg, 1787) Testudinidae Chersine geometrica Psammobates geometricus (Linnaeus, 1758) Testudinidae Chersine graeca Testudo hermanni Gmelin, 1789 Testudinidae Chersine marginata Testudo marginata Schoepff, 1793 Testudinidae Chersine Mühlenbergii Glyptemys muhlenbergii (Schoepff, 1801) Emydidae Chersine planitia Macrochelys temminckii (Troost, 1835) Chelydridae Chersine punctularia Rhinoclemmys punctularia (Daudin, 1802) Geoemydidae Chersine pusilla Testudo graeca Linnaeus, 1758 Testudinidae Chersine retusa Cylindraspis indica (Schneider, 1783) Testudinidae Chersine rotunda Cylindraspis peltastes (Duméril & Bibron, 1835) or Cylindraspis Testudinidae vosmaeri (Suckow, 1798)? Chersine scorpioides Kinosternon scorpioides (Linnaeus, 1766) Kinosternidae Chersine signata Homopus signatus (Gmelin, 1789) Testudinidae Chersine tessellata Chelonoidis denticulata (Linnaeus, 1766) Testudinidae Chersine tetradactlya Homopus areolatus (Thunberg, 1787) Testudinidae More than a century after the original description of Chersine, Lindholm (1929) fixed ‘Testudo graeca “L.” auct. = Testudo hermanni Gmelin’ as its type species. Bour & Ohler (2008) argue that this nomenclatural act would represent an ‘unforeseen designation’ and were ‘certainly not in accordance with Merrem’s design when he created the genus.’ But Lindholm (1929) clearly referred to the reversal of the meaning of T. graeca that occurred only a few years before his own paper was published. Merrem (1820) evidently meant the taxonomic species T. hermanni that was previously known under the name of T. graeca. We cite here Fritz & Bininda-Emonds (2007: pp. 303–304): “In support of erecting the new genus Eurotestudo (type species: T. hermanni Gmelin, 1789) in the face of two older names being available (Chersine Merrem, 1820 and Medaestia Wussow, 1916), Lapparent de Broin et al. (2006) claim that these latter names were based on the type species T. graeca Linnaeus, 1758 and, therefore, cannot be applied to T. hermanni. In so doing, Lapparent de Broin et al. (2006) have unfortunately added a further chapter to the complicated and confusing nomenclatural history of Testudo. The name of T. graeca was misapplied for decades after its description by Linnaeus (1758) by virtually all subsequent authors (e.g., Schoepff 1792–1801; Hermann 1804; Merrem 1820; Fitz- inger 1826; Gray 1844, 1870; Strauch 1862; Schreiber 1875; Lortet 1887; Boulenger 1889; Siebenrock 1909), who used this name for the species now known as T. hermanni Gmelin, 1789. It was not until the early 20th century that Siebenrock (1913) and Flower (1925, 1926) recognized this error. Whereas Siebenrock (1913) hesitated to change the long-estab- lished usage of the name T. graeca, Flower (1925, 1926) pointed out that the name T. graeca has to be applied to the spe- cies previously known under the name of T. ibera Pallas, 1814 and that T. hermanni Gmelin, 1789, a name treated for 66 · Zootaxa 1893 © 2008 Magnolia Press FRITZ & KRAUS more than a century as junior synonym of T. graeca Linnaeus, 1758, has to be used for what was known before as T. graeca. […] As regards the names Chersine Merrem, 1820 and Medaestia Wussow, 1916, it is obvious from the period when they were proposed that they were based on the misapplication of the name T. graeca Linnaeus, 1758 and that actu- ally T. hermanni was meant.” That Merrem (1820: p. 31) referred to the taxonomic species Testudo hermanni indeed is demonstrated by his spe- cies diagnosis, translated as follows (italics as in the original; Fig. 1): “graeca 38. T. C. Shell broad, inversed ovoid, rear humpy. Areoles of scutes coarse; feet with scales. Lives in the neighbourhood of the Mediterranean Sea. Shell distinctly domed. Plastron anteriorly weakly, posteriorly distinctly emarginated. Tail short, cone- shaped, with bent, horny tip. Claws5, 4, sometimes 4, 4.” FIGURE 1. Reproduction of the bilingual diagnosis of Chersine graeca by Merrem (1820). It is obvious from the diagnosis that the taxonomic species Testudo hermanni was meant (see text). That Merrem named the species in German ‘echte’ (= true) [tortoise] may have prompted Lindholm (1929) to designate T. hermanni as the type species of Chersine. The horny, claw-like scale at the tip of the tail is diagnostic for T. hermanni (Cheylan 2001) and provides unambigu- ous evidence that this species, and not T. graeca, was meant. Considering this, it is obvious that Lindholm (1929) delib- erately used the wording ‘Testudo graeca “L.” auct. = Testudo hermanni Gmelin’; he intended to point out that Merrem (1820) actually referred to the taxonomic species T. hermanni when using the name graeca. Thus, Lindholm fulfilled the requirements of Article 69.2.4 of the Code (ICZN 1999): “If an author subsequently designates as type species a species originally included […] as an expressly stated misidentification or misapplication of a previously established nominal species, the species so designated is the nominal species denoted by the name of the taxonomic species actually involved (and not the nominal species cited).” In consequence, the type species of Chersine Merrem, 1820 is clearly T. hermanni Gmelin, 1789. Which is the type species of Medaestia Wussow, 1916? Medaestia is a genus-group name for tortoises that was never used as valid except in its original description, an arti- cle about the captive maintenance of tortoises (Wussow 1916). The following species were originally included (as spelled by Wussow 1916: p. 170; in parentheses the current names of the taxonomic species actually involved): ‘Medaes- tia graeca’ (= Testudohermanni), ‘M. horsfieldi’ (= T. horsfieldii), ‘M. ibera’ (= T. graeca), and ‘M. leithi’ (= T.klein- manni). Mertens (1949) formally designated the type species of Medaestia as follows: ‘I fix it here as graeca (hermanni Gmelin).’ By adding ‘hermanni Gmelin’ in brackets, Mertens (1949) clearly expressed that he was aware that Wussow (1916) used the name graeca in the contemporary sense, to denote the taxonomic species later known under the name hermanni. This act is in accordance with Article 69.2.4 of the Code (ICZN 1999) as well, which is why Testudo her- manni Gmelin, 1789 is the valid type species of Medaestia Wussow, 1916. COMMENTS ON CHERSINE Zootaxa 1893 © 2008 Magnolia Press · 67 Conclusions Chersine Merrem, 1820, Medaestia Wussow, 1916, and Eurotestudo Lapparent de Broin, Bour, Parham & Perälä, 2006 are objective synonyms. They are all based on the same type species, Testudo hermanni Gmelin, 1789, for which prior to Flower (1925, 1926) the name T. graeca Linnaeus, 1758 was widely used. The phylogenetic analyses of Fritz & Bininda-Emonds (2007) based on a five-gene data set (mtDNA: 12S rRNA, 16S rRNA, cyt b; nDNA: C-mos, Rag2) rep- resenting approximately two-thirds of all extant testudinid species and all species of Testudo did not support the removal of T. hermanni from Testudo, rendering the discussion started by Bour & Ohler (2008) academic. If within Testudo a nomenclatural distinction were desired for the clade comprising T. hermanni andT. horsfieldii, the oldest available name, Chersine, would have to be applied in subgeneric rank. 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