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Zootaxa 1752: 66
& ANNEMARIE OHLER
In the tenth Edition of his ‘Systema Naturae’, Carolus Linnaeus presented among the species of the genus Testudo
(including all the turtles and tortoises, thus corresponding to the present order of Chelonians) the species Testudo graeca
(Linnaeus 1758). The reference was the description and the plate published by George Edwards (1751) of a tortoise from
North Africa, more precisely from the old fort of Santa-Cruz (presently Djebel Murdjadjo or “pic de l’Aïdour” 35°42’N,
0°45’W, Oran, Algeria). Several hypotheses tried to explain Linnaeus’s choice of the name graeca, one proposing that
the species is the tortoise of the old Greek authors. Nevertheless, since 1758, the identity of Testudo graeca is rather well
established, with a precise type locality, although the pictured holotype is lost. Testudo graeca Linnaeus, 1758 is the type
species of Testudo Linnaeus, 1758 by subsequent designation of Bell (1828).
Johann Friedrich Gmelin gave the name Testudo hermanni to a tortoise of which he saw a specimen in Johann Her-
mann’s collection, in Strasbourg (Gmelin 1789). Johann Gottlob Schneider previously described the same specimen,
without naming it (Schneider 1783). The same tortoise was examined once again, by Johann David Schoepff, who
described and pictured it, but identified the species as Testudo graeca Linnaeus, 1758 (Schoepff 1792). Gmelin’s holo-
type is still extant in the Strasbourg University Zoological Museum (MZS 111) and the type locality was designated as
Collobrières, Var, France (Bour 1987).
Gabriel Bibron was the first to separate both species taxonomically, T. graeca and T. hermanni, but using an inappro-
priate nomenclature (Bibron and Bory 1833; Duméril and Bibron 1835), using T. graeca to name T. hermanni. For about
a century, the European tortoise (Testudo hermanni) was regularly named Testudo graeca, and the Testudo graeca of Lin-
naeus was most often named either Testudo mauritanica Duméril and Bibron, 1835, or Testudo ibera Pallas, 1814. The
mistake was found by Friedrich Siebenrock, who did not dare to change the nomenclature (Siebenrock 1913). That was
done by Stanley Flower, and Testudo hermanni was consistently used since its resurrection (Flower 1925; 1926).
Blasius Merrem wrote an ‘Essay on the systematics of amphibians’ (including reptiles), the publication of which was
certainly delayed: his most recent references were Geoffroy 1809 and Oppel 1811, but Schweigger 1812 was not men-
tioned (Merrem 1820). Within the genus Testudo, Merrem recognized four subgenera, without Testudo as a nominative
subgenus. Land tortoises were included in Chersine, new name (Chersos and Chersinos in Greek mean dry land and [tor-
toise] living on dry land, respectively). Merrem only included Testudo as a synonym of Chersine. Therefore, taxonomi-
cally speaking, Chersine Merrem, 1820 was clearly a nomen novum (replacement name) for Testudo Linnaeus, 1758.
Among the 18 nominal species incorporated into his subgenus Chersine, he put Testudo graeca. And, finally, among the
synonyms of Testudo (Chersine) graeca, Merrem included Testudo hermanni Gmelin, 1789 (Merrem 1820: 31, footnote
p). Following Merrem, Ferdinand von Ritgen (1828) used Chersine to include all land tortoises or ‘Ballentrotten’.
In his genus Testudo, Merrem also incorporated the new subgenus Matamata, with only Chelys Oppel, 1811 (an
unjustified emendation of Chelus Duméril, 1806) as synonym, and two included nominal species. Therefore, taxonomi-
cally speaking, Matamata Merrem, 1820 is clearly a nomen novum (replacement name) for Chelys Oppel, 1811. It was
recognized as such in the checklist of recent turtles and tortoises by Uwe Fritz and Peter Havaš (2007: 327).
John Edward Gray used the name Chersina to include only Testudo angulata Schweigger, 1812, first as subgenus of
, Gray stated that “Merrem gives the above
ting and typographical errors are plentiful in his papers. Further proof appears was evident when Gray wrote ‘Chersina
1. Second one according to the date of the forewords, ‘oct. 1830’ and ‘jan. 1831’, respectively; however, a reference to the second
with a corrective is given in the first one (G
, 1831a: 4). L
(1957) gave the second one as the original
reference for Chersina Gray.
CHERSINE MERREM, 1820 AND CHERSINA GRAY, 1831
(Gray 1831b: 9; 34).
If Chersina Gray, 1831 is recognized as a replacement name for (or an unjustified emendation of) Chersine Merrem,
1820, both “have the same type species, and type fixation for either applies also for the other, despite any statement to the
contrary”, according to the present Code of Zoological Nomenclature (Anonymous 1999: art. 67.8). Leopold Fitzinger
(1843) formally confirmed that Testudo angulata was the type species of Chersina Gray, but because this species was not
mentioned in Merrem’s work, Gray’s action cannot be considered as a subsequent designation of the type species of
Wassili Adolfovitch Lindholm was apparently the first to recognize that Gray ‘borrowed’ Merrem’s nominal genus,
and consequently erected a new name, Goniochersus, to include Testudo angulata (Lindholm 1929). Independently, John
Hewitt reached the same conclusion, creating the new genus Neotestudo (Hewitt 1931). Later, Hewitt used Chersine
(Hewitt 1937a-b) and Robert Mertens and Heinz Wermuth (1955) used Goniochersus, each for the same content. But
eventually, the nominal species Chersina angulata was reinstated by Arthur Loveridge and Ernst Williams (1957) – just
fifty years ago – and since regularly used, so its suppression would threaten the stability of nomenclature. Therefore,
despite the origin of the name as a lapsus, we recommend retaining Chersina as a valid name distinct from Chersine.
In the same paper Lindholm (1929) designated as type species (‘idiogenotyp’ = type by subsequent designation) of
Chersine Merrem, 1820 ‘Testudo graeca “L.” auct. = Testudo hermanni Gmelin’ (Lindholm 1929). Obviously, this desig-
nation was unforeseen, and certainly not in accordance with Merrem’s design when he created the genus, but this point is
outside of the nomenclatural rules. According to Loveridge and Williams (1957:254), Lindholm in this way designated
Testudo graeca Linnaeus, 1758 as type of Chersine Merrem, 1820. On the other hand, Wermuth and Mertens (1977: 20)
stated that Lindholm designated Testudo hermanni Gmelin, 1789 as the type species). France De Lapparent et al. (2006)
followed Loveridge and Williams, recognizing that Testudo graeca was the type species of Chersine, and erected the new
genus Eurotestudo to include Testudo hermanni and related species.
In a completely opposite treatment of the similar case involving the genus Matamata Merrem, 1820, Fritz and Havaš
(2007: 327) did not recognize that Chersine could be a nomen novum for Testudo. Rather, they argued that the designa-
tion of T. hermanni as type species of Chersine was nomenclaturally valid, because Lindholm emphasized that it was not
Testudo graeca of Linnaeus and because T. hermanni was included (as a synonym) in the original description of Chers-
ine. On the other hand, they rejected the taxonomic identity of the monophyletic lineage of Testudo hermanni proposed
by De Lapparent et al., and therefore overlooked the problem that had arisen as a result of the obvious similarities
between the names Chersine and Chersina.
Finally, in a paper dealing with captive husbandry and Testudo horfieldii, Wussow (1916) introduced Medaestia
(without formal diagnosis) as subgenus of Testudo, including T. horsfieldi Gray, 1844, T. graeca, T. ibera Pallas, 1814,
and T. leithi Günther, 1869. Mertens (1949) designated the type species of Medaestia Wussow 1916 as follows: “I fix it
here as graeca (hermanni Gmelin). Thus Medaestia (…) becomes a synonym of Testudo Linnaeus”. As outlined by
Mertens himself, Medaestia is a synonym of Testudo Linnaeus and has Testudo graeca Linnaeus as type-species. In any
case, T. hermanni was not included in the original description of Medaestia, and the ICZN Code is clear on that point
(art. 67.2): “A nominal species is only eligible to be fixed as the type species of a nominal genus or subgenus if it is an
originally included nominal species” (Anonymous 1999). Much later, Wermuth and Mertens (1977: 20) incorrectly
stated that Mertens designated Testudo hermanni Gmelin as the type species of Medaestia (an invalid designation, in any
case), contrary to the statement of Fritz and Havaš (2007).
Lapparent de Broin et al. (2006) recognized two clades in Testudo. We refrain from commenting here on the status
of these two clades, but will do so in a separate paper. However, if these clades should be named, the clade including
Testudo graeca Linnaeus, 1758 should bear the name Testudo Linnaeus, 1758 with Medaestia Wussow, 1916 as one of its
synonyms. For the clade which includes the species Testudo hermanni Gmelin, 1789, the name Chersine Merrem, 1820
is available and Eurotestudo De Lapparent, Bour, Parham and Perälä, 2006 should be considered its junior objective syn-
Anonymous (1999) International Code of Zoological Nomenclature. Fourth edition. ITZN, London 306 pp.
Bell, T. (1828) Characters of the order, families, and genera of the Testudinata. Zoological Journal, 3, 513–516.
Bibron, G. & Bory de Saint–Vincent, J.B.G.M. (1833) Reptiles. Pp. 57–76, in Geoffroy Saint–Hilaire, E., Geoffroy Saint–
Hilaire, I., Deshayes, G. P., Bibron, G. & Bory de Saint–Vincent, J. B. G. M., Travaux de la Section des Sciences Phy-
Zootaxa 1752 © 2008
siques 3 (1) (Zoologie – 1
section. Animaux vertébrés, Mollusques et Polypes), in Bory de Saint–Vincent, J. B. G.
M., Peytier, Puillon Boblaye, E., Servier, Brullé, A., Virlet, T., Geoffroy Saint–Hilaire, E., Geoffroy Saint–Hilaire,
I., Brongniart, A., Bibron, G., Deshayes, G. P., Guérin, F. E., Chaubard, L. A. & Fauché, Expédition scientifique de
Morée. F. G. Levrault, Paris – Strasbourg.
Bour, R. (1987) L’identité des Tortues terrestres européennes : spécimens-types et localités-types. Revue française
Duméril, A.M.C. & Bibron, G. (1835) Erpétologie générale ou histoire naturelle des Reptiles. Vol. 2. Roret, Paris 680 pp.
Fitzinger, L.J. (1843) Systema reptilium. Fasciculus primus. Amblyglossae (Conspectus geographicus). Braumüller &
Seidel Bibliopolas, Vindobonae 106 pp.
Flower, S.S. (1925) Contributions to our knowledge of the duration of life in vertebrate animals. III. Reptiles. Proceed-
ings of the Zoological Society of London, 1925, 911–981.
Flower, S.S. (1926) Species of land tortoises of the genus Testudo found in the countries bordering the Mediterranean
sea. Copeia, 1926, 133.
Fritz, U. & Havaš, P. (2007) Checklist of chelonians of the world. Vertebrate Zoology, 57, 2, 149–368.
Geoffroy Saint–Hilaire, É. (1809) Mémoire sur les tortues molles, nouveau genre sous le nom de Trionyx, et sur la forma-
tion des carapaces. Annales du Muséum d’Histoire naturelle, 14, 1–20.
Gmelin, J.F. (1789) Caroli a Linné Systema per Regna tria Naturae. Tomus I. Pars III. G. E. Beer, Lipsiae 484 pp.
Gray, J. E. (1831a) Synopsis of the Species of the Class Reptilia. Pp. 1–110, in E. Griffith & E. Pidgeon, The Class Rep-
Gray, J.E. (1831b) Synopsis Reptilium; or short descriptions of the species of Reptiles. Part I. Cataphracta. Tortoises,
Crocodiles, and Enaliosaurians. Treuttel, Wurtz, and Co., London 85 pp.
Hewitt, J. (1931) Description of some African tortoises. Annals of the Natal Museum, 6, 3, 461–506.
Hewitt, J. (1937a) A note on the relationship of the Cape genera of land–tortoises. South African Journal of Sciences, 33,
Lapparent de Broin, F. de, Bour, R., Parham, J.F. & Perälä, J. (2006) Eurotestudo, a new genus for the species Testudo
Lindholm, W.A. (1929) Revidiertes Verzeichnis der Gattungen der rezenten Schildkröten nebst Notizen zur Nomenklatur
einiger Arten. Zoologischer Anzeiger, 81, 2, 275–295.
Linnaeus, C. (1758) Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum charac-
Loveridge, A. & Williams, E.E. (1957) Revision of the African tortoises and turtles of the suborder Cryptodira. Bulletin
Merrem, B. (1820) Versuch eines Systems der Amphibien. Johann Christian Krieger, Marburg 191 pp.
Mertens, R. (1949) Medaestia Wussow, 1916, a synonym of Testudo Linnaeus, 1758. Copeia, 1949, 3, 232.
Oppel, M. (1811) Die Ordnungen, Familien und Gattungen der Reptilien als Prodrom einer Naturgeschichte derselben.
J. Lindauer, München 86 pp.
Ritgen, F.A.M.F. von (1828) Versuch einer natürlichen Eintheilung der Amphibien. Nova Acta Physico-Medica Aca-
Schneider, J.G. (1783) Allgemeine Naturgeschichte der Schildkröten, nebst einem systematischen Verzeichnisse der ein-
Schoepff, J.D. (1792–1801) Historia Testudinum Iconibus Illustrata. Ioannis Iacobi Palm, Erlangae 136 pp.
Schweigger, A.F. (1812) Prodromus monographiae cheloniorum. Königsberger Archiv für Naturwissenschaft und Math-
ematik, 1, 271–368, 406–462.
Siebenrock, F. (1913) Schildkröten aus Syrien und Mesopotamien. Annalen des Kaiserlich-Königliches Naturhis-
Wermuth, H. & Mertens, R. (1977) Liste der rezenten Amphibien und Reptilien. Testudines, Crocodylia, Rhynchocepha-
lia. Das Tierreich, 100, 1–174.
Wussow, W. (1916) Meine Erfahrungen mit Testudo horsfieldi. Wochenschrift für Aquarien–Terrarienkunde, 13, 169–