Eurotestudo hermanni Gmelin, 1789, Collobrières, France. Carapace, views: A , dorsal, B , ventral, C , posterior. Plastron, anterior lobe, D , dorsal view. Eurotestudo hermanni Gmelin, 1789, Collobrières, France. Carapace, vues: A , dorsale, B , ventrale, C , postérieure. Lobe antérieur du plastron, D , vue dorsale. 

Contexts in source publication

Context 1

... débute avec Paleotestudo canetotiana [16,19] par la tendance, plus complète que chez les autres espèces de Testudo s.l., à la fusion des trochanters du fémur, puis avec « T. » antiqua [32] par les tendances conjointes à la division externe de la su- pracaudale et à la fusion des suprapygales, enfin avec Eurotestudo n.g., où tous les caractères sont menés à leur terme [20]. Étude cladistique préalable . Une étude cladistique préalable, détaillée par ailleurs [20], inclut un nombre significatif de spécimens des espèces des lignées de Testudo s.s ., d ’ Agrionemys et d ’ hermanni (voir la version en anglais) et certains de ses possibles alliés [5,16], Paleotestudo canetotiana et Testudo promarginata . Les outgroups sont Manouria impressa, Indotestudo elongata et « Ergilemys » [7] (sensu [5]) bruneti . Tous par- tagent des caractères de Testudinidae terrestres [20]. Testudo s.l. partage des caractères avec Indotestudo et « Er. bruneti » et d ’ autres avec le seul « Er. bruneti ». Le genre Testudo s.l . est défini, les caractères partagés sont donnés ainsi que des particularités des formes de la lignée d ’ Eurotestudo n.g. [5,10,11,13,16 – 18,32]. Le point de séparation de Testudo s.s. et de la lignée Eurotestudo par rapport à Paleotestudo canetotiana [16] est examiné, ainsi que l ’ intégration de « T. » antiqua [32] dans la lignée. vant appuyer la séparation générique . Les caractères, mentionnés ci-dessus et retenus, sont examinés, ainsi que d ’ autres, à écarter de la diagnose, tels les tubercules des cuisses (apomorphie de Testudo s.s.), l ’ éperon caudal présent, mais variable, chez Agrionemys, Eurotestudo et T. kleinmanni et les modes de réduction de la main: réduction à quatre doigts ( Agrionemys ) ou seule- ment partielle chez Eurotestudo (ongle du 1 er et/ou du 5 e doigts, éventuellement réduits à absents) [1,3,4,9,23,24,27]. Les relations phylétiques des trois lignées par les différentes approches (morphologie des actuels ou/et des fossiles [9,23,24], analyse moléculaire [14,22, etc.]), ne peuvent être définies. Il apparaît que, suivant les taxons inclus et en fonction des méthodes utilisées, le groupe actuel hermanni ( hermanni seule ou avec boettgeri ) peut, soit être rapproché d ’ Agrionemys [9], mais aussi d ’ Indotestudo et d ’ autres taxons [14,22], soit être le groupe frère d ’ Agrionemys et de Testudo s.s. [24]. D ’ après l ’ étude sur laquelle est fondée la présente diagnose [20], les trois lignées sont bien séparées, après Manouria impressa, Indotestudo et « Er. » bruneti , en un groupe « Testudo s.l. ». Soit la lignée d ’ Eurotestudo n.g. est rapprochée de Testudo s.s., soit les trois lignées de Testudo s.l. sont en irrésolution, si l ’ on supprime le taxon asiatique fossile « T. » turgaica , moins bien connu. Testudo s.s. et Eurotestudo n.g. acquièrent un même mode de recourbement progressif du bourrelet épiplas- tral dorsal. Le caractère est constamment mené à son terme chez Testudo s.s. dès son apparition (présence d ’ une poche gulaire, recourbement jusqu ’ à l ’ entoplastron), moins souvent chez Eurotestudo n.g. Agrionemys présente le stade le moins avancé du processus évolutif et dans une conformation différente du lobe antérieur plastral (plus large avec bords latéraux plus convergents et entoplastron moins réduit). Il y a hétérochronie dans l ’ apparition des stades évolutifs de plusieurs caractères homoplasiques dans les deux groupes. Le patron de coloration commun de la carapace de type « Testudo » milite aussi en faveur de l ’ union de Testudo s.s. et Eurotestudo . En tout état de cause, le point de séparation de la lignée d ’ Eurotestudo n.g. par rapport aux formes asiatiques originelles remonte à une époque indétermi- née, mais antérieure à l ’ Oligocène. The principal aim of this work is to create a new genus Eurotestudo for the so-called hermanni group of testudinids, because it forms a distinct evolutionary lineage without an available name. Some valid names that seemed available for Testudo hermanni Gmelin, 1789 such as Chersine Merrem, 1820 and Medaestia Wussow, 1916, have Testudo graeca Linnaeus, 1758 as type species ([23], A. Rhodin in litt. to J.P.). Eurotestudo n.g. is part of Testudo s.l. (sensu [17,23,24]). This is a diverse group of terrestrial Palaearctic testudinids which, besides (a) Eu. hermanni and affiliated taxa [1 – 6,10,11,13,16,17,19,24,32] includes (b) the western hinged form Testudo Linnaeus, 1758, s.s., type species T. graeca , a genus extant in the southern-oriental Mediterranean Basin eastward to the Middle East [1,6,8,9, 15,17,24], and (c) Agrionemys Khozatsky & Mlynarski, 1966 [13], type species Testudo horsfieldii Gray, 1844, a western Central Asiatic extant genus, only represented in Europe as fossil (eastern part). Some recent studies [23 – 26] have elevated many subspecies to the rank of species within Agrionemys, Testudo s.s., and ‘ T. ’ hermanni Gmelin, 1789, while new species have also been described recently [29]. T. hermanni (osteological Fig. 1) was separated from T. boettgeri Mojsisovics, 1889 (osteological Fig. in [9] as T. hermanni ), and the name T. hercegovinensis Werner, 1899 was resurrected for a population previously attributed to boettgeri [28] (not included in the analysis in [20]). The taxon of upper rank (according to the ICZN) to unite the extant and fossil species in the hermanni complex is a genus, necessary in accordance with previous opinions [16,17, 23,24], that agree with the various hypotheses about phylogenetic relationships among the three groups [14,17,22,24]. Examination of fossil lineages, into which we can integrate the extant species, shows that there is a clear separation of the three groups, each one inclusive of a succession of valid species: the separation occurred, at least, since the Upper Miocene, but probably the Oligocene. It is the date of the appearance of the oldest attested Testudo s.s., Testudo marmorum Gaudry, 1862 (Greece). In Africa, Testudo s.s. is definitely known from the Pliocene (Morocco). However, Testudo ( ‘ s.l. ’ ) semenensis Bergounioux, 1955, from the Upper Miocene (Tunisia), may be attributable to Testudo s.s [15,20]. Agrionemys is firstly known from the Upper Miocene of the Republic of Moldova, by ‘ Testudo ’ bessarabica Riabinin 1915 [30] ( Protestudo Chkhikvadze, 1970) [6], and of Afghanistan ([17] and Rage & Lapparent de Broin, in prep.]. Agrionemys might be related to ‘ Testudo ’ turgaica Riabinin, 1926 [31], from the ‘ Middle ’ Miocene of Khazakstan as well as to other Asiatic or eastern European forms [8,34]. The stem lineage of Eurotestudo n.g. is identified in the ‘ Middle ’ Miocene with the appearance of Paleotestudo canetotiana (Lartet, 1851), France [5,16] and in the Upper Miocene with ‘ Testudo ’ antiqua Bronn, 1831, Germany) [32]. Other older extinct western European species, such as ‘ Testudo ’ promarginata Rein- ach, 1900, from the Lower Miocene (Germany, France) [5], may also be on the stem of Eurotestudo n.g. However, ‘ T ’ . promarginata may predate the split between Eurotestudo n.g. and Testudo s.s. [20]. The diagnosis of the extant group hermanni has al- ready been established on the basis of the morphological study of osteology [5,6,9,10,18,19,23,24] and external characters such as horny appendices, coloration and scales [1 – 4,24,26]. The present diagnosis of Eurotestudo n.g. is principally based on characters of the carapace, preserved in the fossils: plates and scute outlines as well as proportions, features which no doubt charac- terize the whole genus. Many specimens of the fossil species and extant populations of the lineages of Testudo, Agrionemys and Eurotestudo n.g. have been exam- ined. The diagnosis of each lineage includes some un- ambiguous characters and various homoplastic characters: their appearance in the lineages is asynchro- nous and their variability has been established for each population. Order Chelonii Brongniart (Latreille), 1800 Superfamily Testudinoidea Batsch, 1788 Family Testudinidae Batsch, 1788 Infrafamily Testudininei Batsch, 1788 Eurotestudo new genus Etymology : from ‘ Europe ’ , the continent of biogeo- graphic origin, and ‘ Testudo ’ Type species : Testudo hermanni Gmelin, 1789, type locality: Collobrières, Var, France Named valid species (sensu ICZN) : The Eurotestudo ’ hermanni group: extant Eu. hermanni (Fig. 1) and boettgeri from which ‘ T. ’ hercegovinensis Werner, 1899 may be disassociated [28], and the fossil Eu. pyrenaica (Depéret & Donnezan, 1890), Pliocene of Per- pignan (MN 15), Eu. globosa (Portis, 1890), Plio-Pleis- tocene boundary, Le Ville, Upper Valdarno, Eu. lunellensis (Almera & Bofill, 1903), ‘ Middle ’ Pleistocene of Caverna de Gràcia, and Eu. szalai Mlynarski, 1955, Pliocene of Weze (MN 15). The extant species plus Eu. pyrenaica and Eu. lunellensis are or can be (respectively) well diagnosed. Eu. globosa (one male specimen in Le Ville, thick bones; referred fragments in other localities from Valdarno) may be a junior sy- nonym of Eu. hermanni. Eu. szalai (some fragments) cannot be sufficiently diagnosed [1,4,5,10 – 12,21]. Material referred to the genus : all the specimens in the literature referred to T. hermanni and Testudo sp. from the Quaternary of Europe, which present the characters of Eu. hermanni exposed in the present diagnosis: in particular the populations of T. hermanni from the Quaternary of France, especially the populations from l ’ Escale (ca 0.6 Myr) and Lunel-Viel (ca 0.3 to 0.34 Myr), and ‘ Testudo cf. hermanni ’ from Soave (Zoppega 2, Italy) (early Middle Pleistocene) [10,11, 33]. The unnamed Lunel-Viel and Soave populations certainly represent distinct diagnosable species. The ‘ hermanni lineage ’ initially includes Paleotestudo canetotiana [16] by the trend towards the fusion of the trochanters, more complete than in Agrionemys and Testudo s.s. [19], then ‘ T. ’ antiqua [32] by the common trend towards an external division of the supracaudal and fusion of the suprapygals and ...

Context 2

... évolutifs de plusieurs caractères homoplasiques dans les deux groupes. Le patron de coloration commun de la carapace de type « Testudo » milite aussi en faveur de l ’ union de Testudo s.s. et Eurotestudo . En tout état de cause, le point de séparation de la lignée d ’ Eurotestudo n.g. par rapport aux formes asiatiques originelles remonte à une époque indétermi- née, mais antérieure à l ’ Oligocène. The principal aim of this work is to create a new genus Eurotestudo for the so-called hermanni group of testudinids, because it forms a distinct evolutionary lineage without an available name. Some valid names that seemed available for Testudo hermanni Gmelin, 1789 such as Chersine Merrem, 1820 and Medaestia Wussow, 1916, have Testudo graeca Linnaeus, 1758 as type species ([23], A. Rhodin in litt. to J.P.). Eurotestudo n.g. is part of Testudo s.l. (sensu [17,23,24]). This is a diverse group of terrestrial Palaearctic testudinids which, besides (a) Eu. hermanni and affiliated taxa [1 – 6,10,11,13,16,17,19,24,32] includes (b) the western hinged form Testudo Linnaeus, 1758, s.s., type species T. graeca , a genus extant in the southern-oriental Mediterranean Basin eastward to the Middle East [1,6,8,9, 15,17,24], and (c) Agrionemys Khozatsky & Mlynarski, 1966 [13], type species Testudo horsfieldii Gray, 1844, a western Central Asiatic extant genus, only represented in Europe as fossil (eastern part). Some recent studies [23 – 26] have elevated many subspecies to the rank of species within Agrionemys, Testudo s.s., and ‘ T. ’ hermanni Gmelin, 1789, while new species have also been described recently [29]. T. hermanni (osteological Fig. 1) was separated from T. boettgeri Mojsisovics, 1889 (osteological Fig. in [9] as T. hermanni ), and the name T. hercegovinensis Werner, 1899 was resurrected for a population previously attributed to boettgeri [28] (not included in the analysis in [20]). The taxon of upper rank (according to the ICZN) to unite the extant and fossil species in the hermanni complex is a genus, necessary in accordance with previous opinions [16,17, 23,24], that agree with the various hypotheses about phylogenetic relationships among the three groups [14,17,22,24]. Examination of fossil lineages, into which we can integrate the extant species, shows that there is a clear separation of the three groups, each one inclusive of a succession of valid species: the separation occurred, at least, since the Upper Miocene, but probably the Oligocene. It is the date of the appearance of the oldest attested Testudo s.s., Testudo marmorum Gaudry, 1862 (Greece). In Africa, Testudo s.s. is definitely known from the Pliocene (Morocco). However, Testudo ( ‘ s.l. ’ ) semenensis Bergounioux, 1955, from the Upper Miocene (Tunisia), may be attributable to Testudo s.s [15,20]. Agrionemys is firstly known from the Upper Miocene of the Republic of Moldova, by ‘ Testudo ’ bessarabica Riabinin 1915 [30] ( Protestudo Chkhikvadze, 1970) [6], and of Afghanistan ([17] and Rage & Lapparent de Broin, in prep.]. Agrionemys might be related to ‘ Testudo ’ turgaica Riabinin, 1926 [31], from the ‘ Middle ’ Miocene of Khazakstan as well as to other Asiatic or eastern European forms [8,34]. The stem lineage of Eurotestudo n.g. is identified in the ‘ Middle ’ Miocene with the appearance of Paleotestudo canetotiana (Lartet, 1851), France [5,16] and in the Upper Miocene with ‘ Testudo ’ antiqua Bronn, 1831, Germany) [32]. Other older extinct western European species, such as ‘ Testudo ’ promarginata Rein- ach, 1900, from the Lower Miocene (Germany, France) [5], may also be on the stem of Eurotestudo n.g. However, ‘ T ’ . promarginata may predate the split between Eurotestudo n.g. and Testudo s.s. [20]. The diagnosis of the extant group hermanni has al- ready been established on the basis of the morphological study of osteology [5,6,9,10,18,19,23,24] and external characters such as horny appendices, coloration and scales [1 – 4,24,26]. The present diagnosis of Eurotestudo n.g. is principally based on characters of the carapace, preserved in the fossils: plates and scute outlines as well as proportions, features which no doubt charac- terize the whole genus. Many specimens of the fossil species and extant populations of the lineages of Testudo, Agrionemys and Eurotestudo n.g. have been exam- ined. The diagnosis of each lineage includes some un- ambiguous characters and various homoplastic characters: their appearance in the lineages is asynchro- nous and their variability has been established for each population. Order Chelonii Brongniart (Latreille), 1800 Superfamily Testudinoidea Batsch, 1788 Family Testudinidae Batsch, 1788 Infrafamily Testudininei Batsch, 1788 Eurotestudo new genus Etymology : from ‘ Europe ’ , the continent of biogeo- graphic origin, and ‘ Testudo ’ Type species : Testudo hermanni Gmelin, 1789, type locality: Collobrières, Var, France Named valid species (sensu ICZN) : The Eurotestudo ’ hermanni group: extant Eu. hermanni (Fig. 1) and boettgeri from which ‘ T. ’ hercegovinensis Werner, 1899 may be disassociated [28], and the fossil Eu. pyrenaica (Depéret & Donnezan, 1890), Pliocene of Per- pignan (MN 15), Eu. globosa (Portis, 1890), Plio-Pleis- tocene boundary, Le Ville, Upper Valdarno, Eu. lunellensis (Almera & Bofill, 1903), ‘ Middle ’ Pleistocene of Caverna de Gràcia, and Eu. szalai Mlynarski, 1955, Pliocene of Weze (MN 15). The extant species plus Eu. pyrenaica and Eu. lunellensis are or can be (respectively) well diagnosed. Eu. globosa (one male specimen in Le Ville, thick bones; referred fragments in other localities from Valdarno) may be a junior sy- nonym of Eu. hermanni. Eu. szalai (some fragments) cannot be sufficiently diagnosed [1,4,5,10 – 12,21]. Material referred to the genus : all the specimens in the literature referred to T. hermanni and Testudo sp. from the Quaternary of Europe, which present the characters of Eu. hermanni exposed in the present diagnosis: in particular the populations of T. hermanni from the Quaternary of France, especially the populations from l ’ Escale (ca 0.6 Myr) and Lunel-Viel (ca 0.3 to 0.34 Myr), and ‘ Testudo cf. hermanni ’ from Soave (Zoppega 2, Italy) (early Middle Pleistocene) [10,11, 33]. The unnamed Lunel-Viel and Soave populations certainly represent distinct diagnosable species. The ‘ hermanni lineage ’ initially includes Paleotestudo canetotiana [16] by the trend towards the fusion of the trochanters, more complete than in Agrionemys and Testudo s.s. [19], then ‘ T. ’ antiqua [32] by the common trend towards an external division of the supracaudal and fusion of the suprapygals and finally Eurotestudo n.g. [20] where these characters are the best realized. Eurotestudo n.g. is diagnosed by the obligatory com- bination of the following characters: (1) narrowed vertebral series, narrower than the costal series as a whole (in all populations; an apomorphic character); (2) fusion of the suprapygals into a trapezoid with a straight posterior border: the fusion varies from occasional in fossils (but often incompletely preserved) to most often present in extant populations (a rare homoplasy in Testudo s.s. and Agrionemys ); (3) the quadrangular pygal becomes hexagonal with small latero-anterior sides (often present in all populations, a rare homoplasy in Testudo s.s. and Agrionemys ), and which are sometimes covered by the 11th marginals; (4) tendency to having a divided supracaudal, externally and eventually internally; frequency of inner division of the supracaudal varies from rare to frequent depending on population (present in ‘ Ergilemys ’ , but presumably not by the same evolutionary process, see [20]); external division of the supracaudal occasional to constant, according to population, constant in hermanni and in the majority of cases in boettgeri (a rare homoplasy in Testudo s.s. and extant Agrionemys and in some other Testudininei such as Pyxis ); (5) ventral surface of the gulars, making a posteriorly pointed triangle, frequently ventrally in relief relative to horizontal plan, often present in all populations (a homoplasy in extant Agrionemys ); there is often a medial anterior bend between the gulars (a homoplasy in Testudo s.s. and in extant Agrionemys ) (Fig. 1). silized, possibly generic . Synapomorphies for the extant species (unique among the Testudinidae) are: (1) the small scales on the outer area of the front face of the forearm (in addition to the large and regular ones): irregular antero-distal area of small scales in Eu. hermanni , all smaller and very numerous in Eu. boettgeri [3]; (2) the fragmented, almost indistinct frontal scale; (3) the color pattern of the plastron with two parasagittal dark bands, each one whole or broken up [1,3,4]. A cladistic analysis, previously performed and de- tailed elsewhere [20], includes, in the ingroup the species: Testudo turgaica, Agrionemys bessarabica, A. horsfieldii, A. kazachstanica Chkhikvadze, 1988; T. marmorum , T. marginata Schoepff, 1793, T. weissingeri Bour, 1995, T. antakyensis Perälä, 1996, T. kenitrensis Gmira 1993, T. graeca (s.l.) from the Maghreb, T. promarginata, Paleotestudo canetotiana, T. antiqua and the hermanni group (above mentioned valid species), and three outgroup taxa of terrestrial testudinids: Manouria impressa (Günther, 1882), Indotestudo elongata (Blyth, 1853) and ‘ Er. ’ bruneti Broin, 1977, a species attributed to the genus Ergilemys Chkikvadze, 1972 [7] sensu [5], Oligocene, La Millo- que, France. Many Miocene and Oligocene fossil species, all insufficiently known, although potentially belonging to the lineage of Eurotestudo n.g., were disregarded. Among them, some Oligo-Miocene frag- mentary specimens from France, attributed to ‘ Ergilemys ’ sp., have a hinge similar to that of Testudo s.s. only [5 (pl. 25, 28)], even in relatively young adults. The relationships of these specimens with ‘ Er. ’ bruneti and the Eurotestudo n.g. lineage are unclear. All the above taxa are ...

Context 3

... identified in the ‘ Middle ’ Miocene with the appearance of Paleotestudo canetotiana (Lartet, 1851), France [5,16] and in the Upper Miocene with ‘ Testudo ’ antiqua Bronn, 1831, Germany) [32]. Other older extinct western European species, such as ‘ Testudo ’ promarginata Rein- ach, 1900, from the Lower Miocene (Germany, France) [5], may also be on the stem of Eurotestudo n.g. However, ‘ T ’ . promarginata may predate the split between Eurotestudo n.g. and Testudo s.s. [20]. The diagnosis of the extant group hermanni has al- ready been established on the basis of the morphological study of osteology [5,6,9,10,18,19,23,24] and external characters such as horny appendices, coloration and scales [1 – 4,24,26]. The present diagnosis of Eurotestudo n.g. is principally based on characters of the carapace, preserved in the fossils: plates and scute outlines as well as proportions, features which no doubt charac- terize the whole genus. Many specimens of the fossil species and extant populations of the lineages of Testudo, Agrionemys and Eurotestudo n.g. have been exam- ined. The diagnosis of each lineage includes some un- ambiguous characters and various homoplastic characters: their appearance in the lineages is asynchro- nous and their variability has been established for each population. Order Chelonii Brongniart (Latreille), 1800 Superfamily Testudinoidea Batsch, 1788 Family Testudinidae Batsch, 1788 Infrafamily Testudininei Batsch, 1788 Eurotestudo new genus Etymology : from ‘ Europe ’ , the continent of biogeo- graphic origin, and ‘ Testudo ’ Type species : Testudo hermanni Gmelin, 1789, type locality: Collobrières, Var, France Named valid species (sensu ICZN) : The Eurotestudo ’ hermanni group: extant Eu. hermanni (Fig. 1) and boettgeri from which ‘ T. ’ hercegovinensis Werner, 1899 may be disassociated [28], and the fossil Eu. pyrenaica (Depéret & Donnezan, 1890), Pliocene of Per- pignan (MN 15), Eu. globosa (Portis, 1890), Plio-Pleis- tocene boundary, Le Ville, Upper Valdarno, Eu. lunellensis (Almera & Bofill, 1903), ‘ Middle ’ Pleistocene of Caverna de Gràcia, and Eu. szalai Mlynarski, 1955, Pliocene of Weze (MN 15). The extant species plus Eu. pyrenaica and Eu. lunellensis are or can be (respectively) well diagnosed. Eu. globosa (one male specimen in Le Ville, thick bones; referred fragments in other localities from Valdarno) may be a junior sy- nonym of Eu. hermanni. Eu. szalai (some fragments) cannot be sufficiently diagnosed [1,4,5,10 – 12,21]. Material referred to the genus : all the specimens in the literature referred to T. hermanni and Testudo sp. from the Quaternary of Europe, which present the characters of Eu. hermanni exposed in the present diagnosis: in particular the populations of T. hermanni from the Quaternary of France, especially the populations from l ’ Escale (ca 0.6 Myr) and Lunel-Viel (ca 0.3 to 0.34 Myr), and ‘ Testudo cf. hermanni ’ from Soave (Zoppega 2, Italy) (early Middle Pleistocene) [10,11, 33]. The unnamed Lunel-Viel and Soave populations certainly represent distinct diagnosable species. The ‘ hermanni lineage ’ initially includes Paleotestudo canetotiana [16] by the trend towards the fusion of the trochanters, more complete than in Agrionemys and Testudo s.s. [19], then ‘ T. ’ antiqua [32] by the common trend towards an external division of the supracaudal and fusion of the suprapygals and finally Eurotestudo n.g. [20] where these characters are the best realized. Eurotestudo n.g. is diagnosed by the obligatory com- bination of the following characters: (1) narrowed vertebral series, narrower than the costal series as a whole (in all populations; an apomorphic character); (2) fusion of the suprapygals into a trapezoid with a straight posterior border: the fusion varies from occasional in fossils (but often incompletely preserved) to most often present in extant populations (a rare homoplasy in Testudo s.s. and Agrionemys ); (3) the quadrangular pygal becomes hexagonal with small latero-anterior sides (often present in all populations, a rare homoplasy in Testudo s.s. and Agrionemys ), and which are sometimes covered by the 11th marginals; (4) tendency to having a divided supracaudal, externally and eventually internally; frequency of inner division of the supracaudal varies from rare to frequent depending on population (present in ‘ Ergilemys ’ , but presumably not by the same evolutionary process, see [20]); external division of the supracaudal occasional to constant, according to population, constant in hermanni and in the majority of cases in boettgeri (a rare homoplasy in Testudo s.s. and extant Agrionemys and in some other Testudininei such as Pyxis ); (5) ventral surface of the gulars, making a posteriorly pointed triangle, frequently ventrally in relief relative to horizontal plan, often present in all populations (a homoplasy in extant Agrionemys ); there is often a medial anterior bend between the gulars (a homoplasy in Testudo s.s. and in extant Agrionemys ) (Fig. 1). silized, possibly generic . Synapomorphies for the extant species (unique among the Testudinidae) are: (1) the small scales on the outer area of the front face of the forearm (in addition to the large and regular ones): irregular antero-distal area of small scales in Eu. hermanni , all smaller and very numerous in Eu. boettgeri [3]; (2) the fragmented, almost indistinct frontal scale; (3) the color pattern of the plastron with two parasagittal dark bands, each one whole or broken up [1,3,4]. A cladistic analysis, previously performed and de- tailed elsewhere [20], includes, in the ingroup the species: Testudo turgaica, Agrionemys bessarabica, A. horsfieldii, A. kazachstanica Chkhikvadze, 1988; T. marmorum , T. marginata Schoepff, 1793, T. weissingeri Bour, 1995, T. antakyensis Perälä, 1996, T. kenitrensis Gmira 1993, T. graeca (s.l.) from the Maghreb, T. promarginata, Paleotestudo canetotiana, T. antiqua and the hermanni group (above mentioned valid species), and three outgroup taxa of terrestrial testudinids: Manouria impressa (Günther, 1882), Indotestudo elongata (Blyth, 1853) and ‘ Er. ’ bruneti Broin, 1977, a species attributed to the genus Ergilemys Chkikvadze, 1972 [7] sensu [5], Oligocene, La Millo- que, France. Many Miocene and Oligocene fossil species, all insufficiently known, although potentially belonging to the lineage of Eurotestudo n.g., were disregarded. Among them, some Oligo-Miocene frag- mentary specimens from France, attributed to ‘ Ergilemys ’ sp., have a hinge similar to that of Testudo s.s. only [5 (pl. 25, 28)], even in relatively young adults. The relationships of these specimens with ‘ Er. ’ bruneti and the Eurotestudo n.g. lineage are unclear. All the above taxa are Testudininei by characters given in the analysis [20]. The characters of the clades are present in some other Testudininei. Out of the 18 characters of the analysis, Testudo s.l. shares with Indotestudo and ‘ Er. ’ bruneti: (1) the coincidence of the costal-marginal scute sulci and the pleur- al-peripheral sutures and (2) the fusion of the two 12th marginals into a supracaudal. With ‘ Er. bruneti ’ , Testudo s.l. shares the shell form: more elevated than that found in M. impressa , with elevated peripherals and marginals, arched with domed lateral pleural slopes and with two anterior and posterior slopes meeting at a domed more or less flattened part, and more or less strong protuberances below the vertebrals 2 or 3 or 4 and eventually the costals; basically quadrangular and moderately wide, not looking narrow or round; posterior border moderately postero-laterally expanded in dorsal view. In Agrionemys , the shell is rounded and shortened at the level of the bridge, with an elevated bridge and more convergent plastral lobes and a larger entoplastron. In the T. marginata group, the shell becomes elongated (much postero-laterally expanded at the peripheral border), differently from I. elongata (pos- tero-medially expanded). In the antiqua group, the shell widens. In Paleotestudo, the posterior border is not at all expanded. As a member of Testudo s.l., the Eurotestudo n.g. lineage shares with Testudo s.s. and Agrionemys particularly the following characters: (1) the posteriorly as- cending dorsal epiplastral lip with a slightly convex surface: stopping its ascension abruptly (a) and, being more or less curved (b), located above the posterior surface of the epiplastron which is not thickened (c) – elements (a), (b), and (c) differentiate these chelonians from Indotestudo (Fig. 1D, figures in [5,9,20]) – ; (2) the typically sinuous sulcus between the abdominal and femoral, with the latero-anterior sinuosity clearly extended on the hypoplastron and anterior to the ingu- inal notch (Fig. 1B); however, in Testudo s.s., the curve tends to be reduced, with the presence of the hypo-xi- phiplastral hinge, particularly in the marginata group; in Agrionemys the hypoplastral overlap by the femorals is apparently more extensive, partly because the hypoplastron is shortened; (3) the possible posterior reduction of the series of eight neurals (Fig. 1A) to 7 or 6; this character is very rare in Eurotestudo n.g., but it is the norm in extant Testudo species (in time after the fossil species T. kenitrensis and T. marmorum and some fossil T. graeca from Morocco), and in the extant Agrionemys species (evolving after the fossil species bessarabica ); (4) the ‘ Testudo s.l. ’ type of suprapygal – pygal, as opposed to the ‘ geoemydine ’ (in Manouria impressa ) and ‘ Geochelone ’ (in Indotestudo and ‘ Er. ’ bruneti ) types: both suprapygals constitute one trapezoid structure, with straight borders, in front of the pygal (Fig. 1C), that is completely elongated throughout its width and not only laterally as in the ‘ Geochelone ’ type (see [5,9,18] and other references included); consequently, the posterior border of the vertebral 5 is con- fluent with the limits of the suprapygal – pygal structure (complete coincidence of sutures ...

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... nouveau genre est créé pour le groupe d espèces hermanni , en le séparant de Testudo s.s. [20]. Il existe trois lignées distinctes, séparées au sein de Testudo s.l. (sensu [17,23,24]), Testudinidae terrestre d ’ origine paléarctique : (a) Testudo s.s., connu en Europe [1,24] depuis le Miocène supérieur au moins [17], et en Afri- que sûrement depuis le Pliocène, mais probablement dès le Miocène [15]; (b) Agrionemys [12], connu en Afghanistan et en Moldavie au Miocène supérieur [8,17,30], lié ou non à des formes orientales plus an- ciennes dont « T. » turgaica du Miocène « moyen » [17,31,34] ; (c) la lignée d ’ Eurotestudo n.g., représentée en Europe au plus tard depuis le Miocène supérieur et moyen [5,16,32] de France et d ’ Allemagne et probablement dès l ’ Oligocène et actuellement représentée en Europe au moins par « T. » hermanni Gmelin, 1789 et « T. » boettgeri Mojsisovics, 1889 [1 – 3,28]. Des formes du Miocène inférieur [5] peuvent se situer, soit dans cette lignée, soit dans une lignée commune avec celle de Testudo s.s., suivant leur point de séparation [20]. La définition du nouveau genre proposée ici est fondée sur des caractères ostéologiques [5,6,9,10,18,19,23,24], grâce à une nouvelle analyse cladistique incluant des espèces fossiles [5,7,16,30 – 32] et actuelles de Testudo s.l., ce qui a permis de polariser les caractères et de faire la part des homoplasies. Certains caractères externes des espèces actuelles peuvent participer à la diagnose [1 – 4,9,23,24,27]. Eurotestudo n.g. Espèce type: Testudo hermanni Gmelin, 1789 Espèces valides incluses Le groupe hermanni: Eu. hermanni (Fig. 1), Eu. boettgeri (dont Eu . hercegovinensis tend à être séparé [28]), Eu. pyrenaica, Eu. globosa, Eu. lunellensis, Eu. szalai . Les espèces actuelles et les fossiles Eu. pyrenaica et Eu. lunellensis , sont ou peuvent être bien définies. Eu. globosa (un seul spécimen, mâle, os épais) se présente comme un représentant de Eu. hermanni. Eu. szalai (fragments isolés) n ’ est pas assez préservé pour être diagnostiqué [1 – 3,5,11,13,21]. Eurotestudo n.g. est caractérisé par la nécessaire combinaison de (1) la série des vertébrales étrécie ; (2) la fusion des suprapygales en un trapèze à bord postérieur rectiligne (ou l ’ état de la plus forte tendance à la fusion précédant celui de celle-ci) ; (3) la pygale quadrangulaire tendant à devenir hexagonale, à petits côtés antérieurs (alors souvent chevauchés par les mar- ginales 11) ; (4) la tendance à la division de la supra- caudale, externe (souvent) et interne (moins souvent) ; (5) la surface ventrale des gulaires formant un triangle dirigé postérieurement et souvent saillant ventralement, souvent avec inflexion médiane antérieure du bord, saillant à l ’ avant, des gulaires. Les caractères 2 à 5 sont indépendamment homoplasiques chez Agrionemys et/ ou Testudo s.s., rarement, et jamais tous ensemble (Fig. 1A, B, C, D). Caractères externes additifs, non fossilisés (synapomorphies des espèces actuelles potentiellement génériques) : (1) sur la face frontale du bras, distale par rapport aux grandes écailles, une aire antéro-distale limitée, avec, soit des écailles petites et irrégulières ( Eu. hermanni ), soit uniquement de nombreuses écailles très petites ( Eu. boettgeri) , alors que toutes les écailles sont grandes et régulières chez les autres espèces de Testudo s.l. ; (2) écaille frontale fragmentée, presque indistincte ; (3) patron de coloration du plastron avec deux bandes foncées parasagittales, entières ou fragmentées [1,3,4]. Matériel référé au genre: tous les spécimens référés à T. hermanni et à Testudo sp. du Quaternaire d ’ Europe ayant les caractères donnés dans la diagnose et notam- ment ceux de l ’ Escale, Lunel-Viel [10,11] et Soave [33]. Les populations de Lunel-Viel et de Soave repré- sentent de bonnes espèces, suffisamment préservées pour être diagnostiquées. La lignée hermanni : elle débute avec Paleotestudo canetotiana [16,19] par la tendance, plus complète que chez les autres espèces de Testudo s.l., à la fusion des trochanters du fémur, puis avec « T. » antiqua [32] par les tendances conjointes à la division externe de la su- pracaudale et à la fusion des suprapygales, enfin avec Eurotestudo n.g., où tous les caractères sont menés à leur terme [20]. Étude cladistique préalable . Une étude cladistique préalable, détaillée par ailleurs [20], inclut un nombre significatif de spécimens des espèces des lignées de Testudo s.s ., d ’ Agrionemys et d ’ hermanni (voir la version en anglais) et certains de ses possibles alliés [5,16], Paleotestudo canetotiana et Testudo promarginata . Les outgroups sont Manouria impressa, Indotestudo elongata et « Ergilemys » [7] (sensu [5]) bruneti . Tous par- tagent des caractères de Testudinidae terrestres [20]. Testudo s.l. partage des caractères avec Indotestudo et « Er. bruneti » et d ’ autres avec le seul « Er. bruneti ». Le genre Testudo s.l . est défini, les caractères partagés sont donnés ainsi que des particularités des formes de la lignée d ’ Eurotestudo n.g. [5,10,11,13,16 – 18,32]. Le point de séparation de Testudo s.s. et de la lignée Eurotestudo par rapport à Paleotestudo canetotiana [16] est examiné, ainsi que l ’ intégration de « T. » antiqua [32] dans la lignée. vant appuyer la séparation générique . Les caractères, mentionnés ci-dessus et retenus, sont examinés, ainsi que d ’ autres, à écarter de la diagnose, tels les tubercules des cuisses (apomorphie de Testudo s.s.), l ’ éperon caudal présent, mais variable, chez Agrionemys, Eurotestudo et T. kleinmanni et les modes de réduction de la main: réduction à quatre doigts ( Agrionemys ) ou seule- ment partielle chez Eurotestudo (ongle du 1 er et/ou du 5 e doigts, éventuellement réduits à absents) [1,3,4,9,23,24,27]. Les relations phylétiques des trois lignées par les différentes approches (morphologie des actuels ou/et des fossiles [9,23,24], analyse moléculaire [14,22, etc.]), ne peuvent être définies. Il apparaît que, suivant les taxons inclus et en fonction des méthodes utilisées, le groupe actuel hermanni ( hermanni seule ou avec boettgeri ) peut, soit être rapproché d ’ Agrionemys [9], mais aussi d ’ Indotestudo et d ’ autres taxons [14,22], soit être le groupe frère d ’ Agrionemys et de Testudo s.s. [24]. D ’ après l ’ étude sur laquelle est fondée la présente diagnose [20], les trois lignées sont bien séparées, après Manouria impressa, Indotestudo et « Er. » bruneti , en un groupe « Testudo s.l. ». Soit la lignée d ’ Eurotestudo n.g. est rapprochée de Testudo s.s., soit les trois lignées de Testudo s.l. sont en irrésolution, si l ’ on ...

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... (voir la version anglaise) Eurotestudo n.g. Espèce type: Testudo hermanni Gmelin, 1789 Espèces valides incluses Le groupe hermanni: Eu. hermanni ( Fig. 1), Eu. boettgeri (dont Eu. hercegovinensis tend à être séparé [28]), Eu. pyrenaica, Eu. globosa, Eu. lunellensis, Eu. szalai. Les espèces actuelles et les fossiles Eu. pyrenaica et Eu. lunellensis, sont ou peuvent être bien définies. Eu. globosa (un seul spécimen, mâle, os épais) se pré-sente comme un représentant de Eu. hermanni. Eu. ...

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... (moins souvent) ; (5) la surface ventrale des gulaires formant un triangle dirigé postérieurement et souvent saillant ventralement, souvent avec inflexion médiane antérieure du bord, saillant à l'avant, des gulaires. Les caractères 2 à 5 sont indépendamment homoplasiques chez Agrionemys et/ ou Testudo s.s., rarement, et jamais tous ensemble (Fig. 1A, B, C, ...

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... Gray, 1844, a western Central Asiatic extant genus, only represented in Europe as fossil (eastern part). Some recent studies [23][24][25][26] have elevated many subspecies to the rank of species within Agrionemys, Testudo s.s., and 'T.' hermanni Gmelin, 1789, while new species have also been described recently [29]. T. hermanni (osteological Fig. 1) was separated from T. boettgeri Mojsisovics, 1889 (osteological Fig. in [9] as T. hermanni), and the name T. hercegovinensis Werner, 1899 was resurrected for a population previously attributed to boettgeri [28] (not included in the analysis in [20]). The taxon of upper rank (according to the ICZN) to unite the extant and fossil ...

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... valid species (sensu ICZN): The 'Eurotestudo' hermanni group: extant Eu. hermanni ( Fig. 1) and boettgeri from which 'T.' hercegovinensis Werner, 1899 may be disassociated [28], and the fossil Eu. pyrenaica (Depéret & Donnezan, 1890), Pliocene of Perpignan (MN 15), Eu. globosa (Portis, 1890), Plio-Pleistocene boundary, Le Ville, Upper Valdarno, Eu. lunellensis (Almera & Bofill, 1903), 'Middle' Pleistocene of Caverna de ...

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... Testudininei such as Pyxis); (5) ventral surface of the gulars, making a posteriorly pointed triangle, frequently ventrally in relief relative to horizontal plan, often present in all populations (a homoplasy in extant Agrionemys); there is often a medial anterior bend between the gulars (a homoplasy in Testudo s.s. and in extant Agrionemys) (Fig. ...

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... the following characters: (1) the posteriorly ascending dorsal epiplastral lip with a slightly convex surface: stopping its ascension abruptly (a) and, being more or less curved (b), located above the posterior surface of the epiplastron which is not thickened (c) -elements (a), (b), and (c) differentiate these chelonians from Indotestudo (Fig. 1D, figures in [5,9,20]) -; (2) the typically sinuous sulcus between the abdominal and femoral, with the latero-anterior sinuosity clearly extended on the hypoplastron and anterior to the inguinal notch (Fig. 1B); however, in Testudo s.s., the curve tends to be reduced, with the presence of the hypo-xiphiplastral hinge, particularly in the marginata ...

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... posterior surface of the epiplastron which is not thickened (c) -elements (a), (b), and (c) differentiate these chelonians from Indotestudo (Fig. 1D, figures in [5,9,20]) -; (2) the typically sinuous sulcus between the abdominal and femoral, with the latero-anterior sinuosity clearly extended on the hypoplastron and anterior to the inguinal notch (Fig. 1B); however, in Testudo s.s., the curve tends to be reduced, with the presence of the hypo-xiphiplastral hinge, particularly in the marginata group; in Agrionemys the hypoplastral overlap by the femorals is apparently more extensive, partly because the hypoplastron is shortened; (3) the possible posterior reduction of the series of eight ...

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... 1B); however, in Testudo s.s., the curve tends to be reduced, with the presence of the hypo-xiphiplastral hinge, particularly in the marginata group; in Agrionemys the hypoplastral overlap by the femorals is apparently more extensive, partly because the hypoplastron is shortened; (3) the possible posterior reduction of the series of eight neurals (Fig. 1A) to 7 or 6; this character is very rare in Eurotestudo n.g., but it is the norm in extant Testudo species (in time after the fossil species T. kenitrensis and T. marmorum and some fossil T. graeca from Morocco), and in the extant Agrionemys species (evolving after the fossil species bessarabica); (4) the 'Testudo s.l.' type of ...

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... and in the extant Agrionemys species (evolving after the fossil species bessarabica); (4) the 'Testudo s.l.' type of suprapygal-pygal, as opposed to the 'geoemydine' (in Manouria impressa) and 'Geochelone' (in Indotestudo and 'Er.' bruneti) types: both suprapygals constitute one trapezoid structure, with straight borders, in front of the pygal (Fig. 1C), that is completely elongated throughout its width and not only laterally as in the 'Geochelone' type (see [5,9,18] and other references included); consequently, the posterior border of the vertebral 5 is confluent with the limits of the suprapygal-pygal structure (complete coincidence of sutures and sulci); however, in extant ...