C. R. Palevol 5 (2006) 803 811
http://france.elsevier.com/direct/PALEVO/
Systematic Paleontology (Vertebrate Paleontology)
Eurotestudo, a new genus for the species
Testudo hermanni Gmelin, 1789 (Chelonii, Testudinidae)
a, b c d
France de Lapparent de Broin *, Roger Bour , James F. Parham , Jarmo Per�l�
a
� Pal�obiodiversit� et pal�oenvironnements , d�partement � Histoire de la Terre du Mus�um national d histoire naturelle,
UMS 203, UMR 5143 du CNRS, CP 38, 8, rue Buffon, 75231 Paris cedex 05, France
b
Laboratoire des reptiles et amphibiens, d�partement � syst�matique et �volution du
Mus�um national d histoire naturelle, UMS 602, CP 30, 25 rue Cuvier, 75231 Paris cedex 05, France
c
Museum of Paleontology, University of California, Berkeley, CA, 94720, USA/ Department of Evolutionary Genomics,
DOE Joint Genome Institute and Lawrence Berkeley National Laboratory, 2800 Mitchell Drive, Walnut Creek, CA, 94598, USA
d
Department of Biological and Environmental Sciences, PO Box 65 (Biocenter 3, Viikinkaari 1), FIN-00014 university of Helsinki, Finland
Received 18 November 2005; accepted after revision 21 February 2006
Available online 02 May 2006
Presented by Philippe Taquet
Abstract
The new genus is created to include the species of the hermanni group, which is within Testudo s.l., a Palearctic genus,
consequently separated from both Testudo s.s. and Agrionemys. A preliminary cladistic analysis of the osteological characters,
including fossil species, demonstrated the splitting of the three lineages, probably since the Oligocene and surely at the Upper
Miocene. Diagnosis of the new genus is based on a collection of features. The main stages of evolution leading to the three
lineages are provided. We also describe external characters of the extant species that could be considered as diagnostic. However,
phylogenetic relationships between genera are not definitively established. To cite this article: F. de Lapparent de Broin et al.,
C. R. Palevol 5 (2006).
� 2006 Acad�mie des sciences. Published by Elsevier SAS. All rights reserved.
R�sum�
Eurotestudo, nouveau genre pour l espŁce Testudo hermanni Gmelin, 1789 (Chelonii, Testudinidae). Ce nouveau genre est
cr�� pour le groupe d espŁces hermanni, le s�parant de Testudo s.s. et d Agrionemys au sein de Testudo s.l., genre pal�arctique.
Une analyse cladistique des caractŁres ost�ologiques, men�e au pr�alable et incluant des espŁces fossiles, a montr� la s�paration
des trois lign�es, probablement depuis l OligocŁne et s�rement le MiocŁne sup�rieur. La diagnose du nouveau genre est �tablie sur
une conjonction de caractŁres. Les principales �tapes de l �volution menant aux trois genres sont donn�es. Les caractŁres externes
des espŁces actuelles pouvant participer ą la diagnose sont examin�s. Les relations phyl�tiques entre les genres ne sont pas �tablies
d�finitivement. Pour citer cet article : F. de Lapparent de Broin et al., C. R. Palevol 5 (2006).
� 2006 Acad�mie des sciences. Published by Elsevier SAS. All rights reserved.
Keywords: Turtles; Eurotestudo n.g.; Testudo s.l.; Testudinidae; Europe; Tertiary Extant
Mots cl�s : Tortues ; Eurotestudo n.g. ; Testudo s.l. ; Testudinidae ; Europe ; Tertiaire Actuel
*
Auteur correspondant.
E-mail address: fdelap@mnhn.fr (F. de Lapparent de Broin).
1631-0683/$ - see front matter � 2006 Acad�mie des sciences. Published by Elsevier SAS. All rights reserved.
doi:10.1016/j.crpv.2006.03.002
804 F. de Lapparent de Broin et al. / C. R. Palevol 5 (2006) 803 811
Version fran�aise abr�g�e (2) la fusion des suprapygales en un trapŁze ą bord
post�rieur rectiligne (ou l �tat de la plus forte tendance
Introduction ą la fusion pr�c�dant celui de celle-ci) ; (3) la pygale
quadrangulaire tendant ą devenir hexagonale, ą petits
Un nouveau genre est cr�� pour le groupe d espŁces
c�t�s ant�rieurs (alors souvent chevauch�s par les mar-
hermanni, en le s�parant de Testudo s.s. [20]. Il existe
ginales 11) ; (4) la tendance ą la division de la supra-
trois lign�es distinctes, s�par�es au sein de Testudo s.l.
caudale, externe (souvent) et interne (moins souvent) ;
(sensu [17,23,24]), Testudinidae terrestre d origine pa- (5) la surface ventrale des gulaires formant un triangle
l�arctique : (a) Testudo s.s., connu en Europe [1,24]
dirig� post�rieurement et souvent saillant ventralement,
depuis le MiocŁne sup�rieur au moins [17], et en Afri- souvent avec inflexion m�diane ant�rieure du bord, sail-
que s�rement depuis le PliocŁne, mais probablement
lant ą l avant, des gulaires. Les caractŁres 2 ą 5 sont
dŁs le MiocŁne [15]; (b) Agrionemys [12], connu en
ind�pendamment homoplasiques chez Agrionemys et/
Afghanistan et en Moldavie au MiocŁne sup�rieur
ou Testudo s.s., rarement, et jamais tous ensemble
[8,17,30], li� ou non ą des formes orientales plus an- (Fig. 1A, B, C, D).
ciennes dont � T. turgaica du MiocŁne � moyen
CaractŁres externes additifs, non fossilis�s (synapo-
[17,31,34] ; (c) la lign�e d Eurotestudo n.g., repr�sent�e morphies des espŁces actuelles potentiellement g�n�ri-
en Europe au plus tard depuis le MiocŁne sup�rieur et ques) : (1) sur la face frontale du bras, distale par rap-
moyen [5,16,32] de France et d Allemagne et probable- port aux grandes �cailles, une aire ant�ro-distale
ment dŁs l OligocŁne et actuellement repr�sent�e en limit�e, avec, soit des �cailles petites et irr�guliŁres
Europe au moins par � T. hermanni Gmelin, 1789 et (Eu. hermanni), soit uniquement de nombreuses
� T. boettgeri Mojsisovics, 1889 [1 3,28]. Des formes �cailles trŁs petites (Eu. boettgeri), alors que toutes les
du MiocŁne inf�rieur [5] peuvent se situer, soit dans �cailles sont grandes et r�guliŁres chez les autres
cette lign�e, soit dans une lign�e commune avec celle espŁces de Testudo s.l. ; (2) �caille frontale fragment�e,
de Testudo s.s., suivant leur point de s�paration [20]. La presque indistincte ; (3) patron de coloration du plastron
d�finition du nouveau genre propos�e ici est fond�e sur avec deux bandes fonc�es parasagittales, entiŁres ou
des caractŁres ost�ologiques [5,6,9,10,18,19,23,24], fragment�es [1,3,4].
gr�ce ą une nouvelle analyse cladistique incluant des Mat�riel r�f�r� au genre: tous les sp�cimens r�f�r�s
espŁces fossiles [5,7,16,30 32] et actuelles de Testudo ą T. hermanni et ą Testudo sp. du Quaternaire d Europe
s.l., ce qui a permis de polariser les caractŁres et de faire ayant les caractŁres donn�s dans la diagnose et notam-
la part des homoplasies. Certains caractŁres externes ment ceux de l Escale, Lunel-Viel [10,11] et Soave
des espŁces actuelles peuvent participer ą la diagnose [33]. Les populations de Lunel-Viel et de Soave repr�-
[1 4,9,23,24,27]. sentent de bonnes espŁces, suffisamment pr�serv�es
pour ętre diagnostiqu�es.
Syst�matique (voir la version anglaise) La lign�e hermanni : elle d�bute avec Paleotestudo
canetotiana [16,19] par la tendance, plus complŁte que
Eurotestudo n.g.
chez les autres espŁces de Testudo s.l., ą la fusion des
EspŁce type: Testudo hermanni Gmelin, 1789
trochanters du f�mur, puis avec � T. antiqua [32] par
EspŁces valides incluses
les tendances conjointes ą la division externe de la su-
Le groupe hermanni: Eu. hermanni (Fig. 1), Eu.
pracaudale et ą la fusion des suprapygales, enfin avec
boettgeri (dont Eu. hercegovinensis tend ą ętre s�par�
Eurotestudo n.g., oł tous les caractŁres sont men�s ą
[28]), Eu. pyrenaica, Eu. globosa, Eu. lunellensis, Eu.
leur terme [20].
szalai. Les espŁces actuelles et les fossiles Eu. pyrenai-
ca et Eu. lunellensis, sont ou peuvent ętre bien d�finies. Comparaisons morphologiques
Eu. globosa (un seul sp�cimen, m�le, os �pais) se pr�-
sente comme un repr�sentant de Eu. hermanni. Eu. sza- �tude cladistique pr�alable. Une �tude cladistique
pr�alable, d�taill�e par ailleurs [20], inclut un nombre
lai (fragments isol�s) n est pas assez pr�serv� pour ętre
significatif de sp�cimens des espŁces des lign�es de
diagnostiqu� [1 3,5,11,13,21].
Testudo s.s., d Agrionemys et d hermanni (voir la ver-
Diagnose du genre sion en anglais) et certains de ses possibles alli�s [5,16],
Paleotestudo canetotiana et Testudo promarginata. Les
Eurotestudo n.g. est caract�ris� par la n�cessaire outgroups sont Manouria impressa, Indotestudo elon-
combinaison de (1) la s�rie des vert�brales �tr�cie ; gata et � Ergilemys [7] (sensu [5]) bruneti. Tous par-
F. de Lapparent de Broin et al. / C. R. Palevol 5 (2006) 803 811 805
tagent des caractŁres de Testudinidae terrestres [20]. coloration commun de la carapace de type � Testudo
Testudo s.l. partage des caractŁres avec Indotestudo et milite aussi en faveur de l union de Testudo s.s. et Euro-
� Er. bruneti et d autres avec le seul � Er. bruneti . testudo. En tout �tat de cause, le point de s�paration de
Le genre Testudo s.l. est d�fini, les caractŁres partag�s la lign�e d Eurotestudo n.g. par rapport aux formes
sont donn�s ainsi que des particularit�s des formes de la asiatiques originelles remonte ą une �poque ind�termi-
lign�e d Eurotestudo n.g. [5,10,11,13,16 18,32]. Le n�e, mais ant�rieure ą l OligocŁne.
point de s�paration de Testudo s.s. et de la lign�e Euro-
1. Introduction
testudo par rapport ą Paleotestudo canetotiana [16] est
examin�, ainsi que l int�gration de � T. antiqua [32]
The principal aim of this work is to create a new
dans la lign�e.
CaractŁres externes des repr�sentants actuels pou- genus Eurotestudo for the so-called hermanni group
of testudinids, because it forms a distinct evolutionary
vant appuyer la s�paration g�n�rique. Les caractŁres,
lineage without an available name. Some valid names
mentionn�s ci-dessus et retenus, sont examin�s, ainsi
that seemed available for Testudo hermanni Gmelin,
que d autres, ą �carter de la diagnose, tels les tubercules
des cuisses (apomorphie de Testudo s.s.), l �peron cau- 1789 such as Chersine Merrem, 1820 and Medaestia
dal pr�sent, mais variable, chez Agrionemys, Eurotestu- Wussow, 1916, have Testudo graeca Linnaeus, 1758
as type species ([23], A. Rhodin in litt. to J.P.). Euro-
do et T. kleinmanni et les modes de r�duction de la
main: r�duction ą quatre doigts (Agrionemys) ou seule- testudo n.g. is part of Testudo s.l. (sensu [17,23,24]).
This is a diverse group of terrestrial Palaearctic testudi-
ment partielle chez Eurotestudo (ongle du 1er et/ou du
nids which, besides (a) Eu. hermanni and affiliated taxa
5e doigts, �ventuellement r�duits ą absents)
[1 6,10,11,13,16,17,19,24,32] includes (b) the western
[1,3,4,9,23,24,27].
hinged form Testudo Linnaeus, 1758, s.s., type species
T. graeca, a genus extant in the southern-oriental Med-
Discussion et conclusion
iterranean Basin eastward to the Middle East [1,6,8,9,
Les relations phyl�tiques des trois lign�es par les 15,17,24], and (c) Agrionemys Khozatsky & Mlynarski,
diff�rentes approches (morphologie des actuels ou/et 1966 [13], type species Testudo horsfieldii Gray, 1844,
des fossiles [9,23,24], analyse mol�culaire [14,22, a western Central Asiatic extant genus, only represented
etc.]), ne peuvent ętre d�finies. Il appara�t que, suivant in Europe as fossil (eastern part). Some recent studies
les taxons inclus et en fonction des m�thodes utilis�es, [23 26] have elevated many subspecies to the rank of
le groupe actuel hermanni (hermanni seule ou avec species within Agrionemys, Testudo s.s., and  T. her-
boettgeri) peut, soit ętre rapproch� d Agrionemys [9], manni Gmelin, 1789, while new species have also been
mais aussi d Indotestudo et d autres taxons [14,22], soit described recently [29]. T. hermanni (osteological
ętre le groupe frŁre d Agrionemys et de Testudo s.s. Fig. 1) was separated from T. boettgeri Mojsisovics,
[24]. D aprŁs l �tude sur laquelle est fond�e la pr�sente 1889 (osteological Fig. in [9] as T. hermanni), and the
diagnose [20], les trois lign�es sont bien s�par�es, aprŁs name T. hercegovinensis Werner, 1899 was resurrected
Manouria impressa, Indotestudo et �Er. bruneti, en for a population previously attributed to boettgeri [28]
un groupe � Testudo s.l. . Soit la lign�e d Eurotestudo (not included in the analysis in [20]). The taxon of
n.g. est rapproch�e de Testudo s.s., soit les trois lign�es upper rank (according to the ICZN) to unite the extant
de Testudo s.l. sont en irr�solution, si l on supprime le and fossil species in the hermanni complex is a genus,
taxon asiatique fossile � T. turgaica, moins bien con- necessary in accordance with previous opinions [16,17,
nu. Testudo s.s. et Eurotestudo n.g. acquiŁrent un męme 23,24], that agree with the various hypotheses about
mode de recourbement progressif du bourrelet �piplas- phylogenetic relationships among the three groups
tral dorsal. Le caractŁre est constamment men� ą son [14,17,22,24]. Examination of fossil lineages, into
terme chez Testudo s.s. dŁs son apparition (pr�sence which we can integrate the extant species, shows that
d une poche gulaire, recourbement jusqu ą l entoplas- there is a clear separation of the three groups, each one
tron), moins souvent chez Eurotestudo n.g. Agrionemys inclusive of a succession of valid species: the separation
pr�sente le stade le moins avanc� du processus �volutif occurred, at least, since the Upper Miocene, but prob-
et dans une conformation diff�rente du lobe ant�rieur ably the Oligocene. It is the date of the appearance of
plastral (plus large avec bords lat�raux plus convergents the oldest attested Testudo s.s., Testudo marmorum
et entoplastron moins r�duit). Il y a h�t�rochronie dans Gaudry, 1862 (Greece). In Africa, Testudo s.s. is defi-
l apparition des stades �volutifs de plusieurs caractŁres nitely known from the Pliocene (Morocco). However,
homoplasiques dans les deux groupes. Le patron de Testudo ( s.l. ) semenensis Bergounioux, 1955, from
806 F. de Lapparent de Broin et al. / C. R. Palevol 5 (2006) 803 811
Fig. 1. Eurotestudo hermanni Gmelin, 1789, CollobriŁres, France. Carapace, views: A, dorsal, B, ventral, C, posterior. Plastron, anterior lobe, D,
dorsal view.
Fig. 1. Eurotestudo hermanni Gmelin, 1789, CollobriŁres, France. Carapace, vues: A, dorsale, B, ventrale, C, post�rieure. Lobe ant�rieur du
plastron, D, vue dorsale.
the Upper Miocene (Tunisia), may be attributable to ined. The diagnosis of each lineage includes some un-
Testudo s.s [15,20]. Agrionemys is firstly known from ambiguous characters and various homoplastic
the Upper Miocene of the Republic of Moldova, by characters: their appearance in the lineages is asynchro-
nous and their variability has been established for each
 Testudo bessarabica Riabinin 1915 [30] (Protestudo
population.
Chkhikvadze, 1970) [6], and of Afghanistan ([17] and
Rage & Lapparent de Broin, in prep.]. Agrionemys
2. Systematics
might be related to  Testudo turgaica Riabinin, 1926
[31], from the  Middle Miocene of Khazakstan as well
Order Chelonii Brongniart (Latreille), 1800
as to other Asiatic or eastern European forms [8,34].
Superfamily Testudinoidea Batsch, 1788
The stem lineage of Eurotestudo n.g. is identified in
Family Testudinidae Batsch, 1788
the  Middle Miocene with the appearance of Paleotes-
Infrafamily Testudininei Batsch, 1788
tudo canetotiana (Lartet, 1851), France [5,16] and in
Eurotestudo new genus
the Upper Miocene with  Testudo antiqua Bronn,
Etymology: from  Europe , the continent of biogeo-
1831, Germany) [32]. Other older extinct western Eur-
graphic origin, and  Testudo
opean species, such as  Testudo promarginata Rein-
Type species: Testudo hermanni Gmelin, 1789, type
ach, 1900, from the Lower Miocene (Germany, France)
locality: CollobriŁres, Var, France
[5], may also be on the stem of Eurotestudo n.g. How-
ever,  T . promarginata may predate the split between
2.1. Included species
Eurotestudo n.g. and Testudo s.s. [20].
The diagnosis of the extant group hermanni has al-
Named valid species (sensu ICZN): The  Eurotestu-
ready been established on the basis of the morphologi-
do hermanni group: extant Eu. hermanni (Fig. 1) and
cal study of osteology [5,6,9,10,18,19,23,24] and exter-
boettgeri from which  T. hercegovinensis Werner,
nal characters such as horny appendices, coloration and
1899 may be disassociated [28], and the fossil Eu. pyr-
scales [1 4,24,26]. The present diagnosis of Eurotestu-
enaica (Dep�ret & Donnezan, 1890), Pliocene of Per-
do n.g. is principally based on characters of the cara- pignan (MN 15), Eu. globosa (Portis, 1890), Plio-Pleis-
pace, preserved in the fossils: plates and scute outlines
tocene boundary, Le Ville, Upper Valdarno, Eu.
as well as proportions, features which no doubt charac- lunellensis (Almera & Bofill, 1903),  Middle Pleisto-
terize the whole genus. Many specimens of the fossil cene of Caverna de Grącia, and Eu. szalai Mlynarski,
species and extant populations of the lineages of Testu- 1955, Pliocene of Weze (MN 15). The extant species
do, Agrionemys and Eurotestudo n.g. have been exam- plus Eu. pyrenaica and Eu. lunellensis are or can be
F. de Lapparent de Broin et al. / C. R. Palevol 5 (2006) 803 811 807
(respectively) well diagnosed. Eu. globosa (one male populations (a homoplasy in extant Agrionemys); there
specimen in Le Ville, thick bones; referred fragments is often a medial anterior bend between the gulars (a
in other localities from Valdarno) may be a junior sy- homoplasy in Testudo s.s. and in extant Agrionemys)
nonym of Eu. hermanni. Eu. szalai (some fragments) (Fig. 1).
cannot be sufficiently diagnosed [1,4,5,10 12,21].
Additional characters: external characters, not fos-
Material referred to the genus: all the specimens in silized, possibly generic. Synapomorphies for the extant
the literature referred to T. hermanni and Testudo sp. species (unique among the Testudinidae) are: (1) the
from the Quaternary of Europe, which present the char- small scales on the outer area of the front face of the
acters of Eu. hermanni exposed in the present diagno- forearm (in addition to the large and regular ones): irre-
sis: in particular the populations of T. hermanni from gular antero-distal area of small scales in Eu. hermanni,
the Quaternary of France, especially the populations all smaller and very numerous in Eu. boettgeri [3]; (2)
from l Escale (ca 0.6 Myr) and Lunel-Viel (ca 0.3 to the fragmented, almost indistinct frontal scale; (3) the
0.34 Myr), and  Testudo cf. hermanni from Soave color pattern of the plastron with two parasagittal dark
(Zoppega 2, Italy) (early Middle Pleistocene) [10,11, bands, each one whole or broken up [1,3,4].
33]. The unnamed Lunel-Viel and Soave populations
certainly represent distinct diagnosable species.
2.3. Morphological comparisons
The  hermanni lineage initially includes Paleotestu-
do canetotiana [16] by the trend towards the fusion of
A cladistic analysis, previously performed and de-
the trochanters, more complete than in Agrionemys and
tailed elsewhere [20], includes, in the ingroup the spe-
Testudo s.s. [19], then  T. antiqua [32] by the common
cies: Testudo turgaica, Agrionemys bessarabica,
trend towards an external division of the supracaudal
A. horsfieldii, A. kazachstanica Chkhikvadze, 1988;
and fusion of the suprapygals and finally Eurotestudo
T. marmorum, T. marginata Schoepff, 1793,
n.g. [20] where these characters are the best realized.
T. weissingeri Bour, 1995, T. antakyensis Per�l�, 1996,
T. kenitrensis Gmira 1993, T. graeca (s.l.) from the
2.2. Diagnosis
Maghreb, T. promarginata, Paleotestudo canetotiana,
T. antiqua and the hermanni group (above mentioned
Eurotestudo n.g. is diagnosed by the obligatory com- valid species), and three outgroup taxa of terrestrial tes-
bination of the following characters: (1) narrowed ver- tudinids: Manouria impressa (G�nther, 1882), Indotes-
tudo elongata (Blyth, 1853) and  Er. bruneti Broin,
tebral series, narrower than the costal series as a whole
1977, a species attributed to the genus Ergilemys
(in all populations; an apomorphic character); (2) fusion
of the suprapygals into a trapezoid with a straight pos- Chkikvadze, 1972 [7] sensu [5], Oligocene, La Millo-
terior border: the fusion varies from occasional in fos- que, France. Many Miocene and Oligocene fossil spe-
cies, all insufficiently known, although potentially be-
sils (but often incompletely preserved) to most often
present in extant populations (a rare homoplasy in Tes- longing to the lineage of Eurotestudo n.g., were
disregarded. Among them, some Oligo-Miocene frag-
tudo s.s. and Agrionemys); (3) the quadrangular pygal
becomes hexagonal with small latero-anterior sides (of- mentary specimens from France, attributed to  Ergil-
ten present in all populations, a rare homoplasy in Tes- emys sp., have a hinge similar to that of Testudo s.s.
only [5 (pl. 25, 28)], even in relatively young adults.
tudo s.s. and Agrionemys), and which are sometimes
The relationships of these specimens with  Er. bruneti
covered by the 11th marginals; (4) tendency to having
a divided supracaudal, externally and eventually intern- and the Eurotestudo n.g. lineage are unclear. All the
ally; frequency of inner division of the supracaudal var- above taxa are Testudininei by characters given in the
ies from rare to frequent depending on population (pre- analysis [20]. The characters of the clades are present in
some other Testudininei.
sent in  Ergilemys , but presumably not by the same
evolutionary process, see [20]); external division of Out of the 18 characters of the analysis, Testudo s.l.
the supracaudal occasional to constant, according to po- shares with Indotestudo and  Er. bruneti: (1) the coin-
pulation, constant in hermanni and in the majority of cidence of the costal-marginal scute sulci and the pleur-
cases in boettgeri (a rare homoplasy in Testudo s.s. al-peripheral sutures and (2) the fusion of the two 12th
and extant Agrionemys and in some other Testudininei marginals into a supracaudal. With  Er. bruneti , Testu-
such as Pyxis); (5) ventral surface of the gulars, making do s.l. shares the shell form: more elevated than that
a posteriorly pointed triangle, frequently ventrally in found in M. impressa, with elevated peripherals and
relief relative to horizontal plan, often present in all marginals, arched with domed lateral pleural slopes
808 F. de Lapparent de Broin et al. / C. R. Palevol 5 (2006) 803 811
and with two anterior and posterior slopes meeting at a tero-medially so that the supracaudal slightly covers the
domed more or less flattened part, and more or less suprapygal (particularly in A. kazachstanica) with a si-
strong protuberances below the vertebrals 2 or 3 or 4 nuosity [9] and the vertebral 5 may overlap the pygal,
and eventually the costals; basically quadrangular and often in Eurotestudo, sometimes in Testudo; (5) the nar-
moderately wide, not looking narrow or round; poster- rowing of the lateral scute border on the dorsal epiplas-
ior border moderately postero-laterally expanded in tron.
dorsal view. In Agrionemys, the shell is rounded and
In Testudo s.l., the suprapygals (two in general) are
shortened at the level of the bridge, with an elevated
divided by a semicircular (primitively) or a semicircu-
bridge and more convergent plastral lobes and a larger
lar semitransversal, or a transverse line, according to
entoplastron. In the T. marginata group, the shell be- the following evolutionary stages; the most derived
comes elongated (much postero-laterally expanded at
stage is the fusion of the suprapygals into a trapezoid.
the peripheral border), differently from I. elongata (pos- The fusion of the suprapygals into a trapezoid with a
tero-medially expanded). In the antiqua group, the shell
posterior straight border is mostly known in Eurotestu-
widens. In Paleotestudo, the posterior border is not at
do n.g., although it also occurs rarely in some species of
all expanded.
Testudo and Agrionemys. The three genera evolved, in
As a member of Testudo s.l., the Eurotestudo n.g. parallel, the following homoplastic characters (that are
lineage shares with Testudo s.s. and Agrionemys parti- in general very frequently witnessed in Testudininei):
cularly the following characters: (1) the posteriorly as- (1) the partial to complete reduction of the dorsal cervi-
cending dorsal epiplastral lip with a slightly convex sur- cal (constantly or occasionally present in a population);
face: stopping its ascension abruptly (a) and, being (2) a tendency for the pectorals to extend medioanter-
more or less curved (b), located above the posterior sur- iorly toward the entoplastron and onto the entoplastron
face of the epiplastron which is not thickened (c)  ele- (without meeting each other anteromedially), more or
less frequently according to population, and not only
ments (a), (b), and (c) differentiate these chelonians
in Agrionemys and Eurotestudo n.g. (particularly in
from Indotestudo (Fig. 1D, figures in [5,9,20])  ;
the boettgeri, Lunel-Viel and Soave populations), from
(2) the typically sinuous sulcus between the abdominal
which taxa this character is well known, but also in
and femoral, with the latero-anterior sinuosity clearly
extended on the hypoplastron and anterior to the ingu- T. graeca [9].
inal notch (Fig. 1B); however, in Testudo s.s., the curve With Testudo, Eurotestudo n.g. shares an epiplastral
tends to be reduced, with the presence of the hypo-xi- lip that curves onto the entoplastron, overhanging the
phiplastral hinge, particularly in the marginata group; dorsal surface. Below this, there is a depressed gular
in Agrionemys the hypoplastral overlap by the femorals pocket. A tendency toward a gular pocket is obvious
is apparently more extensive, partly because the hypo- in Eurotestudo n.g.: a narrow and weak gular pocket
plastron is shortened; (3) the possible posterior reduc- is particularly found in the Lunel-Viel population, and
tion of the series of eight neurals (Fig. 1A) to 7 or 6; one is often present in P. canetotiana. P. canetotiana
this character is very rare in Eurotestudo n.g., but it is (figures in [5,16]) is considered as belonging to the
the norm in extant Testudo species (in time after the Eurotestudo n.g. lineage despite its similarity with Tes-
fossil species T. kenitrensis and T. marmorum and some tudo. The differentiating conditions are the acquisition
fossil T. graeca from Morocco), and in the extant in Testudo of a characteristic hinge, in both sexes, be-
Agrionemys species (evolving after the fossil species tween the hypo-and xiphiplastra, with (a) a correlative
bessarabica); (4) the  Testudo s.l. type of suprapy- elongation of the posterior lobe, (b) the fusion of the
gal pygal, as opposed to the  geoemydine (in Manour- lateral extremities of the suture (at the hinge) and of
ia impressa) and  Geochelone (in Indotestudo and  Er. the abdomino-femoral sulcus (except in juveniles and
bruneti) types: both suprapygals constitute one trape- in the small-sized T. kenitrensis), and (c) the tendency
zoid structure, with straight borders, in front of the py- to shorten the femorals on the hypoplastron (particu-
gal (Fig. 1C), that is completely elongated throughout larly in the marginata group). In Testudo, the gular
its width and not only laterally as in the  Geochelone pocket is constant, small to strong [6,9,23,24], except
type (see [5,9,18] and other references included); con- in T. antakyensis Per�l�, 1996 (the lip is often not even
sequently, the posterior border of the vertebral 5 is con- curved; Fig. in [9] as T. terrestris Forssk�l, 1775). In
fluent with the limits of the suprapygal pygal structure Agrionemys (figure in [9]) (unknown in A. bessarabica),
(complete coincidence of sutures and sulci); however, the epiplastral lip is never curved up to entoplastron, as
in extant Agrionemys, vertebral 5 is slightly shorter pos- in fossils of the Eurotestudo n.g. lineage ( T. antiqua,
F. de Lapparent de Broin et al. / C. R. Palevol 5 (2006) 803 811 809
Eu. pyrenaica, figures in [5,32]) and the epiplastral en- do not participate in the lateral borders; presently the
toplastral surface is rarely depressed into a gular pocket. gulars occupy either only the complete anterior border
On the other hand, the ventral surface of the gulars in of the lobe, or a narrow slice (protruding or not) in its
relief can also be found in Agrionemys, but never in medial part, the humeral lateral borders being rounded.
Testudo. This morphology is also present in Testudo s.s. (always)
and in A. kazachstanica and some A. horsfieldii, but not
The process of modification of the suprapygal area,
in  T. turgaica and A. bessarabica. However, in
with the tendency to complete fusion of the plates, is
Agrionemys, the anterior lobe always has more conver-
most achieved in Eurotestudo n.g., making it unique
ging lateral borders. T. antiqua belongs more confi-
among Palaearctic forms (present in some African small
dently to the Eurotestudo n.g. lineage: tendency to fea-
endemics [18]). Meanwhile, the process of fusion of the
ture a divided supracaudal, possibility for having fused
last neurals is more achieved in the two other genera,
suprapygals, general aspect of the shell which is of a
although homoplastically because the fusion progres-
quadrangular form, posteriorly expanded according to
sively develops in each lineage separately. In the line-
the norm in Eurotestudo n.g. contrarily to Paleotestudo.
age of Eurotestudo n.g., there is no single species accu-
The epiplastral lip is wide and long, never curved up to
mulating all the more derived states of the homoplasies.
the entoplaston and there is no gular pocket. The shell
P. canetotiana has a higher tendency towards a gular
is particularly wide (width/length). Its femur is unde-
pocket. The species have their particularities:  T. anti-
scribed, and the possible fusion of the suprapygals
qua has a wider shell [32];  Eu. pyrenaica has a trian-
needs to be confirmed. A revision of  T. antiqua (as
gular or trapezoid notch at the nuchal, not affecting the
for some close fossil species) is necessary to reconsider
adjacent peripherals and the cervical is completely lack-
its phylogenetic position with respect to its possible in-
ing, as in the Upper Miocene  T. amberiacensis Dep�r-
tegration into the genus [20].
et, 1894, France [5], which might belong to a pyrenaica
group if it is confirmed that it belongs to Eurotestudo.
The Eu. aff. hermanni populations from Lunel-Viel and 2.4. External characters indicating generic status
Soave are more derived by the progression of the pec-
torals on the entoplastron. The Soave population and
The extant species in the new genus Eurotestudo are
some elements from the Quaternary of the Iberian Pe-
unique among the Testudinidae by the following char-
ninsula (references in [11,19,20] have the most devel-
acters: (1) the scalation of the front face of the forearm
oped epiplastral lip, thick and often very protruding, but
includes a distal area of small and irregular (small
lacking a gular pocket. The extant Eu. hermanni is most
scales in Eu. hermanni, very small and numerous in
advanced by having a trapezoid suprapygal and the
Eu. boettgeri), while there are only large and regular
more consistently divided supracaudal (externally and
scales in other Testudinidae; (2) the frontal scale is
internally), and perhaps also by featuring the very occa- fragmented, almost indistinct, while most tortoises have
sional presence of seven neurals (the series is not well
a large and well-delimited frontal, following two elon-
enough known in other populations to make compari- gated prefrontals; (3) the color pattern of the plastron:
sons). Eu. boettgeri has the femorals much shortened.
from the basic pattern of postero-lateral dark spots, ori-
Concerning fossil relatives of Eurotestudo n.g., Pa- ginating from the areolar zone (basically radiated), de-
leotestudo canetotiana is considered as belonging to the velops a system of parasagittal dark bands that is un-
Eurotestudo lineage by the more advanced fusion of the known among other chelonians. Another external
femoral trochanters and its full aspect; some specimens character supports the separation of the new genus
have the gulars in relief ventrally and one has a hexa- Eurotestudo: the thigh tubercle ( thigh-spur ) may con-
gonal pygal. This is also congruent with its geographi- stitute a basic autapomorphy in the extant Testudo sp.; it
is absent in Agrionemys and Eurotestudo. In return, the
cal context. As seen above, the species is also rather
color pattern of the dorsal carapace of the  Testudo
similar to Testudo graeca (s.l.) except for the absence
type may constitute a basic synapomorphy linking
of hinge and shorter posterior lobe. But it lacks the
Eurotestudo Testudo s.s. Some characters (previously
protuberances and the posteriorly expanded border of
considered as synapomorphies) are actually weakly
the shell. P. canetotiana is the first European form
homoplastic:
which presents the most derived evolutionary state of
the anterior lobe shape: the trapezoid lobe with ante-  the caudal spur is moderate in Agrionemys, strong
riorly prominent gulars, well laterally exposed, be- and lengthened in Eurotestudo and very small in
comes widened at the anterior border and the gulars T. kleinmanni plus T. werneri. It is present and morpho-
810 F. de Lapparent de Broin et al. / C. R. Palevol 5 (2006) 803 811
logically variable in various lineages of terrestrial Tes- Acknowledgements
tudinidae, but also in other chelonians (Kinosternon, the
extinct Meiolania);
To the Academic Kippis Society; Dr M.F. Bonifay
& Prof. E. Bonifay (Marseille, France); Dr S. Calzada
 the reduction of the fingers of the hand, considered
Badia, Dra A. Masriera (Barcelona, Spain); Dr M. Del-
as shared by Agrionemys and Eurotestudo n.g. is
fino (Florence, Italy); Dr A. Rhodin (Lunenburg, MA,
neither a synapomorphy nor a homoplasy: it is not the

USA); Dr L. Sorbini , Dra A. Vaccari (Verona, Italy);
same character:
to Dr M. Pickford (English correction); Dr J.-C. Rage
and Prof. P. Taquet (review of the note).
four fingers in Agrionemys;
five fingers, but nails 1 and/or 5 are often reduced,
References
rarely absent in Eu. hermanni s.s. Testudo has primi-
tively five fingers [1,3,4,9,23,24,27].
[1] R. Bour, L identit� des Tortues terrestres europ�ennes : sp�ci-
mens-types et localit�s-types, Rev. Fr. Aquariol. 13 (4) (1987)
111 122.
3. Discussion and conclusion
[2] R. Bour, Introduction, Manouria 7 (22) (2004) 2.
[3] R. Bour, Testudo boettgeri, Manouria 7 (22) (2004) 9 10.
None of the approaches, either morphological [9,23,
[4] R. Bour, A new character for the identification of populations of
24] or molecular [14,22,etc.], provides a strong hypoth- the Hermann s tortoise, Testudo hermanni Gmelin, 1789 (Che-
lonii, Testudinidae), Salamandra (Frankf.) 40 (1) (2004) 59 66.
esis of inter-relationships of the lineages or within spe-
[5] F. de Broin, Contribution ą l �tude des Ch�loniens. Ch�loniens
cies in the genera. The results differ according to the
continentaux du Cr�tac� et du Tertiaire de France, M�m. Mus.
authors, the taxa included, the type and amount of ge-
natl Hist. nat. Paris, Ser. C 38 (1977) i ix, 1 366.
netic material, the number of specimens and the method
[6] V.M. Chkhikvadze, [Sur l origine des tortues terrestres pal�arc-
tiques actuelles], Bull. Acad. Sci. Georgian SSR 57 (1) (1970)
employed. The hermanni group (including Eu. hermanni
245 247 (in Russian).
alone or with Eu. boettgeri) may be sister to Agrionemys
[7] V.M. Chkhikvadze, Sur la position syst�matique des Tortues
[9,14] or else with Indotestudo or others [22]. Or it may
terrestres g�antes du Tertiaire du Pal�arctique, Soovshch,
be placed as the sister of a clade with Agrionemys and
Akad. Nauk. Gruznuskoj. 65 (3) (1972) 745 748 (in Russian).
Testudo [24]. In recently constructed cladograms [20],
[8] V.M. Chkhikvadze, in: Les tortues fossiles du Caucase et du
Nord de la mer Noire, Metzniereba, Tbilissi, 1983, pp. 1 149
the three genera are well separated after  Er . bruneti
(in Russian).
in every hypothesis:  T. promarginata is either the sister
[9] S. Gmira, �tude des Ch�loniens fossiles du Maroc. Anatomie.
group of both Testudo s.s. and Eurotestudo or the sister
Syst�matique. Phylog�nie, Cahiers de Pal�ontologie, �ditions
taxon of the three genera in politomy if the poorly pre-
du CNRS, Paris, 1995 (140 p.)
served  T. turgaica is excluded from the analysis. Even [10] S. Hervet, Les Tortues du Quaternaire de France : critŁres de
d�termination, r�partitions chronologique et g�ographique, M�-
if the exact link-point between the three lineages is not
sog�e 58 (2000) 3 47.
definitely established, their separation and differentiation
[11] E. Jim�nez Fuentes, J.-L. Cardoso, E.G. Crespo, Presencia de
is well established. We can hypothesize that the shared
Agrionemys (=Testudo) hermanni (Gmelin, 14789) en el Paleo-
origin of Testudo and Eurotestudo n.g. is more probable
l�tico medio de la Gruta Nova da Columbeira (Bombarral, Pro-
than that of Eurotestudo n.g. and Agrionemys, a pre- vincia de Estremadura, Portugal), Stud. Geol. Salmant. 34
(1998) 123 139.
viously proposed hypothesis [9]: according to the new
[12] L.I. Khozatsky, M. Mlynarski, Agrionemys  Nouveau genre de
analysis [20], similarities remaining between Eu. her-
tortues terrestres (Testudinidae), Bull. Acad. Pol. Sci., II, Ser.
manni and extant Agrionemys spp. are mostly primitive.
Sci. Biol. 14 (2) (1966) 123 125.
Beside the derived more curved epiplastral lip, shared by
[13] T. Kotsakis, R�vision des tortues (Emydidae, Testudinidae,
Trionychidae) du Plio-Pl�istocŁne de Valdarno sup�rieur (Tos-
Testudo and Eurotestudo n.g, the derived color pattern of
cane, Italie), Quaternaria 32 (1980) 11 37.
the dorsal carapace of the  Testudo type, shared by ex-
[14] A.C. van der Kuyl, D.L. Ballasina, J.T. Dekker, J. Maas, R.E.
tant species, is also significant. In any case, various
Willemsen, J. Goudsmit, Phylogenetic relationships among the
homoplastic characters have evolved asynchronously in
species of the genus Testudo (Testudines: Testudinidae) inferred
these three lineages. Although the relationships between
from mitochondrial 12S rRNA gene sequences, Mol. Phylogen-
et. Evol. 22 (2) (2002) 174 183.
Eurotestudo n.g., Testudo, and Agrionemys cannot be
[15] F. de Lapparent de Broin, African chelonians from the Jurassic
firmly established, these lineages are clearly and consis-
to the Present. A preliminary catalog of the African fossil che-
tently separated according to all the approaches. What-
lonians, Paleontol. Afr. 36 (2000) 43 82.
ever their inter-relationships may be, the common origin
[16] F. de Lapparent de Broin, Les Ch�loniens de Sansan, M�m.
of the three genera is in Asia before the Oligocene. Mus. natn. Hist. nat., Paris 183 (11) (2000) 219 261.
F. de Lapparent de Broin et al. / C. R. Palevol 5 (2006) 803 811 811
[17] F. de Lapparent de Broin, The European turtle fauna from the [26] J. Per�l�, Morphological variation among Middle Eastern Testu-
Triassic to the Present, Dumerilia 4 (3) (2001) 155 216. do graeca L., 1758 (sensu lato), with a focus on taxonomy,
[18] F. de Lapparent de Broin, The Miocene chelonians from the Chelonii 3 (2002) 78 108.
southern Namibia, B. Senut, M. Pickford (Eds.), Faunas from [27] J. Per�l�, Occurrence and taxonomic significance of thigh-spurs
the southern Namibia, Mem. Geol. Surv. Namibia 19 (2003) in Testudo marginata Schoepff, 1792 and Testudo weissingeri
67 102. Bour, 1995, Herpetozoa 14 (3/4) (2002) 123 126.
[19] F. de Lapparent de Broin, M.T. Antunes, Pleistocene Chelo- [28] J. Per�l�, Testudo hercegovinensis, Manouria 7 (22) (2004) 19
nians from Gruta da Figueira Brava (Arr�bida, Portugal), 20.
Mem. Acad. Cienc. Lisb. Cl. Cienc. 38 (2000) 101 112. [29] A. Pieh, J. Per�l�, Variabilit�t der Maurischen Landschildkr�ten
[20] F. de Lapparent de Broin, R. Bour, J. Per�l�, Definition of the (Testudo graeca Linnaeus, 1758  Komplex) im zentralen Mar-
genus Eurotestudo (Testudinidae, Chelonii): a morphological okko mit Beschreibung zweier neuer Taxa, Herpetozoa 17 (1/2)
approach, Ann. Paleontol (in press). (2004) 19 47.
[21] M. Mlynarski, Tortoises from the Pliocene of Poland, Acta [30] A.N. Riabinin, [Sur les tortues des d�p�ts m�otiens de Bessar-
Geol. Pol. 5 (2) (1955) 161 214 (Consp., 46 62). abie], Trudy Geol. min. Muz. Petra Velikago Imper. Akad.
[22] J. Parham, J.R. Macey, T.J. Papenfuss, C.R. Feldman, O. T�r- Nauk 1 (1915 1918) 1 16 (in Russian).
kozan, R. Polymeni, J. Boore, The phylogeny of Mediterranean [31] A.N. Riabinin, [Testudo turgaica nov. sp. du MiocŁne moyen de
and their close relatives based on complete mitochondrial gen- la r�gion de Turgai], Trav. Mus. G�ol., Acad. Sci. URSS 1
ome sequences from museum specimens, Mol. Phylogenet. (1926) 53 62 (in Russian).
Evol. 38 (2006) 50 64. [32] H.H. Schleich, Neogene Testudines of Germany. Their strati-
[23] J. Per�l�, The genus Testudo Linnaeus, 1758 sensu lato (Testu- graphic and ecological evaluation, Stud. Palaeochel. I (1985)
dines: Testudinidae): Phylogeny, Taxonomy, Conservation, Ph. 249 267.
D. thesis, University of Bristol, School of Biological Sciences, [33] L. Sorbini, M.V. Durante Pasa, Le collezioni paleontologiche
2002 (i xiv + 328 p.) quaternarie del Museo Civico di Storia Naturale di Verona, Ori-
[24] J. Per�l�, The genus Testudo (Testudines: Testudinidae): Phylo- gine, inventario, bibliografia, Mus. Civ. Stor. Nat. Verona, Ser.
genetic inferences, Chelonii 3 (2002) 32 39. Cataloghi 1 (1974) 1 53.
[25] J. Per�l�, Biodiversity in relatively neglected taxa of Testudo L., [34] H.-k. Yeh, Fossil turtles of China, Paleontol. Sin. N.S.C. 150
1758 s.l, Chelonii 3 (2002) 40 53. (18) (1963) 1 112.


Wyszukiwarka