Chap. 25
Amphibians and reptiles
Amphibiens et reptiles
p. 443-447
Résumés
The Pre-Pottery Neolithic (PPNA) site of Klimonas yielded a total of 53 herpetofaunal remains representing at least five taxa: an amphibian anuran (Bufotes viridis) and four reptiles [a terrestrial turtle, very likely a member of the subgenus Testudo (T. (Testudo) sp. or Testudo s.s.), the scincid Eumeces schneideri and two snakes, the colubrid cf. Dolichophis jugularis and the viperid Macrovipera lebetina]. The endemic or anthropogenic origins of these taxa are discussed with special attention to the terrestrial tortoise. It was found in a Late Pleistocene level at Aetokremnos as well as in the Early Holocene levels of Klimonas, suggesting that it might have been native to the island in the past, before becoming extinct. Digestion marks on three viper vertebrae suggest that they had been swallowed by a dog.
Le site néolithique pré-céramique de Klimonas a livré un total de 53 restes d’herpétofaune provenant principalement du Bâtiment communautaire et de la fosse dépotoir des bâtiments B25-26. Au moins cinq taxons sont présents sur le site : un amphibien anoure (Bufotes viridis) et quatre reptiles [une tortue terrestre, très probablement un membre du sous-genre Testudo (T. (Testudo) sp. ou Testudo s.s.), le scincidé Eumeces schneideri et deux serpents, le colubridé cf. Dolichophis jugularis et le vipéridé Macrovipera lebetina]. Parmi les taxons déterminés, le crapaud vert (B. viridis), le scinque de Schneider (E. schneideri), la couleuvre Grand Fouet noir (D. jugularis) et la vipère lébétine (M. lebetina) sont actuellement présents à Chypre. Le crapaud vert, le Grand Fouet noir et la vipère lébétine sont connus dans l’île au moins depuis l’Épipaléolithique d’Aetokremnos, tandis que pour le scinque de Schneider, le PPNA de Klimonas constitue la plus ancienne mention de l’espèce dans l’île. L’origine ancienne dans l’île de E. schneideri, D. jugularis et M. lebetina est bien attestée par la nature endémique des sous-espèces connues à Chypre (E. schneideri ssp., D. jugularis cypriacus et M. lebetina lebetina). L’origine de B. viridis reste énigmatique. Tous les spécimens de tortues trouvés récemment à Chypre sont réputés provenir de lignées introduites sur l’île. Les découvertes dans un niveau du Pléistocène tardif d’Aetokremnos ainsi que dans les niveaux du début de l’Holocène de Klimonas, suggèrent que Testudo (très probablement un membre du sous-genre Testudo) pourrait avoir été originaire de l’île dans le passé, puis s’est ensuite éteint. La présence de traces de digestion sur des vertèbres de vipère lébétine, et leur intensité, montrent qu’un mammifère carnivore (probablement un chien) est à l’origine de leur dépôt.
Texte intégral
Introduction
1The different excavation sessions carried out in Klimonas (PPNA, Cyprus), provided a total of 53 bone remains of amphibians and reptiles and a minimum number of 21 individuals. At least five taxa have been identified: the green toad (Bufotes viridis), a tortoise (cf. T. (Testudo)), the Schneider’s skink (Eumeces schneideri), a colubrid with a vertebral morphology close to the large whip snake (cf. Dolicophis jugularis), an indeterminate “colubrine” and a viperid, the blunt-nosed viper (Macrovipera lebetina).
2The vast majority of the herpetofaunal remains came from structures or contexts related to the different buildings discovered at the site, mainly from the Communal building (tab. 25-1).
1. Systematic, ecological and taphonomic data
1.1. Amphibia Linnaeus, 1758
3Anura Duméril, 1805
4Bufonidae Gray, 1825
5Bufotes viridis (Laurenti, 1768), green toad
6Material: 1 coracoid (fig. 25-1, A).
Fig. 25-1 – Bufotes viridis

A: coracoid. Anura indet, B: right humerus, ventral view. cf. Testudo (Testudo) or Testudo s.s., C: right scapula, posterior view; D: right peripheral 2, external view. E: modern Testudo graeca (MNHN-ZA-AC-1891-863), carapace, posterior area. F: Testudo (Testudo) or Testudo s.s. suprapygal 2 from Klimonas, external view. G: modern Testudo graeca (MNHN-ZA-AC-A5163), carapace, posterior area. N7 = neural 7; Sp1 = suprapygal 1; Sp2 = suprapygal 2; P = pygal.
Bufotes viridis, A : coracoïde. Anura indet., B : humérus droit, vue ventrale. cf. Testudo (Testudo) ou Testudo s.s., C : scapula droit, vue postérieure ; D : périphérique 2 droite, vue externe. E : Testudo graeca actuelle (MNHN-ZA-AC-1891-863), carapace, zone postérieure. F : Testudo (Testudo) ou Testudo s.s. suprapygal 2 de Klimonas, vue externe. G : Testudo graeca actuelle (MNHN-ZA-AC-A5163), carapace, zone postérieure. N7 = neural 7 ; Sp1 = suprapygal 1 ; Sp2 = suprapygal 2 ; P = pygal. Clichés S. Bailon, CNRS.
Photos S. Bailon, CNRS.
Tab. 25-1 – Amphibians and reptiles from Klimonas. Number of remains and minimum number of individuals (in brackets); C-B: Communal building; B: Building; WP: Waste pit; OC: Out of context.
C-B | WP-B 25-26 | B-PPNA | B 7-8 | B 8 | B 10 | B 18 | B 25-27 | B 26 | OC | ||
Anura indet. | 2 (1) | ||||||||||
Bufotes viridis | 1 (1) | ||||||||||
Testudo sp. | 2 (1) | 2 (1) | 1 (1) | 1 (1) | 1 (1) | ||||||
Eumeces schneideri | 5 (1) | 2 (2) | 1 (1) | 1 (1) | 3 (1) | ||||||
“Colubrinae” indet. | 3 | ||||||||||
Dolicophis jugularis | 1 (1) | 9 (1) | 1 (1) | 1 (1) | 1 (1) | 1 (1) | |||||
Macrovipera lebetina | 12 (2) | ||||||||||
Snake indet. | 1 | 1 | |||||||||
Total | 23 (6) | 11 (2) | 1 (1) | 1 (1) | 3 (3) | 2 (2) | 1 (1) | 1 (1) | 2 (2) | 8 (2) | 53 (21) |
Amphibiens et reptiles de Klimonas. Nombre de restes et nombre minimal d’individus (entre parenthèses) ; C-B : Bâtiment commautaire ; B : bâtiment ; WP : fosse à déchets ; OC : hors contexte.
7The presence of a glenoid process and an epicoracoid process of similar width on this element allows its attribution to the Bufonidae (according to the criteria of Bailon 1999a). Currently, the green toad (B. viridis) is the only known bufonid in Cyprus.
8The green toad is found in dry and sandy habitats, except during its reproductive period where it is found in aquatic environments.
9A humerus (fig. 25-1, B) and a radio-ulna were attributed to an anuran amphibian, but due to their fragmentarion, precise attribution was not possible.
10For these elements, spongy bone is visible on the articular surfaces and the diaphyseal portions of the bones show some alterations, mainly abrasions due to the chemical action of the sediment and roots, and not linked to the digestion of a predator.
11The status of the current populations of green toad (B. viridis) on the island is still uncertain, although an ancient origin seems to be the most likely (Baier et al. 2009). One bufonid remain has been recorded in a possible Pleistocene deposit at Cape Pyla (Bufo sp. (currently Bufotes sp.; Sanchiz 1984). The Green Toad is also mentioned at the Aetokremnos site (Bailon 1999b), probably from the Epipaleolithic human occupation levels (stratum 1C), and in many wells of the Neolithic Mylouthkia site (between the 10th–9th millennia cal BC; Croft et al. 2017).
1.2. Reptilia Laurentis, 1768
12Chelonii Latreille (1800)
13Testudinidae Batsch, 1788
14Testudo Linnaeus, 1758, cf. T. (Testudo), (or Testudo s.s).
15Material: a right scapula (fig. 25-1, C), a left humerus, a costal pair plate fragment, two plate fragments, a right peripheral 2 (fig. 25-1, D) and a suprapygal 2 (fig. 25-1, F).
16In this study, we use the term Testudo in the traditional meaning of the genera (Testudo sensu lato) and the sub-genera Testudo (or Testudo s.s.) for T. (Testudo) graeca, T. (Testudo) kleinmanni and T. (Testudo) marginata; Chersine for T. (Chersine) hermanni and Agrionemys for T. (Agrionemys) horsfieldii (TTWG 2017).
17The texture and morphology of the plates and bones are exclusive to a terrestrial turtle Testudo s.l.: relatively thick plates with well-developed growth rings and a granular outer surface; deep sulci between the scutes; coincidence of the pleuro-marginal sulci with the costo-peripheral sutures; posterior margin of the fifth vertebral scute overlying the suture between the suprapygal and the pygal; the scapula and the acromion form an obtuse angle of about 110 ° and the medial and lateral tuberosities of the humerus are long and close together (Hervet 2000, Bailon 2010, Pérez-García et al. 2015). These character stages allow us to exclude the assignment of this taxon to the European freshwater turtle Emys orbicularis (Linnaeus, 1758) and Mauremys leprosa (Schweigger, 1912) and to the Trionychidae members. It also differs from Titanochelon (Pérez-Garcia and Vlachos 2014; formerly known as Geochelone) due to its much smaller size.
18The morphology of the suprapygal from Klimonas deserves to be commented. It is trapezoidal, wider than high and with a concave anterior border (fig. 25-1, F). In Testudo s.l., the morphology of the suprapygal region presents a wide variability and represents a typical example of the heterochronic-homoplastic evolutionary process (Lapparent de Broin et al. 2006). The general trend observed by these authors is the presence of a single suprapygal in T. (Chersine) and two suprapygales in T. (Testudo) and T. (Agrionemys) (however, the opposite case is also possible, although with much less frequency). In T. (Agrionemys), the suprapygal 2 is generally lenticular--trapezoidal; trapezoidal with a rounded anterior border or fused (3a, 4a and 5 states, respectively, in Lapparent de Broin et al. 2006), while in T. (Testudo) (or Testudo s.s.) this variability is more important (lenticular (state 1); mixed lenticular on one side and trapezoidal on the other (state 2a); lenticular-trapezoidal with a rounded anterior border (state 3a); trapezoidal with an straight anterior border (state 4a) or fused (state 5) (Lapparent de Broin et al. 2006)]. In the comparative material consulted, this variability is even greater in T. (Testudo) (essentially T. (Testudo) graeca), with a suprapygal 2 with a slightly concave to very concave anterior border and, therefore, a suprapygal 1 posteriorly embraced by the second (fig. 1E and 1G). The morphology of the -suprapygal 2 from Klimonas constitutes an intermediate case between this last morphology and state 4a of Lapparent de Broin et al. 2006. In Klimonas, this suprapygal 2 should partially embrace the posterior region of the suprapygal 1 in the living individual.
19These morphological considerations and the biogeographical data (T. (Testudo) is the only Testudinid present in the western Mediterranean) allow us to infer the presence of a representative of this turtle group in Klimonas.
20According to Baier et al. (2009), all recent Testudo specimens found in Cyprus were most likely introduced by humans, and we have no data to prove the existence of stable current wild populations on the island.
21A likely representative of the subgenre Testudo [T. (Testudo) sp.; Testudo (probably T. graeca/ibera in Reese 1999, after R. Bour identification) could already be present in the Late Pleistocene hippopotamus layer at Aetokremnos. The problem lies in not knowing whether this hippopotamus layer is -contemporary with the arrival of humans on the island (Hadjisterkotis et al. 2000) or earlier (Vigne et al. 2014, Zazzo et al. 2015). In the first case, the introduction of this taxon by the first human colonizers remains possible, while in the second case, it would be a native taxon. The presence of Testudo in Klimonas seems to show a certain continuity of the taxon in Cyprus. Other Testudo remains have been found in archaeological contexts of the Middle and the Late Cypriot Period (essentially tombs and trenches from the Bronze Age; Hadjisterkotis and Reese 1994). These findings suggest that Testudo might have been native to the island in the past but then became extinct.
22Note also that Vigne et al. (2017b) preliminarily reported Mauremys rivulata in Klimonas but it has not been detected in our material. Even though this species is attested (for the first time) in the 8th millennium cal BC deposits of Shillourokambos (Vigne 2011a), the preliminary identification at Klimonas is probably mistaken.
1.3. Squamata Oppel, 1811
23Lacertilia Owen, 1842
24Scincidae Gray, 1825
25Eumeces schneideri (Daudin, 1802), Schneider’s Skink.
26Material: 1 maxilla, 6 dentaries (5 right and 1 left), 1 humerus, 1 ulna, 2 femurs and 1 caudal vertebra.
27The Schenider’s Skink is characterized by the presence of pleurodont, cylindrical, unicuspid, robust and latero-medially enlarged teeth on the maxilla and dentary bones. In addition, the dentary shows, in lateral view, a strong posterior impression of the masticator muscles (fig. 25-2, A, B). This is a terrestrial species, living in dry, bushy areas with rocky outcrops. Currently, only a few isolated populations are recognized in Cyprus, where the species may be in serious threat of extinction (Atatür and Goçmen 2001, Baier et al. 2009). This is the oldest mention of this species in Cyprus and its presence in Klimonas (this work) and the Mylouthkia sites show that it had a wider distribution during the Cypriot Neolithic.
Fig. 25-2 – Eumeces schneideri

A and B: right dentary, lateral and medial views. cf. Dolichophis jugularis, C and D: trunk vertebra, dorsal and ventral views. Macrovipera lebetina, E, F and G: trunk vertebra, dorsal, posterior and left lateral views; H: digested trunk vertebra. Scale= 2mm.
Eumeces schneideri, A et B : dentaire droit, vues latérale et médiale. cf. Dolichophis jugularis, C et D : vertèbre dorsale, vues dorsale et ventrale. Macrovipera lebetina, E, F et G : vertèbre dorsale, vues dorsale, postérieure et latérale gauche ; H : vertèbre dorsale digérée. Échelle = 2 mm. Clichés S. Bailon, CNRS.
Photos S. Bailon, CNRS.
28The surfaces of these remains show many chemical alterations related to sediment and root marks. No trace of digestion was observed on the bones or teeth.
1.4. Serpentes Linnaeus, 1758
1.4.1. Colubridae Oppel, 1811
29cf. Dolichophis jugularis (Linnaeus, 1758), large whip snake.
30Material: 1 trunk vertebra (fig. 25-2, C and D).
31The vertebra belongs to a snake of average size, with a centrum length of 4.6 mm. This vertebra is longer than wide (ratio of centrum length to width of the interzygapophyseal constriction = 1.6). In posterior view, the neural arch is moderately vaulted and has convex posterior edges. In ventral view, the hemal keel is long, narrow and well delimited laterally; the prezygapophyseal processes are conical and moderately elongated. This morphology is reminiscent of the current Dolicophis jugularis vertebrae, but a precise allocation cannot be established without more fossil material. The vertebra is burnt (fig. 25-2, C and D). The large whip snake was found at the Aetokremnos site (stratum 2, probably Epipaleolithic human occupation; Bailon 1999b) and in the Neolithic of Mylouthkia (Croft et al. 2017). Currently the large whip snake is represented in Cyprus by an endemic form (D. jugularis cypriacus), which implies a relatively ancient origin (Göçmen et al. 2008).
32The large whip snake is the most common snake in Cyprus. It is ubiquitous and does not appear to require a particular habitat (Baier et al. 2009).
1.4.2. Colubroidea sp., “Colubrine Type” (sensu Szyndlar, 1991)
33Material: 3 trunk vertebrae.
34These vertebrae are highly fragmented; however, we observed that they are longer than wide and their centrum has a hemal keel but no hypapophysis. These characteristics make it possible to attribute these vertebral fragments to a “colubrine” (sensu Szyndlar 1991) and to differentiate them from other colubroids such as Natrix, elapids and viperids.
1.4.3. Viperidae Oppel, 1811
35Macrovipera lebetina (Linnaeus, 1758), blunt-nosed viper.
36Material: 25 trunk vertebrae.
37Among the Cypriot snakes, the vertebral morphology of the blunt-nosed viper is very characteristic and therefore easily attributed: vertebrae with a centrum length greater than 5 mm in adults; neural arc flattened in posterior view; prezygapophyses and postzygapophyses dorsally inclined; centrum short and robust and well limited laterally, and displaying a well-developed neural spine and hypapophysis (Szyndlar and Rage 1999, Bailon 1999b, Bailon et al. 2010) (fig. 25-2, E–H).
38The blunt-nosed viper is a highly euryoeceous species. It inhabits flat meadows, pasture, pine forests, scrubland and rocky areas in close vicinity to water (Baier et al. 2009). This species is known on the island since at least since the Late Pleistocene in Aetokremnos (Bailon 1999b).
39Three of the studied vertebrae were digested (fig. 25-2, H). They show traces of dissolution, thinning and bone loss related to the action of digestive juices. The intensity of digestion is moderate, like that produced by a carnivorous mammal, although humans cannot be excluded. These digested vertebrae came from the Communal building and the waste pit of buildings B25-26 in Sector B.
2. Conclusions
40The Pre-Pottery Neolithic (PPNA) site of Klimonas yielded a total of 53 herpetofaunal remains, mainly from the Communal building and the waste pit of buildings B25-26. At least five taxa were present on the site: an amphibian anuran (Bufotes viridis) and four reptiles (a terrestrial turtle, very likely a member of the subgenus Testudo (T. (Testudo) sp. or Testudo s.s.), the scincid Eumeces schneideri and two snakes, the colubrid cf. Dolichophis jugularis and the viperid Macrovipera lebetina).
41Among the determined taxa, the green toad (B. viridis), the Schneider’s skink (E. schneideri), the large whip snake (D. jugularis) and the blunt-nosed viper (M. lebetina) are currently present in Cyprus. The green toad, the large whip snake and the blunt-nosed viper are known on the island since at least the Epipaleolithic of Aetokremnos (Bailon 1999a), while for the Schneider’s skink the Neolithic of Klimonas constitutes the earliest mention of the species on the island. The ancient origin of E. schneideri, D. jugularis and M. lebetina on the island is well attested by the endemic nature of the subspecies known in Cyprus (E. schneideri ssp., D. jugularis cypriacus and M. lebetina lebetina) (Göçmen et al. 2008). The origin of B. viridis is still unsolved.
42After Baier et al. (2009), all tortoise specimens found recently in Cyprus were most likely introduced to the island, and it is unclear whether tortoises were native to the island in the past. Findings in a Late Pleistocene level of Aetokremnos, as well as in the early Holocene levels of Klimonas, suggest that Testudo (very likely a member of the subgenus Testudo) might have been native to the island in the past but then became extinct.
43The presence of digestion traces on the blunt-nosed viper vertebrae, and their intensity, show that a carnivorous mammal (probably a dog) is at the origin of their deposit.
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