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Biotype, nomenclature and taxonomic diagnostic characters of Testudo hermanni hermanni GMELIN 1789 in Southern France with preliminary notes on comparative egg morphology with T. h. boettgeri MOJSISOVICS 1889.

A C Highfield


The distribution of Testudo hermanni hermanni GMELIN 1789 in France is restricted to the area of Provence and the Massif des Maures, between Toulon and Saint-Raphael on the coast to Draguignan, Le Luc and Sollies Pont inland. The main concentration of tortoises occurs in sites containing suitable habitat between Collobrieres, La Garde Freinet, Gonfaron and Gogolin. Few tortoises are seen close to human habitation, even where the habitat appears suitable although there are some localised exceptions to this. The majority inhabit the relatively inaccessible hillsides and mountains which are a feature of this region at a preferred altitude of between 500 to 700 metres.


Two principal habitats are preferred, densely wooded hillsides and coarsely vegetated gentle slopes. On an expedition in August we found tortoises in both situations. Tortoises were found frequently in heavy scrub and bramble thicket - making the location of specimens for study a somewhat uncomfortable experience. At this time of year the vegetation on the lower slopes is extremely dry and even in the more densely vegetated areas humidity was minimal. Earlier in the year green vegetation is more abundant and the tortoises are much more active. In the very hot and dry conditions of mid-summer much of the day is spent in retreat. Maximum activity occurs early in the morning or in late afternoon.
Plant species sympatric with Testudo hermanni in southern France include; Opuntia ficus-indica, Sedum reflexum, Sedum album, Lavandula staechus, Rubus spp., Thymus vulgaris, Helichrysum staechus, Euphorbia characias, Genista scorpius, Salix spp., Cistus monspeliensis and Juniperus oxycedrus. Tortoises are rarely encountered in habitats thickly populated with Beech trees or where the ground is heavily covered by ferns, favouring a more open aspect. Preferred nesting sites include sunny slopes, deserted terraced olive groves or woodland openings. We found several vacated eggshells in such locations. Most of these appeared to have been deposited by predators, although in one location may have been the result of a successful hatching. Other reptiles sympatric with Testudo hermanni include Emys orbicularis (European pond terrapin), Natrix spp., Laecerta spp. and Vipera spp. Predators of T. hermanni include badgers (Meles meles), beech martens (Martes foina) and weasels (Mustela nivalis). Of these, badgers and beech martens are particularly prevalent predators of nests, in some areas attacking up to 90% of sites (Stubbs and Swingland, 1984). Analysis of faeces samples from wild T. hermanni in France revealed traces of grass, opuntia seed and snail shells in addition to high concentrations of helminths; no external parasites were observed (Bruekers, 1986).


For some years this tortoise (along with those of Spain and the Balearic islands) has been identified as Testudo hermanni robertmertensi WERMUTH 1952. The eastern population of hermanni was referred to T. hermanni hermanni GMELIN 1789. Recently however work by Roger Bour (1987) has demonstrated that these attributions are erroneous, and that the nominotypical species is actually that of southern France, previously referred to T. h. robertmertensi.
Following the discovery in the collection of the Strasbourg Zoological Museum of the holotype of T. hermanni GMELIN 1789 (specimen reference MZUS 121) it became clear that this specimen was derived from the French and not the eastern population of the species. The nomenclature of the original publication therefore stands for this population by virtue of priority over Wermuths later name robertmertensi which is consequently invalid. Of the eastern population, this should be referred to Testudo hermanni boettgeri MOJSISOVICS 1889 (lectotype Testudo graeca var. boettgeri, specimen reference Senckenberg Museum, Frankfurt No. SMF 7836).
The respective type localities of each population are assigned to Collobrieres, France for T. hermanni hermanni GMELIN 1789 and to Orsova, Romania for T. hermanni boettgeri MOJSISOVICS 1889.

Diagnostic characters

1. Supracaudal

It is frequently alleged that it is possible to distinguish between the southern and eastern populations of T. hermanni by determining if the supracaudal shield is divided or undivided. It is also sometimes claimed that T. hermanni can be distinguished from T. graeca using the same criteria (Devaux, 1988 p.22 fig. 1). In fact, neither is true and this character is of no value in specific determination (Highfield, 1990a).

2. Plastron

The plastral markings of Testudo hermanni hermanni GMELIN 1789 are characteristically formed of two almost solid dark bands running longitudinally down the plastron (plate 2). Every specimen of this sub-species examined by the author (several hundred, in situ) have possessed this feature.
The plastral markings of the eastern form, T. hermanni boettgeri MOJSISOVICS 1889 appear to be somewhat more variable but the characteristic form is far less dense and well defined than that of the French population. Some specimens examined have possessed plastrons with dense markings which almost approach that of T. h. hermanni however, so this character should not employed in isolation to diagnose speciation.

3. Dimensions

There appears to be considerable divergence in maximum size between the eastern and southern forms, the former consistently attaining a greater maximum observed size than those of the latter. Heron (1968) examined over 400 French specimens and found that large females attained only circa 155mm and males only circa 130mm. The largest animal of this sample measured only 160mm. More recent observations indicate that the largest size attained by females is 190mm and males 168mm (see Devaux, 1988 also Cheylan, 1981). The specimens observed during field studies in France by the present author all measured less than 170mm.
Measurements of Testudo hermanni hermanni from the Balearic islands and Spain tend to confirm that these dimensions are fairly typical of the western population as a whole (Vroom, 1983, Kramer & Vickers, 1983, Pelaz, 1988).

By comparison the eastern populations of Testudo hermanni boettgeri MOJSISOVICS 1889 can attain remarkable sizes; the author has previously described a specimen which measured 264mm long and weighed 3,420g. (Highfield, 1988) - believed to be a record for the species. Numerous other specimens measuring over 200mm have also been observed.

4. Colouration & carapace markings

The groundcolour of T. h. hermanni is typically a bright golden yellow. This feature was very obvious in all of the specimens observed in France; it contrasts sharply with most specimens of the eastern T. h. boettgeri where the groundcolour could best be described as a greenish-yellow. Similarly, the carapace markings of the French population seem to be unusually clear and well defined compared to most eastern specimens. One feature particularly noted in all of the French animals examined was a sharply defined "keyhole" marking in bright yellow on the lower central posterior vertebral scute (just above the supracaudal). We have not observed this character so sharply defined in eastern specimens.

5. External cranial characters

There is a marked difference in appearance between the heads of the western and eastern populations of T. hermanni. The western population have much more elongate and smoothly contoured heads which are typically snake-like in general appearance. The eastern populations have a shorter, more bulbous head of quite distinctive construction. The internal characters of the skull of T. hermanni hermanni have been described by Bour (1989).


The southern French population of Testudo hermanni hermanni GMELIN 1789 is clearly distinguished from the more easterly populations of T. h. boettgeri MOJSISOVICS 1889 by a variety of diagnostic characters. Of these, the relative difference in size between similarly aged adults of the two populations and the unusually bright markings of the southern population are particularly notable. Recent studies also reveal a major difference in egg morphology between the French and eastern European populations as shown in table 1. (Highfield, 1990b and in litt).
Within the eastern populations of T. h. hermanni GMELIN 1789 there are considerable local geographic variant forms. These may well eventually prove to be further sub-species.


The author would like to thank Roger Bour, Bernard Devaux and Eric Gibson for their assistance during his visit to France.


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