SYSTEMATIC DESCRIPTION OF THE HERPETO FAUNA FROM THE KÄRLICH MAIN SECTION

The fossil material was identified through comparisons with specimens of extant species from western and central Europe. Due to the limited comparisons, the occurrence of bufonids and lacertids restricted to other parts of Europe and regions such as the Near East cannot be excluded with certainty.

The fossil material is stored in the collection of the Faculty of Archeology (FdA), Leiden University, The Netherlands.

Class Amphibia Linnaeus, 1758
Order Caudata Oppel, 1811
Family Salamandridae Gray, 1825

The family Salamandridae has a wide distribution at present, occurring in Britain, Eurasia, northwestern Africa in the Old World, and from southern Canada through the United States into northern Mexico in the New World (Frost 1985). Species that presently occur in Germany are Salamandra salamandra, S. atra, Triturus alpestris, T. carnifex, T. cristatus, T. helveticus, and T. vulgaris (Günther 1996). Triturus cristatus, T. vulgaris, and T. vulgaris or T. helveticus were recovered from the Pleistocene deposits at Kärlich.

Triturus cristatus (Laurenti, 1768)
Northern Crested Newt

Material. One trunk vertebra from Kä G. (FdA-Kä G nr. 8001).

Remarks. Trunk vertebrae of Triturus cristatus differ from those of T. alpestris, T. helveticus, T. marmoratus, and T. vulgaris in having the posterior border of the neural arch flared and continuing well behind the level of the postzygapophyses (see Holman and Stuart 1991: figure 1a). The trunk vertebrae of T. cristatus further differ from those of T. alpestris, T. helveticus, and T. vulgaris in being larger and in having a much lower neural spine. Northern crested newts often are aquatic throughout the year, although in some situations they become terrestrial during the non-breeding season. These large newts prefer still or slowly flowing water with large amounts of aquatic vegetation (Arnold and Burton 1978). Triturus cristatus previously was called the 'warty newt' or the 'great crested newt,' but the common name now is standardized as 'the northern crested newt' (Frank and Ramus 1995). Triturus cristatus presently occurs in the vicinity of Kärlich (Grosse and Günther 1996).

cf. Triturus vulgaris (Linnaeus, 1758)
Smooth Newt

Material. One trunk vertebra from Kä G. (FdA-KäG nr. 8002).

Remarks. The trunk vertebrae of Tritutus vulgaris may be distinguished from those of T. helveticus on the basis of having a deeper and less broadly U-shaped notch in the posterior border of the neural arch (Holman and Stuart 1991, figures 1b and 1c). This notch also tends to be deeper in T. vulgaris than in T. alpestris, but there is some overlap in this character. Nevertheless, T. alpestris, a medium-sized salamander, has larger vertebrae than the smaller species, T. vulgaris. The smooth newt presently occurs in the Kärlich area (Buschendorf and Günther 1996), spends more time on land than most other species of European newts, and lives in a variety of damp terrestrial habitats. It breeds in shallow, still water with ample vegetation.

Triturus vulgaris (Linnaeus, 1758) or T. helveticus (Razoumowsky, 1789)
Smooth Newt or Palmate Newt

Material. A fragmentary vertebra from Kä H. (FdA-KäH nr. 8001).

Remarks. This small vertebrae represents either a smooth newt or a palmate newt, but the bone is not complete enough to be identified to the specific level. Both forms occur at or near the Kärlich area today (Buschendorf and Günther 1996; Schlüpmann et al. 1996).

Order Anura Rafinesque, 1815
Family Bufonidae Gray, 1825

The family Bufonidae is cosmopolitan in distribution except for Australia, Madagascar, and isolated oceanic islands (Frost 1985). Species that presently occur in Germany are Bufo bufo, B. calamita, and B. viridis (Günther 1996). Bufo bufo and Bufo sp. indet. were found in the Pleistocene at Kärlich.

Bufo bufo (Linnaeus, 1758)
Common European Toad

Material. One right distal humerus of a male individual from KäG, (FdA-KäG nr. 8003) (Figure 3).

Remarks. The humerus is identified as a male on the basis of its enlarged mesial crest (Bailón 1986). The fossil humerus may be separated from those of male specimens of the two other European toads (Bufo calamita Laurenti and B. viridis Laurenti) because the shaft is more robust and less curved, the distal condyle is wider, the radial epicondyle is more distally swollen, and the mesial crest is shorter and wider (see Bailón 1986, figure 4).

Bufo bufo occurs in the Kärlich area today (Günther and Geiger 1996). The common European toad is the most widespread of the three European toad species and occurs in a large variety of habitats, some being rather dry. In southern Europe, Bufo bufo may reach a very large size (Arnold and Burton 1978). Bufo bufo previously was called the 'common toad,' but its vernacular name was recently standardized as the 'common European toad' (Frank and Ramus 1995).

Bufo sp. indet.

Material. A damaged right distal humerus of an immature individual (FdA-KäG nr. 8004) and a damaged right scapula (FdA-KäG nr. 8005) from Kä G. A fragmentary scapula of a juvenile individual (FdA-KäE nr. 8001) from E.

Remarks. Unfortunately the humerus is too damaged to identify to the specific level. We used characters given by Hallock et al. (1990) to distinguish the scapula of Bufo from Rana and other European anurans. The authors state that "scapulae of Bufo may be separated from those of Rana on the basis that Bufo lacks the ridge on the dorsomedial side of the scapula that occurs in Rana. Moreover, the ventral end of the scapula is more deeply notched in Bufo than it is in Rana."

Class Reptilia Laurenti, 1768
Order Sauria McCartney, 1802
Family Lacertidae Bonaparte, 1831

The Lacertidae is a widespread Old World lizard group that occurs in Europe, Africa, and much of Asia (Estes 1983). The six species that occur in Germany today are Lacerta agilis, L. bilineata, L. horvathi, L. viridis, L. vivipara, and Podarcis muralis (Günther 1996). Cf. Lacerta agilis, Lacerta sp. (large species), L. vivipara, and Lacerta or Podarcis species have been identified in the Pleistocene record of Kärlich.

cf. Lacerta agilis Linnaeus, 1758
Sand Lizard

Material. One essentially complete right dentary (FdA-KäG nr. 8006), one partial left dentary (FdA-KäG nr. 8007) (Figure 4), and one partial maxilla (FdA-KäG nr. 8008) from Kä G.

Remarks. The right dentary has 21 teeth and alveolar spaces, and the tooth row is 5.6 mm in length. The dentary differs from the large Lacerta species L. lepida and L. viridis in being smaller (much smaller than in a modern adult L. lepida specimen Michigan State University (MSU l643) which has a tooth row 22.0 mm long), by having fewer dentary teeth (many fewer than in two modern L. lepida (MSU l643, and Museo Nacional de Ciencias Naturales , Madrid (MNCN) l6506) which, respectively, have 27 and 28 dentary teeth, and by lacking tricuspid teeth in the dentary (tricuspid teeth are present in L. lepida and L. viridis).

The right dentary differs from the small Lacertidae species, L. vivipara and Podarcis muralis in being larger (tooth row usually less than 5.0 mm in L. vivipara and P. muralis), in having all of the dentary teeth more swollen and robust, in having the anterior teeth less pointed and posteriorly curved, and in having the Meckelian groove with its borders more robust. The less complete dentary agrees with the right dentary in all of the characters above.

The partial maxilla has only unicuspid and bicuspid teeth present as in L. agilis and P. muralis, but differs from P. muralis in having larger and more robust teeth. The teeth fossils appear identical to those in modern adult specimens of L. agilis from MNCN and National Museum of Natural History Leiden (NMNH). At present, Lacerta agilis occurs throughout most of Germany, including the Kärlich area (Ebling et al. 1996). The sand lizard lives throughout its range in rather dry habitats, i.e., in areas with sandy, well-drained soils. In England and northern Europe, it is usually found on coastal sand dunes with some plant cover or in sandy heathland (Arnold and Burton 1978).

Lacerta sp. (large species)

Material. Six partial maxillae (all with teeth) (FdA-KäG nr. 8009 - 8014) (Figure 5), and one vertebra (FdA-KäG nr. 8015) from Kä G.

Remarks. These six maxillae represent a large species of Lacerta. They bear teeth that are the same size and appear identical to those of an adult Lacerta viridis from the Natural History Museum Naturalis, Leiden with a skull length of 23.2 mm and are larger than those of an adult L agilis from the NMNH with a skull length of l9.0 mm. The maxillae further differ from those of L. agilis in having tricuspid teeth present on the maxillary (Figure 5.1–5.3); tricuspid teeth are lacking in adult L. agilis. On the other hand, these teeth are much smaller than those of an adult L. lepida from the MSU collection, a specimen that also lacks tricuspid maxillary teeth. The vertebra has a centrum length of 3.3 mm, and, thus, is larger than those in available L. agilis specimens, but still much smaller than those of L. lepida. Specific identification of the large Lacerta awaits more fossil and modern comparative material.

Lacerta vivipara Jaquin, 1787
Viviparous Lizard

Material. Two left (FdA-KäG nr. 8016 - 8017) (Figure 6.1) and four right partial dentaries (FdA-KäG nr. 8018 - 8021) (Figure 6.2); two left (FdA-KäG nr. 8022 - 8023) and two right maxillae (FdA-KäG nr. 8024 - 8025); and one maxillary fragment (FdA-KäG nr. 8026), all bearing teeth, from Kä G.

Remarks. The dentaries differ strongly from L. agilis, L. lepida, and L. viridis in being smaller (much smaller than in L. lepida and L. viridis), having the teeth much more gracile, with the anterior teeth more pointed and usually more posteriorly curved, and with the borders of the Meckelian groove much less swollen and robust. The dentaries differ from P. muralis in having several tricuspid teeth in each of the dentary bones, whereas P. muralis has all of its teeth in the maxillary and the dentary bones unicuspid or bicuspid.

The maxillae also are smaller and their teeth much more gracile than in L. agilis, L. lepida, and L. viridis. The maxillae further differ from L. agilis and P. muralis in bearing tricuspid teeth. All of these elements seem identical to those in four modern adult MSU Lacerta vivipara skeletal specimens.

Lacerta vivipara occurs in the Kärlich area today (Günther and Völkl 1996a). At present it is a ground-dwelling lizard that is found in many kinds of open situations such as heathlands, open woods, and grasslands.

cf. Lacerta or Podarcis sp. indet.
Small Lacertid species

Material. A small, fragmentary vertebra from Kä H (FdA-KäH nr. 8002).

Remarks. This triangular-shaped lizard vertebra represents either Podarcis or one of the small species of Lacerta, such as L. vivipara.

Family Anguidae Gray, 1825

The subfamily Anguinae is a Holarctic group that is widespread in the United States and Mexico and is represented by both limb-bearing and limbless species. In Europe, however, there are only two monotypic genera, Anguis fragilis, and Ophisaurus apodus, both of which are limbless. Only one species, Anguis fragilis, presently occurs in Germany, and that species was found in the Pleistocene of the Kärlich locality.

Anguis fragilis Linnaeus, 1758
Slow Worm

Material. One hundred and twelve trunk vertebrae, 21 caudal vertebrae, and five osteoscutes from Kä G. Nine trunk vertebrae are from Kä H (FdA-KäH nr. 8003-8011).

Remarks. Trunk vertebrae, caudal vertebrae, and osteoscutes of Anguis fragilis are all characteristic, and can only be confused with those of Ophisaurus apodus, a very much larger form. Holman (1998) gives characters that distinguish the vertebrae of the two species. The trunk vertebrae are particularly diagnostic, being quadrangular in shape, having very large prezygapophyseal and postzygapophyseal articular surfaces, and with the ventral surface of the centrum very flat and smooth. The caudal vertebrae have conspicuous fracture planes in them and the tails of slow worms are easily broken, allowing the animals to escape from predators that seize them behind the anal region.

The limbless slow worm lizard prefers habitats that are well covered with vegetation and other ground cover, and prefers somewhat damper environs than the previous lizard species. The species occurs in the Kärlich area today (Günther and Völkl 1996b).