Jozef Klembara

Revision of the Anguine Lizard Pseudopus laurillardi (Squamata, Anguidae) from the Miocene of Europe, with Comments on Paleoecology

Abstract A revision of the Early-Middle Miocene anguine, Pseudopus laurillardi (Lartet, 1851), is presented based on a detailed anatomical analysis of one newly discovered articulated specimen and numerous disarticulated cranial and postcranial elements from several localities in Germany, as well as on the restudy of the original skeletal material of Lartet. The articulated specimen represents the first record of an articulated anguine from the Neogene. We demonstrate that the contemporaneous anguine Propseudopus fraasii ( = Pseudopus fraasi, Pseudopus moguntinus or Ophisaurus moguntinus) from Germany and elsewhere in Europe represents a junior synonym of P. laurillardi. Three species of Pseudopus can by discriminated in the Cenozoic of Eurasia: P. laurillardi (Early-Middle Miocene of Europe), P. pannonicus (Late Miocene-Middle Pleistocene of central and eastern Europe) and P. apodus (Late Pleistocene-Recent, from Eastern Europe to central Asia). Eleven morphological characters of the skull have been identified that distinguish Pseudopus laurillardi from P. pannonicus and P. apodus. Four of these characters regard the frontal and parietal bones, whereas all other characters regard the dentary and dentition. The genus Pseudopus represents the largest and most robust taxon in the subfamily Anguinae and first occurs in central Europe at the beginning of MN 4 (∼18.5 Ma). In contrast to the extant species, P. apodus, the fossils P. laurillardi and P. pannonicus had a greater ecological plasticity and lived in various types of environments. The fossil remains of these taxa are most frequently found in localities characterized by sub-humid to humid climate, which may indicate that their preferential habitats include forested environments.

A CHORISTODERAN REPTILE (REPTILIA: DIAPSIDA) FROM THE LOWER MIOCENE OF NORTHWEST BOHEMIA (CZECH REPUBLIC)

Abstract Choristoderan material from a microvertebrate horizon in the Czech Republic extends the history of the group into the early-middle Miocene. On cranial and vertebral morphology, the new Czech material most closely resembles Lazarussuchus inexpectatus from the Upper Oligocene of France. The morphological similarity supports attribution of the Czech material to the genus Lazarussuchus, but the combination of a smaller and more rounded upper temporal fenestra and the absence of accessory articular facets on the vertebrae distinguish the Czech form from the type species. The articulated skull of L. inexpectatus, re-examined for this study, and the disarticulated material of L. dvoraki sp. nov. complement each other, and permit a better understanding of skull morphology in this enigmatic genus. They demonstrate clearly that the lower temporal fenestra in Lazarussuchus was secondarily closed by apposition of the squamosal, jugal, quadratojugal, and postorbitofrontal.

Salamander-like development in a seymouriamorph revealed by palaeohistology

The amniotes generally lay eggs on land and are thereby differentiated from lissamphibians (salamanders, frogs and caecilians) by their developmental pattern. Although a number of 330–300-Myr old fossils are regarded as early tetrapods placed close to amniotes on the basis of anatomical data, we still do not know whether their developmental pattern was more similar to those of lissamphibians or amniotes. Here we report palaeohistological and skeletochronological evidence supporting a salamander-like development in the seymouriamorph Discosauriscus. Its long-bone growth pattern, slow diaphyseal growth rate and delayed sexual maturity (at more than 10 years old) are more comparable with growth features of extant salamanders rather than extant amniotes, even though they are mostly hypothesized to be phylogenetically closer to living amniotes than salamanders.

Anguimorph lizards (Squamata, Anguimorpha) from the Middle and Late Eocene of the Hampshire Basin of southern England

The Hampshire Basin in southern England is the only sequence virtually spanning the Eocene-Oligocene boundary in western Europe. It has provided an opportunity to observe changes in populations across the Grande Coupure event, as an overturn in the squamate assemblage of continental Europe has also been noted. Five Tertiary formations have yielded lizard material: Creechbarrow Limestone, Barton Clay, Headon Hill, Bembridge Limestone and Bouldnor Formation. These formations span the period from the late Middle Eocene to the earliest Oligocene. Localities in the Hampshire Basin represent a depositional environment with an increasing tendency towards lacustrine and riparian influences in later horizons. The anguimorph fauna includes one new anguine, Headonhillia parva gen. et sp. nov., diagnosed by: (1) small size, (2) large parietal area levis, and (3) short parietal crista postfovealis. Another anguine, Ophisaurus sp., is identified on the basis of the morphology of the parietal. Three further morphotypes of anguines have been distinguished based on dentary, vertebrae and osteoderm specimens. Two types of indeterminate Anguidae are distinguished on the basis of maxillae and dentaries and three morphotypes of indeterminate Anguidae on the basis of vertebrae. Glyptosaurines have been identified in three members on the basis of osteoderms. ‘Necrosaurs’ have also been identified from the Hampshire Basin: two types, distinguished on the basis of dentary material and osteoderms, are identified as indeterminate ‘necrosaurs’. One type of indeterminate Varanoidea is distinguished on the basis of dentaries. The presence of anguids in the Hampshire Basin is well documented, and they appear to form a consistently diverse assemblage throughout the sequence. Anguine osteoderms are present in both pre- and post-Grande Coupure material. Pre-Grande Coupure vertebrae and osteoderms are similar in size to those of adult Recent anguine Pseudopus apodus, although many of the Hampshire Basin anguines, including Headonhillia parva, represent the smallest anguines known to date.

The Early Miocene squamates of Amöneburg (Germany): the first stages of modern squamates in Europe

Squamate faunas from the MN 1–3 interval (earliest Miocene) are poorly known in Europe and worldwide. Our research brings the first report on a complete squamate fauna from the MN 2 zone in Europe. It comprises a relatively large number of specimens from Wiesbaden-Amöneburg in western Germany. MN 2 is part of a time interval (MP 29–MN 2) covering the latest Oligocene–earliest Miocene, which has been labelled the ‘Dark Period’ as far as snakes are concerned. Unexpectedly, a high diversity of squamates was discovered at Amöneburg. This fauna fills an important gap in our knowledge of herpetofaunas from this time interval in Europe, and it represents the first true beginnings of the modern fauna. The new finds are important for the two reasons. Firstly, the beginning of the Miocene heralds the temporary return to a paratropical humid climate after the relatively cool and dry Oligocene, and the Amöneburg material provides us with the opportunity to observe changes in herpetofaunas during this crucial interval. The fauna comprises some survivors from the Oligocene, and especially new forms which may have either originated locally, in Europe, or dispersed from the East. Secondly, the Early Miocene was marked by the collision of Eurasia with Africa, an event that potentially allowed immigration of African squamates into Europe. Since taxa with African affinities such as Chamaeleonidae and Cordylidae are missing from Wiesbaden-Amöneburg, this suggests that African taxa did not reach Europe in MN 2, but were able to enter it during MN 3. One new species, Blanus thomaskelleri sp. nov., is described. http:\\zoobank.org/urn:lsid:zoobank.org:pub:426A24DC-719B-4D13-86BF-E2858F1F1D49

Comparative Anatomy of the Lower Jaw and Dentition of Pseudopus apodus and the Interrelationships of Species of Subfamily Anguinae (Anguimorpha, Anguidae)

The morphology of the lower jaw and teeth of the legless lizard Pseudopus apodus (Anguimorpha, Anguidae, Anguinae) from Eurasia are described in detail and compared with those of other species of the subfamily Anguinae. The lower jaw anatomy of Pseudopus, especially the dentary and teeth, clearly differs from the genera Ophisaurus and Anguis. Even so, Ophisaurus is largely uniform in its lower jaw morphology across species. The teeth of North American Ophisaurus are slender cylinders, the shafts are mesiodistally compressed and bulge lingually; the apices are curved lingually and posteriorly and have weakly developed cutting edges. Southeast Asian and North African Ophisaurus present conical teeth, with broadened bases, apices more distinctly curved lingually and posteriorly, and cutting edges that are distinctly developed. The lingual surfaces of the tooth apices are striated in Ophisaurus and Pseudopus. The lower jaw of Ophisaurus is in many respects similar to that in Anguis, however, the teeth of Anguis are longer and markedly curved posteriorly. The result of the phylogenetic analysis rendered six equally parsimonious trees. Anguis appears in three alternative positions, as the sister taxon to a clade formed by Ophisaurus and Pseudopus, as the sister taxon of Ophisaurus, or as forming a clade with Ophisaurus which is the sister group to Pseudopus. Anat Rec, 297:516–544, 2014.

A new species of Pseudopus (Squamata, Anguidae) from the early Miocene of Northwest Bohemia (Czech Republic)

ABSTRACT Three species of the genus Pseudopus are recognized in the Cenozoic of Eurasia: P. laurillardi (early—middle Miocene of Europe), P. pannonicus (late Miocene—middle Pleistocene of Central and Eastern Europe), and P. apodus (late Pleistocene—Recent, from Eastern Europe to Central Asia). Here, a new and oldest species of the genus Pseudopus, P. ahnikoviensis, sp. nov., is described from the early Miocene (MN 3) locality Merkur in Northwest Bohemia in the Czech Republic. The new species is distinguished from the other three species of the genus Pseudopus by five possible autapomorphies: length of sulcus between frontal and frontoparietal shields equals length of sulcus between frontal and interfrontal shields; ornamented surface of parietal consisting of densely arranged and very fine ridges, grooves, and pits; ventrolateral ridge of supratemporal process of parietal coincides with lateral margin of supratemporal process, hence no free ventrolateral surface of supratemporal process present; presence of surangular spine of dentary; and apices of crowns of robust bulbous teeth smooth. Pseudopus ahnikoviensis is most similar to P. laurillardi and represents the smallest species in the genus Pseudopus.

New finds of anguines (Squamata, Anguidae) from the Early Miocene of Northwest Bohemia (Czech Republic)

Abstract New material on anguines is described from two Lower Miocene localities in Northwest Bohemia in the Czech Republic: Merkur (MN 3) and Dolnice (MN 4). Although the material is disarticulated, it was possible to assign several elements to one species based on similar ornamentation of the skull roof bones and similar morphology of the teeth. Two new species, Ophisaurusholeci nov. sp. and Pseudopus rugosus nov. sp. are described. Pseudopus rugosus becomes the second species of the genus Pseudopus described from the Merkur locality. In addition to the new Ophisaurus species above, the species O. spinari is described on the basis of the parietal and Ophisaurus sp. 1 and Ophisaurus sp. 2 on the basis of the frontal bones. A further five anguines, designated Anguine morphotypes 1 to 6, are described on the basis of the marginal teeth bearing elements (premaxilla, maxilla and dentary) and on the pterygoid, the angular bone and elements forming the posterior portion of the lower jaw. The described specimens present a relevant contribution to our knowledge of the diversity of anguines in the Early Neogene of Europe.KurzfassungNeues Material von Anguinen aus zwei untermiozänen Lokalitäten, Merkur (MN 3) und Dolnice (MN 4), im nordwestlichen Böhmen in der Tschechischen Republik wird beschrieben. Obwohl das Material disartikuliert ist, war es möglich, einige Elemente aufgrund ähnlicher Ornamentation der Schädeldachknochen und ähnlicher Morphologie der Zähne zu einer Art zuzuordnen. Zwei neue Arten, Ophisaurus holeci nov. sp. und Pseudopus rugosus nov. sp., werden beschrieben. Pseudopus rugosus ist die zweite Art der Gattung Pseudopus, die aus der Lokalität Merkur beschrieben wird. Abgesehen von der neue Art von Ophisaurus wird die Art O. spinari anhand von Parietalknochen beschrieben und Ophisaurus sp. 1 und Ophisaurus sp. 2 anhand der Frontalknochen. Fünf weitere Anguinen, bezeichnet als Anguine Morphotyp 1 bis 6, werden anhand von zahntragenden Elementen (Premaxillare, Maxillare und Dentale) sowie dem Pterygoid, dem Angulare und den Elementen des posterioren Unterkiefers beschrieben. Die beschriebene Anguinen präsentieren einen relevanten Beitrag zu unserer Kenntnis der Diversität von Anguinen in dem unteren Neogen von Europa.

NEW STRUCTURES AND RECONSTRUCTIONS OF THE SKULL OF THE SEYMOURIAMORPH SEYMOURIA SANJUANENSIS VAUGHN

ABSTRACT Several cranial structures are described for the first time in the Lower Permian seymouriamorph tetrapod Seymouria sanjuanensis: 1) septomaxilla, 2) suborbital fenestra, 3) small denticles on the palate densely spaced in compact rows radiating from a point near the midwidth of the posterior margin of the palatal ramus of the pterygoid, 4) long cultriform process, and 5) posterior Meckelian fenestra. The possible presence of the paraquadrate foramen is also described. In addition, a more detailed description is given for the relationships between the maxilla, jugal, and quadratojugal on the ventral rim of the skull and between the nasolacrimal canal and the anteroventral orbital notch. Reconstructions of the skull include revised dorsal and lateral views and a first time ventral view.

COMPARISON OF CRANIAL ANATOMY AND PROPORTIONS OF SIMILARLY SIZED SEYMOURIA SANJUANENSIS AND DISCOSAURISCUS AUSTRIACUS

ABSTRACT Similarly sized skulls of juvenile specimens of the seymouriid and discosauriscid seymouriamorph tetrapods Seymouria and Discosauriscus are compared for the first time. Comparisons are based on a single juvenile skull of Seymouria sanjuanensis from the Lower Permian Bromacker locality, Germany, measuring 56 mm in length and several skulls of Discosauriscus austriacus from the Lower Permian Boskovice Furrow, Czech Republic, reaching a maximum length of 62 mm. Establishment of their stage of development as juvenile is based on the small size of S. sanjuanensis compared to adults of the same species and previous comparisons of various-sized specimens of D. austriacus to complete ontogenetic series of several temnospondyls. In addition, a similar degree of cranial ossification, although slightly more advanced in the S. sanjuanensis skull, is also cited as evidence of very similar ontogenetic stages of development. However, D. austriacus is remarkable in retaining remnants of the lateral line canals, suggesting that a longer premetamorphic period of its life history was aquatic. Despite representing similar, juvenile stages of development, the S. sanjuanensis and D. austriacus specimens can be distinguished clearly from one another on structural and proportional cranial features, thus dispelling suggestions that Seymouria may represent the adult stage of development of Discosauriscus, which is known only by larval and juvenile stages of development.