Abstract

In memory of Oldřich Fejfar

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Pannonicola and Ischymomys are two extinct genera of the family Cricetidae, and both of them were placed at the base of the evolutionary trunk of the subfamily Arvicolinae. Over the last 100 years, the latter has provided the most important fossils for the biostratigraphy of terrestrial Neogene and Quaternary deposits of the Holarctic. The type material of Pannonicola comprises only one m2 and one m3. The first finds of Ischymomys were initially rare, not figured, and their nomenclatural status was faced with various problems. Some authors finally assumed that Ischymomys was a junior synonym of Pannonicola. In this paper, we use new finds from the territory of Ukraine and previously published data to show that Ischymomys is an independent genus, and that the finds described as Pannonicola should probably be placed in the genus Microtodon.

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Shrews of the tribe Nectogalini rank among the rarest items of the late Cenozoic fossil record, and their interpretation is traditionally accompanied by numerous controversies. Here, the data from the Czech Republic and Slovakia covering in total 83 MNI from 45 Pliocene and Quaternary (MN 15 – Q 4 biozones) community samples of 25 sites are reported. A detailed biometric analysis of both Asoriculus spp. (MN 15 – Q 1) and Neomys spp. (Q 1 – Recent) was undertaken to reveal patterns of phenotype variation in particular samples, and their relations to extant mid-European species Neomys fodiens and N. milleri. For that purpose, we examined the variation pattern of 239 dental, rostral and mandibular variables in a large set of extant species (n = 135), quantified their discrimination capacity and by forward selection established a set of criteria applicable for identification and comparative analyses of fragmentary fossil specimens.
We found that both extant species occurred in Central Europe with roughly equal number of records both in the Last Glacial and since the beginning of the Holocene.
The late Biharian and early Toringian samples exhibited statistical homogeneity and differences from extant N. milleri – they are reported here as N. newtoni. Tentatively, we proposed a separate status also for the items of the latest Villanyian – early Biharian age (as Asoriculus castellarini). Compared to both extant Neomys sp. and MN 15–17 Asoriculus gibberodon, the above-mentioned samples seem to be characterized by a broad phenotype variation, particularly in the characters associated with assumed Asoriculus-Neomys transition (mandibular unicuspids, distal mandibular structures). Contrary to the hypothesis on parallel appearance of both genera in the Early Pleistocene and the extinction of Asoriculus by the end of the Early Biharian, we consider the Early Pleistocene diversity of the clade as a stage of a broad gradual transition subsequently driven by adaptive advances of Neomys constitution.
In any case, the morphometric comparisons with the non-European representatives of Nectogalini confirmed close relations between both the European genera and distant divergence of the European clade from the extant Oriental clades.

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The genus Aztlanolagus from North America is synonymized with Pliopentalagus based on similar morphology of dental characters including (1) general form of five reentrant angles and outline of p3, (2) an enamel lake at the position of PIR, (3) general form of enamel crenulations of the anterior wall of the talonid on p4–m2, and (4) an AER on p4–m2 on some specimens. However, the species A. agilis is valid because of its much smaller size. The general morphology and size of this species, transferred to Pliopentalagus, did not change for about the last 2.5 million years. Three Asiatic species of Pliopentalagus (Pl. huainanensis, Pl. dajushanensis, and Pl. anhuiensis, from geologically older to younger) were described from the latest Miocene (ca. 6 Ma) to Late Pliocene (ca. 3 Ma) at Dajushan in Anhui Province, China. These taxa represent a gradually evolving lineage, an example of phyletic gradualism. Pliopentalagus okuyamai was recently described based on a fragmentary skull with upper dentition from ca. 3.5 Ma deposits in Japan and represents a possible ancestral form of the living type species, Pentalagus furnessi. Fossil Pliopentalagus dietrichi is known from the Early Pliocene in Europe and likely evolved from Chinese Pl. dajushanensis. However, Pl. dietrichi lacks an enamel lake on p3 and an AER on p4–m2; this assumes that the ancestral population dispersing to Europe probably did not have a gene controlling the appearance of an enamel lake on p3 and an AER on p4–m2. Thus, Pliopentalagus diversified and dispersed widely across the Holarctic region over the last 6 million years, disappeared near the end of the Pleistocene, and left a single descendant, Pentalagus furnessi in Japan.

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Fluviatile deposits of the Krutaya Gorka regional formation host a new locality of Early Pliocene continental biota from southern Western Siberia. The Novaya Livenka site provides important data on palynology, paleocarpology, molluscs, fishes, and small mammals, mostly coming from a single fossiliferous bed. This study is the first successful insight into regional Early Pliocene palynology reliably controlled by the stage of mammalian evolution. The micromammal association is dominated by remains of pika Ochotona sp. and the primitive brachyodont vole Promimomys. Important biochronological and paleoecological elements of the fauna also include a shrew Parasoriculus, a hazel dormouse Muscardinus, a small beaver Trogontherium and other forms. The vole P. cf. davakosi characterizes a regional phyletic stage of Arvicolinae between a more brachyodont P. antiquus of earlier Ruscinian and forms of the vole radiation event of the late Ruscinian. Synthetic biotic data enable a reconstruction of a smaller water body with a slow current fringed by patches of broad-leaved and conifer forest, and meadow and steppe-like open landscapes under a humid and warm temperate climate. Biochronologic signals of mammals and palynology imply the Early Pliocene age of the biota and its correlation with late early Ruscinian ELMA (MN 14) and mid-Zanclean between ca. 4.5 and 4.3 Ma.

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Csakvaromys sciurinus is an enigmatic stem ground squirrel from the Late Miocene of Hungary that has previously only been briefly described. The restudy of the holotype allows a detailed redescription and comparison with different xerines. Csakvaromys sciurinus is recognized as a junior synonym of Csakvaromys bredai and diagnosed by the following combination of characters: medium size, close to the maximum limits of C. bredai; massive body of the mandible with deep diastemal portion and shallow diastemal depression; elevated area of the mental foramen; nonconfluent lower and upper masseteric crests separated by the attachment area for the anterior fibers of the anterior deep masseter muscle; transversely compressed lower incisor with longitudinally striated anterior enamel face; and brachyodont cheek teeth with mesoconids, rudimentary entoconids, and well-developed anteroconulid of p4.

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Fossils of Metatheria and Eulipotyphla from the karstic fissures MWQ2/2003, MCQ3/2005 and MWQ4/2018 of Mokrá-Quarry (South Moravia, the Czech Republic) are studied. These Early Miocene fissures have yielded remains of the herpetotheriid Amphiperatherium frequens, together with the erinaceids Amphechinus cf. baudeloti, Galerix exilis, and G. symeonidisi. The talpid Talpa sp. has also been identified. The rare remains of Amphiperatherium frequens represent the first description of this species from the Early Miocene of the Czech Republic. The erinaceid Amphechinus cf. baudeloti could be a convenient structural ancestor of A. baudeloti, although the scarcity of remains hampered a clear taxonomic verdict. In addition, the relatively abundant remains of Galericinae in the studied fissures made it possible to identify a mixture of two Galerix species, G. exilis and G. symeonidisi. Their cooccurrence is evidenced at least in MWQ2/2003. The dentognathic and postcranial remains of Talpidae clearly point to Talpa sp. The relatively low diversity of insectivores recovered from Mokrá-Quarry fissures agree with previous studies concerning the paleoenvironmental reconstruction, depicting Mokrá-Quarry as a dry karst plateau close to water bodies and patches of woodland. The wide range of small mammal species found in Mokrá-Quarry fissures suggests a diverse regional environment, making Mokrá-Quarry a reference locality in the Early Miocene of Central Europe.

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Rivoli Veronese (Verona, north-eastern Italy) is a karst fissure filling that yielded an Early Pleistocene fauna, including many remains belonging to the genus Talpa, assigned to the late Villanyian rodent age. Some of these remains, characterized by tiny size and primitive features, were already in the past suggested to represent a new mole species. To better characterize this mole, a re-analysis of the most representative fossil and living populations of the genus Talpa was performed. This allowed confirmation that it represents a new species, which was named Talpa masinii n. sp., to infer that this species existed in earlier periods in Central Europe too, and to better define the taxonomy and biochronology of the fossil Talpa populations.

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Ochotonidae and Leporidae are two living families belonging to the Order Lagomorpha, an ancient group of mammals originating in the Paleogene of Asia. Those families diversified in the Oligocene and Miocene. More primitive stem lagomorphs inhabited Asia during a time of tropical environmental conditions. A Mid-Cenozoic change towards more continental and arid climate in parallel with Antarctic glaciation resulted in significant reorganisation of paleoenvironmental and climatic conditions in Asia and involved the opening of terrestrial connections between Asia, Europe, and North America, allowing faunal exchanges. This promoted diversification and speciation of lagomorphs in the northern continents. Pronounced lagomorph turnover is documented for Central Mongolia, with new data obtained for the Baikalian region as well as adjacent areas. The earliest lagomorphs were represented by archaic stem genera including paleolagids. These were successively replaced by modern Ochotonidae and Leporidae that flourished during the Miocene and Pliocene. The diversity and abundance of ochotonids and leporids decreased during the Pleistocene and of ochotonids, only the pika genus Ochotona survived to the present.

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Studies during the 1970’s and 1980’s concluded that the continental Bou Hanifia Formation spanned the Vallesian and Turolian stages, the majority early opinion being that it was basal Vallesian and that it had yielded the oldest known remnants of the equid Hipparion from Africa. This opinion was taken to be backed-up by radio-isotopic age determinations of ca. 12.2 Ma obtained from volcanic tuffs that were originally mapped as being near the base of the formation. However, later suggestions in the literature indicated that some of the large mammals could be as young as late Turolian to Ventian (8–7 Ma). If so, then a major revision of North African biostratigraphy would be required. This paper focuses on the large mammals and avian eggshells recovered from the Bou Hanifia region, taking into account recent revisions of African faunas and a much augmented data base about their stratigraphic distribution, and it is concluded that the Bou Hanifia fossils have the closest relations to material from the late Turolian and Ventian Stages, and are thus likely to be of latest Miocene (Ventian) age (MN 13).

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Dento-gnathic remains of a fossil suine currently housed at the Université d’Alger are accompanied by a label similar to those associated with other fossils excavated by Arambourg in 1951 at Bou Hanifia from deposits that were, at the time, correlated to the upper Vindobonian or late Middle Miocene. The undescribed suine fossils are enigmatic, in that they do not correspond in geological age to other fossils with similar labels. Several questions are posed including the possibility of mis-labelling. The fossils described herein are attributed to the suine Kolpochoerus maroccanus and correspond closely in terms of morphology and dimensions to material of Kolpochoerus heseloni from the Early Pleistocene of Eastern Africa (Shungura Formation D-G, Ethiopia; Burgi level, Kenya).

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The morphometric variability of the canines and cheek teeth of the fossil and modern dhole (Cuon alpinus) is considered. It was not possible to detect sexual size dimorphism in the species. Geographical variability of dental parameters established two taxon groups: northern, which included two subspecies (C. a. alpinus, C. a. hesperius), and southern, which includes all other recent subspecies. Within the southern group, animals of Southeast Asia (Malacca, Sumatra, and Java) are distinguished by their smaller size. The dhole from the Late Pleistocene of Europe (C. a. europaeus) is close in dental characteristics to representatives of the northern group, and the Late Pleistocene fossil dhole from North America (Mexico) is close to the modern nominotypic subspecies C. a. alpinus. A hypothesis of early, mid-Pleistocene divergence between dholes from the north and north-west of the range and dholes from Southeast Asia is formulated.

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Four sites in northern Greece, presented here, are of particular interest, due to the presence of fossil primates and carnivores in Late Miocene and Pliocene assemblages. The Thermopigi site in eastern Macedonia is particularly important, due to its demonstrable Late Miocene assemblage of more than twenty different species of large mammals, including the postcranial remains (humerus, tibia, calcaneus) of the colobine Mesopithecus delsoni. Carnivores from this site, represented by hyenids, felids, and mustelids, are also described here. Among them, the hyper-carnivorous and widely spread Adcrocuta eximia, an important biostratigraphic marker of Late Miocene, is compared with the single skull from the nearby site of Platania (Drama), both of great taphonomic interest. The site of Milia (Grevena), in Western Macedonia, is also of particular importance, as it includes the earliest Villafranchian age assemblage. New primate material, an ulna and tibia attributed to aff. Dolichopithecus sp. are also described here. Among the new carnivore finds, an incomplete distal tibia belongs to a new felid for the Milia locality, most likely attributable to either a female Megantereon or a new species of Puma. Further investigations are necessary to confirm either attribution. Finally, the new Pliocene site at Promachonas (Serres), mentioned here for the first time, yielded dental material, most likely belonging to cf. Dolichopithecus balcanicus. This study of new material from these sites contributes to a better knowledge regarding the occurrence of primates and carnivores and their paleoenvironments in northern Greece.

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Psammornis rothschildi is an avian oospecies, the type material of which was collected from the desert surface in 1909 (i.e., without stratigraphic context), 20 miles east of Touggourt, Algeria. At 3.2–3.3 mm thick, the eggshell is considerably thicker than those of extant ostriches. Thin sections of the eggshell have been illustrated, and descriptions of the surface features are available, but these have not included images of the external and internal surfaces. This lack has caused uncertainty among authors who have dealt with other occurrences of the taxon, because it has rendered it difficult to compare specimens from other localities with the type material. In order to rectify the situation, images of the original specimens are provided, their curation history is summarised, and a lectotype is nominated so as to stabilise the nomenclature.

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Information on mammal remains from 665 sites/localities was used to estimate the range of size diversity of European species in the Eem interglacial (MIS 5e) and the GS 2.1 stadial (MIS 2). Review of faunal composition and comparison of the rank distributions of the occurrence of species remains across sites allowed determination of the composition of groups of typical and indicator species for each of the time intervals. Entropy, dominance index and self-organization index were calculated from the rank distributions of the occurrence of large, small and medium-sized herbivorous, carnivorous and representatives of the order Eulipotyphla. The parameters of the non-linear function describing the increase in the average number of species that can be found in 1, 2, 3, etc. randomly selected sites were calculated. The relationship between the parameters of these functions and between them and species richness is shown. The paper discusses hypotheses about the relationship between parameter variations and changes in range size ratios in different mammal groups during the Eem interglacial and at the end of the Last Glacial Maximum.

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The mica-clay fossil site Groß Pampau is famous for its later Miocene marine mammal record, which represents the fauna of the then-southern North Sea Basin. Since the 1980s, fossils have been collected and later systematically excavated. For a long time, only marine taxa were identified. In 2017, two nicely preserved remains of terrestrial artiodactyls were discovered as the so far only terrestrial input and northernmost European terrestrial mammal record of the pre-Quaternary Cenozoic. Here, we figure, describe, and compare morphology and dimensions of both specimens. We identified them as fragment of a hemimandible with two lower molars and a distal phalanx of a cervid, but were unable to assign them to a genus or species, due to limited contemporaneous comparative material.

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Dental remains of the large anthracothere, Brachyodus, from the French Faluns (Touraine, Anjou) and the Sables de l’Orléanais, show a broad range of morphological variation, in particular in the degree of enamel wrinkling, the strength of enamel beading on crests and cingula, the form and volume of styles in the upper molars, and occasionally the addition of accessory cusps or cusplets, notably in the P4/. A third lower molar from La Fuye is highly unusual however, in possessing a greatly enlarged entoconid and posthypocristid which radically alter the appearance of the tooth. The specimen is described and compared with other third molars of Brachyodus. It possibly represents a case of teratogeny related to localised injury to, or alteration of, the odontoblast and ameloblast complexes. It is less likely to reflect inbreeding which is hypothesised to result in some cases of anomalous dental formation. Aspects of occlusion and life history of the individual from La Fuye are explored.

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An isolated tooth, of morphology unusual for cave bears, is described from Late Pleistocene deposits (MIS 3) of Pobeda Cave (54.1000° N, 56.5100° E) in the Urals. Analysis of ancient DNA showed that it belongs to the Ural cave bear Ursus kanivetz, which was widespread in the Pleistocene in the Urals. The fossil find was identified as an anomalous upper canine tooth, which got its unusual appearance due to a deviation from normal development under the influence of some unknown internal or external causes.

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