Middle Miocene tragulid remains from Abocador de Can Mata: The earliest record of Dorcatherium naui from Western Europe

In the past, the tragulid species Dorcatherium crassum has been considered a synonym of Dorcatherium naui, although they are currently considered distinct species. The former has an exclusively Aragonian (MN4 to MN6) stratigraphic range, whereas D. naui is considered a typically Vallesian species (MN9 and MN10) that survives until the Turolian (MN11). There are only a few previous citations of Dorcatherium from the European latest Aragonian (MN7+8), and only that of Przeworno (Poland) can be securely attributed to D. naui. Here we describe dentognathic and postcranial remains of Dorcatherium from several localities from the stratigraphic series of the Abocador de Can Mata (ACM), in els Hostalets de Pierola (Vallès-Penedès Basin, Catalonia, Spain). These Dorcatherium remains, from both the lower (ca. 11.9-12.5 Ma, MN7) and upper (ca. 11.6 Ma, MN8) portions of the ACM series, constitute the earliest record of D. naui from the Iberian Peninsula and Western Europe. They most closely resemble D. naui from the Vallesian, but still share several features with D. crassum. On the basis of the unique combination of features displayed by the ACM material, a new subspecies, D. naui meini is erected. Its distinct status from the nominotypical subspecies, D. naui naui, is reinforced by the chronological differences between the two taxa. The morphological similarities between Dorcatherium and the extant water chevrotain (Hyemoschus aquaticus) tentatively suggest the presence of forested and humid paleoenvironments in this area of the Vallès-Penedès Basin during the latest Aragonian. Nevertheless, Dorcatherium is very rare in ACM and displays a very patchy chronological distribution. Together with the similarly spotty distribution of both primates and castorids along the ACM series, this suggests that very humid conditions might have been only intermittently present in this area throughout the latest Aragonian.     

New remains of Dorcatherium crassum (Artiodactyla: Tragulidae) from the Early Miocene (MN4) of Els Casots (Subirats,Vallès-Penedès Basin)

New tragulid dental remains from the late Early Miocene (16.5–16.3 Ma, MN4) locality of Els Casots (Vallès-Penedès Basin, Catalonia, Spain) are described. This sample fits well, both in size and occlusal morphology, with the material of Dorcatherium crassum from the type locality (Sansan, France; MN6). We therefore attribute the tragulid from Els Casots to this species, also in agreement with its known chronostratigraphic range throughout Europe (MN4–MN6) and the previous records of this species from other MN4 localities of the Vallès-Penedès Basin. The described remains – which include the postcanine lower deciduous and permanent dentition, as well as several upper cheek teeth – constitute the most complete sample of D. crassum from the Iberian Peninsula. The presence of this species at Els Casots is consistent with the lacustrine depositional environment inferred from sedimentological evidence and associated fauna, and further confirms the nearby presence of densely forested environments with a humid climate with low seasonality.     

Ruminants from the Early Miocene of Napak,Uganda

Since the late 1950's, many ruminant specimens have been recovered from the Early Miocene volcano-sedimentary succession at Napak, Uganda. Beginning in 1958 when the first fossils were recognised by J.G. Wilson, and the subsequent surveys of Bishop and colleagues, more than 340 specimens have been found. Preliminary publications in the 1960's identified three species, Dorcatherium songhorensis, Dorcatherium ? chappuisi, and Palaeomeryx africanus. Napak material hitherto identified as Dorcatherium ? chappuisi represents a new species of tragulid smaller than D. chappuisi, and is here identified as Dorcatherium iririensis nov. sp., while material previously assigned to Palaeomeryx is transferred to Walangania. A new finding for the site is the tiny tragulid, Dorcatherium parvum. There are thus four species of ruminants at Napak.     

The scimitar-toothed cat Machairodus aphanistus (Carnivora: Felidae) in the Vallès-Penedès Basin (NE Iberian Peninsula)

Here, we revise all the published and unpublished scimitar-toothed cat remains from the Vallès-Penedès Basin (NE Iberian Peninsula), in order to confirm their taxonomic attribution to Machairodus aphanistus as well as to provide more precise information about its chronological distribution in this basin. The studied material (including dentognathic as well as postcranial remains) comes from the following localities: Can Mata indeterminate (late MN7+8 or MN9), Creu Conill 22 (MN9), Can Poncic 1 (MN9), Can Llobateres 1 (MN9), Santiga (MN9), La Tarumba 1 (MN10), Viladecavalls (MN10), Ronda Oest Sabadell ROS-D3 (MN10), and Torrent de Febulines (MN10). Most of the studied material fits well with the morphologic and metrical features characteristic of the Vallesian species M. aphanistus, with the exception of the remains from Creu Conill 22 (an undescribed partial P4 formerly attributed to this taxon), which belongs in fact to a medium-sized hyaenid. From a biostratigraphic viewpoint, the removal of the Creu Conill material from the hypodigm of M. aphanistus has important implications, because this locality (11.1 Ma) was considered to record the first appearance datum of this taxon in the Vallès-Penedès Basin. However, the report of a previously unpublished talus from Can Mata (late MN7+8 or MN9) indicates that this taxon was present in this basin at least by the earliest Vallesian. Therefore, our results indicate that the first appearance datum of Machairodus in the Vallès-Penedès Basin might be somewhat younger than previously assumed, although dating uncertainties for the Can Mata remains preclude a more precise assessment. In contrast, the new mandibular remains from ROS-D3 (MN10) are likely coeval with those from La Tarumba 1, with the last appearance datum of M. aphanistus in the Vallès-Penedès Basin corresponding to Torrent de Febulines (ca. 9.1 Ma). The postcranial material described from various Vallès-Penedès localities further indicates that M. aphanistus displayed less cursorial adaptations than its purported descendant Homotherium.     

The heteromorph ammonite genus Ammonitoceras Dumas, 1876 (Ancyloceratidae) in the lower Aptian (Lower Cretaceous) of the Les Ferres Aptian Basin (southeastern France)

Ammonitoceras Dumas, 1876 is a genus of heteromorph ammonites characterized by the presence of a peculiar ontogenetic stage in its inner whorls: the Ammonitoceras stage. But in spite of its wide paleogeographic and biostratigraphic extension throughout the Aptian (Lower Cretaceous), this genus remains poorly known. In the present work we study specimens of Ammonitoceras from the lower Aptian Deshayesites multicostatus (Deshayesites deshayesi Zone) to Dufrenoyia furcata (Dufrenoyia furcata Zone) subzones of the Les Ferres Aptian Basin (southeastern France). The results are as follows: (1) representatives of Ammonitoceras from this area are regarded dimorphic with criocone macroconchs and ancylocone microconchs, (2) their ontogenetic sequence is described, (3) their intraspecific variability is significant and concerns the adult size and the duration of the ontogenetic stages, especially the Ammonitoceras stage, (4) two species are recognized: the earlier Ammonitoceras ucetiae Dumas, 1876, characterized by a brief Ammonitoceras stage on average, and the latter Ammonitoceras lahuseni (Sinzow, 1906), characterized by a longer Ammonitoceras stage on average. The sample of the Deshayesites grandis Subzone (Deshayesites deshayesi Zone) is composed of specimens too fragmentary to be identified at species level.     

The coralline genera Sporolithon and Heydrichia (Sporolithales,Rhodophyta) clarified by sequencing type material of their generitypes and other species

Interspecific systematics in the red algal order Sporolithales remains problematic. To re‐evaluate its species, DNA analyses were performed on historical type material and recently collected specimens assigned to the two genera Sporolithon and Heydrichia. Partial rbcL sequences from the lectotype specimens of Sporolithon ptychoides (the generitype species) and Sporolithon molle, both from El Tor, Egypt, are exact matches to field‐collected topotype specimens. Sporolithon crassum and Sporolithon erythraeum also have the same type locality; material of the former appears to no longer exist, and we were unable to PCR amplify DNA from the latter. A new species, Sporolithon eltorensis, is described from the same type locality. We have not found any morpho‐anatomical characters that distinguish these three species. No sequenced specimens reported as S. ptychoides from other parts of the world represent this species, and likely reports of S. ptychoides and S. molle based on morpho‐anatomy are incorrect. A partial rbcL sequence from the holotype of Sporolithon dimotum indicates it is not a synonym of S. ptychoides, and data from the holotype of S. episporum confirm its specific recognition. DNA sequences from topotype material of Heydrichia woelkerlingii, the generitype species, and isotype material of Heydrichia cerasina confirm that these are distinct species; the taxon reported to be H. woelkerlingii from New Zealand is likely an undescribed species. Type specimens of all other Sporolithon and Heydrichia species need to be sequenced to confirm that they are distinct species; morpho‐anatomical studies have proved inadequate for this task.     

Extant silicoflagellates from the Northeast Aegean (eastern Mediterranean Sea): Morphologies and double skeletons

Silicoflagellate abundance, vertical distribution and morphology were studied during spring (March 2014) at three sampling stations located in the Northeast Aegean Sea adjacent to the Dardanelles Strait and characterized by a variable influx of cold, low-salinity Black Sea water. The silicoflagellate assemblage was dominated by Dictyocha stapedia and Stephanocha speculum with minor contribution of D. aculeata and Octactis pulchra. While specimens of D. stapedia were represented by the typical morphologies described in other areas of the Mediterranean Sea, populations of S. speculum displayed peculiar characters: they were large, predominantly 7-sided, with a small apical ring as well as apical ring spines, concave basal ring sides and non-rotated apical structure. Some of these features have been described for S. speculum at high latitudes, but the combined characters make these specimens slightly different from the high latitude populations. Similar morphologies have been observed in the western Black Sea, thus we can infer that the peculiar specimens detected in the Northeast Aegean are associated with the influx of Black Sea water masses.     

Tragoportax cf. rugosifrons (Schlosser, 1904) from the late Miocene of Cessaniti (Southern Italy)

This paper describes remains attributable to Tragoportax cf. rugosifrons (Schlosser, 1904) found in the late Miocene site of Cessaniti (Vibo Valentia, Calabria) and the surrounding area. The studied specimens come from the Clypeaster sandstones, included in a marine/fluvial succession dated between 8 and 7.2 Ma. At Cessaniti, Tragoportax is associated with Stegotetrabelodon syrticus Petrocchi, 1941; Samotherium cf. boissieri Forsyth-Major, 1888; Bohlinia cf. attica Matthew, 1929; and an undetermined Rhinocerotid still under study. The genus Tragoportax was common in Eurasia and Africa during the late Miocene. The occurrence of Tragoportax cf. rugosifrons at Cessaniti confirms the peculiarity of the assemblage, with its association of species of North African and Pikermian (Greco-Iranian bioprovince) affinities.     

New Alicornops (Rhinocerotidae) remains from Lower and Middle Siwaliks,Pakistan

Unpublished rhinocerotid remains from the Lower and Middle Siwaliks of Pakistan are described in this paper and recognized as two species of Alicornops. Alicornops complanatum (Heissig) is identified in the Dhok Pathan Formation of the Middle Siwaliks and Alicornops laogouense Deng in the Kamlial Formation of the Lower Siwaliks. The Dhok Pathan Formation levels with A. complanatum are roughly correlated with the late Miocene-Pliocene European mammal zones MN10-15. In turn, levels with A. laogouense of the Kamlial Formation would correlate with the middle–late Aragonian (middle Miocene) European MN5. The recognition of the Chinese species A. laogouense in the Potwar Plateau represents the first discovery of this taxon in Pakistan and increases the geographical and stratigraphic distributions of this species, and adds to the rhinocerotid association from the Siwaliks. In turn, the presence of A. complanatum in the Siwaliks of Potwar Plateau also enlarges its geographic distribution in Pakistan, as it was previously known from the Bugti Hills of Balouchistan. The absence of Alicornops from the Siwaliks in the Chinji and Nagri formations (between late MN5 and MN9 zones) might be due to an inadequate fossil record, as other rhinocerotid species are known from Kamlial to Dhok Pathan formations. However, the two recorded species of Alicornops could also reveal two independent migration waves as supported by the appearance of other taxa in different formations. A summary of fossil Cenozoic rhinocerotids from different areas of Pakistan is also presented.     

Taxonomy of subgenus Temnoclemmys Bergounioux, 1958 (Testudines: Geoemydidae: Ptychogasterinae) based on new material from the Vallès-Penedès Basin (NE Iberian Peninsula)

The taxonomy of the freshwater turtle subgenus Ptychogaster (Temnoclemmys) (Testudines, Geoemydidae, Ptychogasterinae) is revised based on new and previously-published material from the Vallès-Penedès Basin (NE Iberian Peninsula). Additional remains of subgenus Ptychogaster, first reported from this basin, are also described. Although the oldest remains of Temnoclemmys are left unassigned, most of the available remains are attributed to its type species, Ptychogaster (Temnoclemmys) batalleri, originally described from the Late Miocene (MN9 and MN10) of this basin. The new material expands the chronostratigraphic range of P. batalleri in the Vallès-Penedès Basin back to the MN7 + 8 and shows that “Testudo” celonica is its junior synonym. Emended diagnoses for Ptychogaster and Temnoclemmys are also provided.     

New anthropological data from Cussac Cave (Gravettian,Dordogne, France): In situ and virtual analyses of Locus 3

Cussac Cave presents a unique combination of parietal art and several hundred parts of scattered human remains, dated to the Middle Gravettian (29–28,000 cal BP). The cave is protected as a National Heritage site. As a result, only noninvasive bioanthropological analyses are allowed, consisting of in situ observations and the study of 3D models obtained by photogrammetry. Here we present the first results of these analyses of the human remains from Locus 3. Only 65 of the 106 human skeletal fragments and bones could be firmly identified. Virtual analyses were carried out on 3D models of 16 skeletal elements so that osteometric data could be provided. Despite the limitations inherent in studying commingled remains and those specific to Cussac Cave, the search for virtual pair-matching, articular congruence, and osteometric sorting allowed the allocation of twelve bones to three individuals, one late adolescent and two adults.     

First record of late middle Miocene Moschidae from Turkey: Micromeryx and Hispanomeryx from Catakbağyaka (Muğla,SW Turkey)

Here, we describe remains of Micromeryx sp. and Hispanomeryx sp. from the middle Miocene locality Catakbağyaka (MN 7/8). This is the first record for Miocene Moschidae for the late middle Miocene in Turkey and verifies the importance of Turkey as a corridor for the dispersal of Eurasian mammals. Furthermore, the record from Catakbağyaka confirms that the sympatric co-occurrence of two Miocene moschid taxa in one locality appears to be a common phenomenon.     

Hrabalicrinus zitti gen. et sp. nov., and other Upper Jurassic crinoids (Echinodermata,Crinoidea) from the Brno area (Czech Republic)

Jurassic (Oxfordian) crinoids from the Brno area (Czech Republic) are described in details for the first time. A rich crinoid assemblage consisting of cups, isolated cup elements, brachial plates, columnals, pluricolumnals, and cirrals is assigned to isocrinids Isocrinus amblyscalaris (Thurmann), Balanocrinus subteres (Münster), B. pentagonalis (Goldfuss), Isocrinida indet., comatulid Hrabalicrinus zitti gen. et sp. nov., cyrtocrinids Lonchocrinus sp., Cyrtocrinus cf. nutans (Goldfuss), Pilocrinus moussoni (Desor), Tetracrinus moniliformis Münster, Cyrtocrinida indet., and millericrinids (Millericrinida indet.). Crinoids already mentioned from the Jurassic strata of the Czech Republic are subsequently revised. Apart from the isolated remains of Isocrinida, Millericrinida, and Thiolliericrinida, the presence of any other taxon reported from this area should be treated with extreme caution. A rare example of non-regenerative columnal healing (the so-called callus) in I. amblyscalaris is also described. Based on sedimentology and microfacies, the Jurassic limestones were deposited in various palaeoenvironments of upper carbonate platform and shelf lagoon (0 to > 50 m palaeodepth).     

Factors of intraspecific variability in ammonites,the example of Gassendiceras alpinum (d’Orbigny, 1850) (Hemihoplitidae,Upper Barremian)

The homogeneity of a large sample of Gassendiceras alpinum is tested statistically. The quantitative analysis did not reject the null hypothesis and the homogeneity of the sample is accepted. As a result, the different taxa used in the literature (G. alpinum, “Barrancyloceras barremense”, Leroyiceras mascarellii and Spinocrioceras sauvanae) all fall into the variability of a single species, although some doubt subsist for L. dolloi. Taxonomically, by the priority principle the only name that should be used is G. alpinum. The results and other examples about the intraspecific variability confirm the importance of the Westermann's laws in ammonites, but they show that there are other important sources of covariation. In Gassendiceras the variability is tripolar (both the classic robust-slender poles of the Westermann's laws plus a third morphological pole with thick section and not so robust ornamentation), which implies that the intraspecific variability is not always uniform or consistent from one group to another, or in any case do not necessarily respond to the same factors in equivalent proportions. Heterochronies, erasing of the ornamentation and uncoiling of the heteromorph ammonites are also considered as factors of the intraspecific variability.     

Late Miocene (Turolian,MN13) squirrels from Moncucco Torinese,NW Italy

Squirrel remains are rare in the fossil record, which hampers a properly documented interpretation of their evolutionary history. The Late Miocene sciurids from Moncucco Torinese (NW Italy) are described based on a diverse tooth assemblage. The data presented herein confirm the presence of Sciurus warthae and Pliopetaurista pliocaenica, and also include the first unquestionable occurrence of Hylopetes hungaricus plus an indeterminate Sciurinae. The report of Sciurus warthae from Moncucco Torinese represents the earliest record of the genus Sciurus in Europe. Moreover, Moncucco Torinese is the first European locality recording the co-occurrence of Sciurini and Pteromyini in the Late Miocene. Comparative analyses support a close relationship between S. warthae and Sciurus vulgaris corroborating the hypothesis of an earliest divergence of the “Sciurus vulgaris clade” in the Late Miocene. Biochronological, paleobiogeographical and paleoecological implications are also discussed.