Blarinoides aliciae sp. nov., a new Soricidae (Mammalia,Lipotyphla) from the Pliocene of Spain

A new species of Blarinoides (Soricidae, Mammalia), Blarinoides aliciae, is described from the Pliocene of Tollo de Chiclana (Guadix Basin, southeastern Spain). It is notably smaller than the single species of this genus known until now, B. mariae. B. aliciae is recorded from several other Spanish localities, but the sample from Tollo de Chiclana-3 is the most numerous. B. aliciae proves the simultaneous existence of two different species of Blarinoides in Europe during the Pliocene: one of large size in central and eastern Europe, and another one of small size in Iberia.     

Arvicolidae (Rodentia) from the Pliocene of Tollo de Chiclana (Granada, SE Spain)

Several new fossiliferous Pliocene localities have been identified in the continental deposits of the area of Tollo de Chiclana (Guadix Basin, SE Spain), that have yielded a rich rodent and insectivore fauna. In this paper, we study the Arvicolidae from these localities. Remains adscribed to the genera Dolomys, Mimomys and Kislangia have been found, which are very interesting from a biostratigraphical point of view.     

A juvenile skull of Acerorhinus yuanmouensis (Mammalia: Rhinocerotidae) from the Late Miocene hominoid fauna of the Yuanmou Basin (Yunnan,China)

A unique juvenile skull bearing both milk premolars and unerupted but fully developed permanent premolars and molars (observed using X-ray microcomputed tomography), and some isolated upper cheek teeth, all from the Late Miocene hominoid fauna of the Yuanmou Basin (Yunnan, China), closely resemble craniodental material of Acerorhinus yuanmouensis Zong, 1998 from the same locality, and are referred to this species. A phylogenetic analysis based on 214 craniodental morphological characters scored for 31 terminal taxa reveals that A. yuanmouensis should be assigned to the genus Acerorhinus indeed. The newly discovered specimens improve our understanding of this species, especially with respect to the morphology of the milk premolars and premolars. Two intraspecific variations in the upper premolars are noted: a lingual bridge may be present or absent, and the lingual cingulum continuous or reduced. The analysis also indicates that: the phylogenetic status of Acerorhinus lufengensis Deng and Qi, 2009 should be reconsidered; “Aceratherium” huadeensis Qiu, 1979 does neither belong to Aceratherium nor Acerorhinus, and its phylogenetic status remains debatable.     

New cranial material of Shansirhinus (Rhinocerotidae, Perissodactyla) from the Lower Pliocene of the Linxia Basin in Gansu, China

A well-preserved skull and articulated mandible of Shansirhinus, a horned aceratherine rhinocerotid, is described from an Early Pliocene (Gaozhuangian; approximately 5.3-4.34 Ma) locality of the Linxia Basin in Gansu, China. Comparing the new material of Shansirhinus ringstromi Kretzoi, 1942, we confirm the synonymy proposed previously for the species Chilotherium yunnanensis Tang et al., 1974, Chilotherium cornutum Qiu and Yan, 1982, and Ch. tianzhuensis Zheng, 1982. New knowledge concerning its cranial and mandibular morphology allows insight into its phylogenetic position among aceratherine rhinocerotids. A sister group relationship between Shansirhinus and Chilotherium is proposed on the basis of cranial, mandibular and dental evidence. Shared derived characters that support this relationship include: an expanded mandibular symphysis with a concave ventral surface; retracted premaxillae lacking upper incisors; a robust and right-angled facial crest; a flat or slightly concave dorsal skull profile; a weak or absent parastyle fold; and a constricted protocone on the premolars. Shansirhinus was probably a grazer, which is evidenced by the high crown, strong wear, well-developed secondary folds, and enamel plications on its teeth. The age of S. ringstromi is Late Miocene to the Early Pliocene, corresponding to the Turolian to Ruscinian, MN12-MN15 of Europe. S. ringstromi is likely the ancestor of the more advanced S. brancoi (Schlosser, 1903).     

A new Ruscinian site in Europe: Baza-1 (Baza basin,Andalusia, Spain)

The Guadix–Baza depression (southeastern Spain) preserves one of the best continental Plio-Pleistocene records of the European continent. The new site, Baza-1, is the first Ruscinian locality with fauna of large vertebrates known in the basin. During the summer seasons of 2001, 2002, 2015 and 2016, systematic excavations were undertaken in the site over an area of 25 m², which provided > 400 fossil remains of Ruscinian mammals. The faunal assemblage is diverse, comprising 17 genera. Rodentia are represented by Ruscinomys sp., Apocricetus barrierei, Debruijnimys julii, Apodemus gorafensis, Castillomys gracilis, Occitanomys cf. brailloni, Paraethomys meini, Paraethomys aff. abaigari, Stephanomys cordii, Trilophomys cf. castroi and Eliomys aff. intermedius. Large mammals are represented by two proboscideans, Anancus arvernensis and Mammut borsoni, the rhino Stephanorhinus sp. cf. S. jeanvireti, the equid cf. Hipparion sp., two bovids, a large sized Alephis sp. and a small-to-middle sized Bovidae indet. (cf. Antilope sp.), and a middle sized deer, Cervinae indet. This record is completed by the presence of a chelonid, Testudinae indet. Biostratigraphic data from micro and macromammals suggest an age for the assemblage between 4 and 4.5 Ma.     

New data on Mio-Pliocene Sciuridae (Rodentia,Mammalia) from southern Spain

Sciurids are very scarce in the fossil record, especially in the basins of southern Spain. The aim of this paper is to review the Sciuridae record in these basins. The Granada and Guadix basins have yielded specimens of Xerini and Pteromyinae, which represent the largest collection of fossil Sciuridae in southern Spain from the Middle Turolian to the Upper Ruscinian. The new discoveries change the currently known geographical and temporal range of some taxons, since we find the oldest evidence of Pliopetaurista pliocaenica in localities from the Late Turolian and of Heteroxerus mariatheresae in a locality of the Middle Turolian. Furthermore, we record the first evidence of Atlantoxerus margaritae in southern Spain.     

Palaeeudyptes klekowskii, the best-preserved penguin skeleton from the Eocene–Oligocene of Antarctica: Taxonomic and evolutionary remarks

A new fossil penguin skeleton from the La Meseta Formation collected at the locality DPV 13/84 (Seymour Island, Antarctic Peninsula) from the crinoid horizon located 40 m above the base of the 145 m-thick Submeseta Allomember (Late Eocene–Early Oligocene?) is described. The specimen is assigned to the species Palaeeudyptes klekowskii Myrcha, Tatur and del Valle, 1990; it is the most complete penguin skeleton ever recovered from Antarctica. Discoveries like this one are significant for the study of the anatomy and evolution of penguins, in particular regarding the Antarctic species included in the genus Palaeeudyptes Huxley, 1859. P. klekowskii closely resembles its smaller congeneric species P. gunnari ( Wiman, 1905), with only the relative concavity of the margo medialis distinguishing the tarsometatarsi of both taxa. However, the results of a geometric morphometric analysis show some intra- and inter-specific variations, making possible the systematic assignment of the majority of the specimens. Size variation is congruent with the presence of two different species.     

A new Late Cretaceous (Maastrichtian) serial planktic foraminifer (Family Heterohelicidae) with early planispiral coil and revision of Spiroplecta Ehrenberg, 1844

A new planktic foraminifer, Hartella harti nov. gen., nov. sp. is described from the Maastrichtian sediments of the Atlantic Ocean. H. harti likely evolved from Fleisherites glabrans (Cushman). Spiroplecta Ehrenberg is reviewed and considered monospecific. The only species assigned to this genus is Spiroplecta americana Ehrenberg, which evolved from Heterohelix globulosa (Ehrenberg). It is demonstrated that the early planispiral coil developed in at least three separate lineages of serial planktic foraminifera in the Late Cretaceous (late Campanian-early Maastrichtian).     

The earliest European badger (Meles meles), from the Late Villafranchian site of Fuente Nueva 3 (Orce, Granada, SE Iberian Peninsula)

Paleontological and molecular data suggest that the divergence of the European and Asian badgers (Meles meles and Meles leucurus, respectively) from their ancestor Meles thorali might have taken place in the Middle to Late Villafranchian boundary (ca. 1.8 Ma). However, the available record of Late Villafrachian European badgers is scanty and poorly known, and this hypothesis needs more paleontological data in order to be tested. The badger hemimandible from the Iberian locality of Fuente Nueva 3, a locality placed in the beginning of the Late Villafranchian, supports the idea of an early divergence between European and Asian badgers.     

Cytogenetics of the Brazilian Bolitoglossa paraensis (Unterstein, 1930) salamanders (Caudata,Plethodontidae)

Plethodontid salamanders of genus Bolitoglossa constitute the largest and most diverse group of salamanders, including around 20% of living caudate species. Recent studies have indicated the occurrence of five recognized species in the Brazilian Amazon Rainforest. We present here the first cytogenetic data of a Brazilian salamander, which may prove to be a useful by contribution to the cytotaxonomy of the genus. Specimens were collected near the “type” locality (Utinga, Belém, PA, Brazil). Chromosomal preparations from duodenal epithelial cells and testes were subjected to Giemsa staining, C-banding and DAPI/CMA₃ fluorochrome staining. All specimens showed a karyotype with 13 bi-armed chromosome pairs (2n = 26). Nucleolar Organizer Regions, evidenced by CMA₃, were located distally on the long arm of pair 7 (7q). DAPI+ heterochromatin was predominantly centromeric, with some small pericentromeric bands. Although the C-banding patterns of other Bolitoglossa species are so far unknown, cytogenetic studies conducted in other Plethodontid salamanders have demonstrated that pericentromeric heterochromatin is a useful cytological marker for identifying interspecific homeologies. Species diversification is usually accompanied by chromosomal changes. Therefore, the cytogenetic characterization of Bolitoglossa populations from the middle and western Brazilian Amazon Basin could identify differences which may lead to the identification of new species.     

Prostrepsiceros and Protragelaphus (Artiodactyla,Mammalia) from the Late Miocene locality of Chomateri (Attica,Greece)

The Late Miocene fossiliferous locality of Chomateri is located close to the classic locality of Pikermi, but unfortunately its faunal context is not sufficiently known. Some fossil remains of spiral-horned antelopes from Chomateri are assigned to Prostrepsiceros rotundicornis and Protragelaphus skouzesi. The co-existence of P. rotundicornis and P. skouzesi is relatively rare in the Late Miocene, but well-documented in Pikermi. Comparisons of the Chomateri material support a middle Turolian age for the Chomateri locality and suggest a close chronological relation to the Pikermi fauna.     

A revision of chamaeleonids from the Lower Miocene of the Czech Republic with description of a new species of Chamaeleo (Squamata, Chamaeleonidae)

A revision of Chamaeleo caroliquarti Moody and Ro?ek is presented. The comparisons of the holotypic left dentary with those of specimens subsequently assigned to C. caroliquarti and of the Recent species of Chamaeleo, Furcifer and Calumma is carried out. It is shown that the type dentaries of C. caroliquarti include two different morphotypes with the absence of unique features. Within the Recent chameleons, the exact determination of the individual species merely on the basis of the dentaries is impossible. The holotypic dentary of C. caroliquarti is basically identical with that of C. calyptratus. However, the same morphology of the dentary as present in C. caroliquarti is also present in other species of different genera such as Calumma globifer and Furcifer pardalis. The paratypic dentaries of C. caroliquarti have a different morphology to the holotype and are indistinguishable from that in the Recent C. chamaeleon. On the other hand, a new species of the genus Chamaeleo, C. andrusovi, is described on the basis of isolated cranial elements, which possess clear autapomorphic features. This material comes from the Lower Miocene (Ottnangian) zone MN 4 in the Dolnice locality of the Czech Republic, and it differs from Recent and fossil chameleons in the following combination of characters: (1) its typically developed strongly pustular ornamentation and its distribution on the external surfaces of the skull roofing bones; (2) the frontoparietal suture is digitiform with a well-developed, anteriorly directed mesial spine, and (3) the parietal bone narrows posteriorly at its midlength, it is not bowed dorsally and it does not contribute posteriorly to a dorsal sagittal crest. This new material expands our knowledge of the cranial anatomy of Lower Miocene chameleons.     

Morphometric analysis of six Gerbillus species (Rodentia,Gerbillinae) from Tunisia

Size and shape changes in the skull of the genus Gerbillus were investigated using geometric morphometrics. Six species from Tunisia were studied (G. gerbillus, G. campestris, G. nanus, G. tarabuli, G. simoni and G. latastei). Statistical analyses of shape variability allowed us to discriminate three morphological groups which are congruent with the three groups suggested by previous morphological and molecular studies. However, our results contrast with previous molecular investigations. In fact, according to results obtained by the use of principal component analysis, canonical variate analysis and UPGMA, we found a higher degree of divergence between the subgenus Dipodillus and the other two subgenera Gerbillus and Hendecapleura. This fact suggests that the morphometric differences observed among species within the genus Gerbillus are not mainly related to phylogeny. To reconciliate the molecular and morphological approaches, we propose a hypothesis of differential rates of phenotypic evolution in the genus Gerbillus. In this view, the species belonging to the subgenus Dipodillus evolved apomorphic features of the skull likely related to a higher degree of habitat specialization. By contrast, the more generalist Gerbillus and Hendecapleura subgenera show less differentiated plesiomorphic morphology.     

Review of the fossil pufferfish genus Archaeotetraodon (Teleostei, Tetraodontidae), with description of three new taxa from the Miocene of Italy

Three new species of the extinct pufferfish genus Archaeotetraodon are described from the Middle to Upper Miocene of Italy (A. bannikovi nov. sp., A. dicarloi nov. sp. and A. zafaranai nov. sp.), and compared in detail with the three previously described taxa of the genus [A. jamestyleri (Bannikov), A. winterbottomi Tyler and Bannikov and A. cerrinaferoni Carnevale and Santini], which are known, respectively, from the Miocene of Ukraine, the Oligocene of Russia, and the Miocene of Algeria. The three new species described herein bring the number of known Archaeotetraodon species to six, thereby making it by far the most speciose extinct genus of the order Tetraodontiformes. The monophyly of Archaeotetraodon is supported by two derived conditions: bifid scale spinules (versus single spinules in all other tetraodontids), and the fusion of the haemal spine of the penultimate vertebra to the centrum (versus autogenous in all other tetraodontids). We compare a large suite of morphological features of Archaeotetraodon to those found within the group of more basal tetraodontid genera to which we believe it belongs, but we are unable to determine the closest relationships of Archaeotetraodon within that group. Morphofunctional and paleoenvironmental considerations suggest that the species of the genus Archaeotetraodon were adapted to a pelagic or deep-sea lifestyle and were probably able to tolerate relatively low oxygen concentrations. The entire fossil record of the family Tetraodontidae is discussed.     

Reassessment of the taxonomic position of Iranocypris typhlops Bruun & Kaiser, 1944 (Actinopterygii,Cyprinidae)

The Iranian cave barb (Iranocypris typhlops Bruun & Kaiser, 1944) is a rare and endemic species of the family Cyprinidae known from a single locality in the Zagros Mountains, western Iran. This species is “Vulnerable” according to the IUCN Red List and is one of the top four threatened freshwater fish species in Iran. Yet, the taxonomic position of I. typhlops is uncertain. We examined phylogenetic relationships of this species with other species of the family Cyprinidae based on the mitochondrial cytochrome b gene. Our results show that I. typhlops is monophyletic and is sister taxon of a cluster formed by Garra rufa (Heckel, 1843) and Garra barreimiae (Fowler & Steinitz, 1956) within a clade that includes other species of the genus Garra. Based on previous molecular and morphological studies, as well as our new results, we recommend that I. typhlops should be transferred to the genus Garra Hamilton, 1822.     

Canis accitanus nov. sp., a new small dog (Canidae, Carnivora, Mammalia) from the Fonelas P-1 Plio-Pleistocene site (Guadix basin, Granada, Spain)

This paper reports a new species of dog (Canis accitanus nov. sp.) from the Fonelas P-1 site (dated close to the Plio-Pleistocene boundary) in Granada, Spain. This new taxon shows cranial features more similar to coyote-like dogs (C. lepophagus, C. priscolatrans, C. arnensis or C. latrans) than to wolf-like dogs (C. etruscus, C. mosbachensis or C. lupus), such as a long and narrow muzzle, a little-developed sagittal crest and frontal bones raised only a little above the rostrum. However, it also shows a series of autapomorphic characteristics in its upper dentition, essentially in the first upper molar, which reflects a trophic adaptation towards a more abrasive diet than that eaten by other species of its genus. This new dog is the smallest representative of the genus Canis ever recorded for the European Pliocene or Pleistocene.     

The origin of bird pollination in Macaronesian Lotus (Loteae, Leguminosae)

The four red-flowered, apparently bird-pollinated Lotus species from the Canary Islands have previously been classified in their own genus, Rhyncholotus. Currently, they are considered as a separate section within genus Lotus, distinct from other herbaceous Canarian congeners which are yellow-flowered and bee-pollinated. A combined analysis of four nuclear regions (including ITS and three homologues of CYCLOIDEA) and three plastid regions (CYB6, matK and trnH-psbA) nests the four bird-pollinated species within a single extant species of bee-pollinated Lotus (L. sessilifolius), in a very extreme example of species paraphyly. Therefore, our data compellingly support the hypothesis that the Macaronesian Lotus species with a bird pollination syndrome are recently derived from entomophilous ancestors. Calibration of the phylogenetic trees using geological age estimates of the most recent islands (La Palma and El Hierro) together with oldest ages of Fuerteventura indicates that bird pollination evolved ca. 1.7 Ma in the Canarian Lotus. These four bird-pollinated species share a most recent common ancestor (MRCA) with L. sessilifolius that dates to about 2.2 Ma. Our analyses further suggest that the evolution of the bird pollination syndrome was likely triggered by the availability of new niches in La Palma and Tenerife as a result of recent volcanic activity.