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.     

The anterior process of the malleus in extant Lagomorpha (Mammalia)

The anterior anchoring of the malleus of 30 extant species of Lagomorpha (rabbits, hares, pikas) has been studied on the basis of histological serial sections and µCT‐scans. It is shown that former studies of Oryctolagus, Lepus, and Ochotona are incomplete, because the rostral part of the processus anterior of the malleus is always lacking due to damage of this extremely delicate structure. Our study shows that in perinatal stages of Leporidae the praearticulare develops a prominent processus internus that fits into a groove at the ventral side of the tegmen tympani; this “tongue and groove”‐arrangement may act as a hinge. In adult stages, the rostral end of the praearticulare fuses synostotically with the medial process of the ectotympanic. Torsional strain produced by rotation around the axis of the middle ear ossicles at sound transmission must, therefore, be experienced by the extremely thin but highly elastic bony pedicle of the processus internus praearticularis. The free ending processus anterior of a late fetal Ochotona shows a short processus internus praearticularis, which does not articulate with the tegmen tympani. During postnatal development the middle ear of Ochotona becomes considerably remodelled: not only does excessive pneumatization of the tegmen tympani and tympanic cavity wall occur, but the short processus anterior is fused synostotically to a bone trabecula of the tegmen tympani meshwork. The thin and elastic bone bridges are not equivalent in Leporidae and Ochotonidae, that is, they must have evolved convergently. Fleischer's classification with Oryctolagus possessing a “freely mobile type” of middle ear ossicles cannot be supported by our observations. The same holds true for Ochotona, which does not represent a “freely mobile type” either. Thus, we suggest for the lagomorph middle ear ossicles a new category: the “bone elasticity type.”     

A new species of Orycteropodidae (Mammalia, Tubulidentata) in the Mio-Pliocene of northern Chad

A new species of fossil Tubulidentata has been found by the Mission Paléoanthropologique Franco-Tchadienne in Northern Chad. It is the first fossil Orycteropodidae (aardvark) from the Mio-Pliocene of Central Africa. The new taxon, Orycteropus abundulafus sp. nov. , is considered in the framework of the available Orycteropodidae fossil record. The Chadian specimen is characterized by the highest dental robustness index among all Tubulidentata, the presence of crests on the pterygoid, the triangular-shaped olecranon fossa and the reduction of the deltoid crest. All of these characters are linked to a less fossorial animal that had a tougher diet. This new African species is closer to the Eurasian O. gaudryi than to any other Tubulidentata. Together they form a clade distinct from that which includes O. afer . This is the first evidence of a relationship for aardvarks between Africa and Eurasia. An initial step is made towards revision of the phylogeny of the order.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 109–131.     

Effects of altitudinal change on the auditory bulla in Ochotona daurica (Mammalia, Lagomorpha)

The lengths and widths of body parts are influenced to a considerable extent by the environment. Along with increasing altitude, wings of arthropods degenerated or even disappeared so that a trend of degeneration on their acoustic systems appeared. We measured 131 adult skulls of daurian pikas, Ochotona daurica , in order to know whether the degeneration will happen in mammals. Our results show that both the size and relative size of auditory bulla of daurian pikas were negatively correlated with altitude and this change was significant. However, the ear length showed an opposite trend. We suggest that lower oxygen concentration, constantly strong winds and lack of resources at high altitude are responsible for the decrease in the size of the auditory bulla. Besides thermoregulation for large different temperatures of day and night in high altitude, larger external ears of the pika help detect the direction of the predator and could compensate for a relatively small bulla.     

Tooth Morphology in Fossil and Extant Lagomorpha (Mammalia) Reflects Different Mastication Patterns

The functional interpretation of the cheek teeth and the mastication cycle of Lagomorpha are deduced from various aspects of tooth morphology of fossil and extant species. Mastication is composed of an almost orthal shearing and transverse grinding in a lingual direction. Shearing blades are not only indicated by facets but as well by thickened enamel. A primary shearing blade (PSB) inherited from stem lagomorphs occurs in all examined species. It can be correlated with facets 1 and 5 (sensu Crompton 1971) and occurs in very few mammals; it is conspicuously absent in the sister-taxon Rodentia. A secondary shearing blade (SSB) occurs in derived Ochotonidae and two basal Leporidae (Romerolagus and Pronolagus) as a convergent pattern. In fossil ochotonids from Europe, the “lagicone structure” is gradually reduced in favor of the SSB. Thus, ochotonids strengthen the shearing ability, whereas most leporids favor the grinding function realized by the partial crenulation of the enamel band of the re-entrant folds. For the mastication cycle, the distinct phases were recognized, related to phase I of the tribosphenic model. The first movement (phase Ia) is directed almost orthally, the second (phase Ib) lingually. Only in Lepus europaeus was an additional phase detected, which might correspond to phase II.     

Eocene Lagomorpha (Mammalia) of Asia: 2. Strenulagus and Gobiolagus (Strenulagidae)

Two species of early lagomorph, Strenulagus solaris sp. nov. (Monogolia, Khaychin-Ula 2 and 3 localities, Middle Eocene) and Gobiolagus hekkeri (Kyrgyzstan, Andarak 2 locality, terminal Lower Eocene), are described. A revision of Romanolagus hekkeri Shevyreva from the Andarak 2 locality has shown that it belongs to the genus Gobiolagus; consequently, the monotypic genus Romanolagus Shevyreva, 1995 is a junior synonym of Gobiolagus Burke, 1941. The diagnoses and species compositions of the genera Strenulagus and Gobiolagus are revised. Zaissanolagus gromovi Erbajeva from the Lower Oligocene of eastern Kazakhstan is shown to be the latest representative of the genus Gobiolagus and the family Strenulagidae.     

Mitochondrial Cytochrome b Gene Sequence Diversity in the Korean Hare, Lepus coreanus Thomas (Mammalia, Lagomorpha)

Partial sequences of the mitochondrial cytochrome b gene of the Korean hare (Lepus coreanus) were analyzed to determine the degree of genetic diversity. Nine haplotypes were observed, and the maximum Tamura-Nei nucleotide distance among them was 2.8%, indicating that genetic diversity of L. coreanus is moderate. In order to clarify the Korean hare's taxonomic status and relationship with the Manchurian hare (L. mandshuricus) and the Chinese hare (L. sinensis), these nine haplotypes of the Korean hare were compared with 13 haplotypes from five other species of eastern Asian Lepus including L. mandshuricus and L. sinensis. The Korean hare was distinct in its cytochrome b gene, and it is confirmed that L. coreanus is a valid species, as noted by Jones and Johnson (1965, Univ. Kansas Publ. (Mus. Nat. Hist.) 16:357). Further analyses of mtDNA cytochrome b gene with additional specimens of L. coreanus from North Korea and other species of Lepus from eastern Asia are needed to clarify the taxonomic status of the divergent mtDNA clades of L. mandshuricus and L. sinensis.     

An analysis of the mole rats (Mammalia: Rodentia) from Langebaanweg (Mio-Pliocene, South Africa)

A study of the taphonomy and age profiles of the mole rat population from Langebaanweg ‘E’ Quarry was undertaken in order to see if these subterranean rodents had been accumulated by predation (coprocoenosis hypothesis) or as a result of drowning in their burrows during a flood event (catastrophocenosis hypothesis). Previous research has cited the bathyergids as being the most common rodent at Langebaanweg, however, a comprehensive study of new micromammal material indicates that murids, rather than mole rats, dominate most of the assemblages. The concentration of mole rat remains in many of the fossil-bearing levels at Langebaanweg is, nevertheless, still extremely high compared to other fossil sites or predation assemblages in the area. A taphonomic examination of mole rat incisor surfaces indicates that digestion occurs on between 44% and 54% of incisors in the different assemblages, and this, together with the degree of digestion suggests that category 1 and category 2 predators had been responsible for the accumulation of the fossil mole rat assemblages. The absence of rounding, polishing or weathering on the fossils indicates that there has been little or no transport by water, and that burial of the bones was rapid. Breakage and damage to the anterior portion of many of the mandibles made it impossible to accurately assess the age of the older mole rats, and the majority of mandibles could provide only a minimum age of the animal at the time of death. The age profile indicates that very young individuals are missing from the assemblages. This allows for refutation of the previously made suggestion that flooding was responsible for the death of the majority of Langebaanweg mole rats and also introduces the possibility that Bathyergus hendeyi may have been a social animal, unlike extant bathyergids which are solitary.     

Early Eocene lagomorph (Mammalia) from Western India and the early diversification of Lagomorpha

We report the oldest known record of Lagomorpha, based on distinctive, small ankle bones (calcaneus and talus) from Early Eocene deposits (Middle Ypresian equivalent, ca 53 Myr ago) of Gujarat, west-central India. The fossils predate the oldest previously known crown lagomorphs by several million years and extend the record of lagomorphs on the Indian subcontinent by 35 Myr. The bones show a mosaic of derived cursorial adaptations found in gracile Leporidae (rabbits and hares) and primitive traits characteristic of extant Ochotonidae (pikas) and more robust leporids. Together with gracile and robust calcanei from the Middle Eocene of Shanghuang, China, also reported here, the Indian fossils suggest that diversification within crown Lagomorpha and possibly divergence of the family Leporidae were already underway in the Early Eocene.     

New species of Amphilagus (Lagomorpha, Mammalia) from the Miocene of the Valley of Lakes, central Mongolia

New lagomorph species, Amphilagus orientalis sp. nov., A. plicadentis sp. nov., and A. magnus sp. nov. from Lower Miocene localities (Biozone D) of the Valley of Lakes in Central Mongolia are described.     

Protein variation and its systematic implications for the South African Leporidae (Mammalia:Lagomorpha)

Analysis of 9 genetic loci by horizontal starch gel electrophoresis was performed on 203 specimens of South African representatives of the leporid genera Lepus and Pronolagus . Three loci were found to be polymorphic within samples of the various species of both genera: transferrin (Tf-I), esterase-1 (Es-I) and esterase-2 (Es-II). Only one system, carbonic anhydrase (CA-I), proved useful in species identification, this being restricted to the genus Lepus . Based on this system all Lepus specimens analysed would be assigned to one of two groups corresponding to the species L. capensis and L. saxatils .     

Protein variation and its systematic implications for the South African Leporidae (Mammalia:Lagomorpha)

Analysis of 9 genetic loci by horizontal starch gel electrophoresis was performed on 203 specimens of South African representatives of the leporid genera Lepus and Pronolagus. Three loci were found to be polymorphic within samples of the various species of both genera: transferrin (Tf-I), esterase-1 (Es-I) and esterase-2 (Es-II). Only one system, carbonic anhydrase (CA-I), proved useful in species identification, this being restricted to the genus Lepus. Based on this system all Lepus specimens analysed would be assigned to one of two groups corresponding to the species L. capensis and L. saxatilis.     

Evolutionary morphology of the Tenrecoidea (Mammalia) carpal complex

The mammalian carpus can be difficult to interpret both phylogenetically and functionally. It is evolutionarily constrained in terms of functional morphology, yet there is considerable variation among many eutherian and metatherian lower and higher level taxa. The ecologically diverse Tenrecoidea (Mammalia) is a useful model for morphological interpretation of the interplay between function and phylogenetic constraint. Elements from the wrist and hand of 13 tenrecoid species, and one species each from Macroscelididae, Solenodontidae, and Erinaceidae, were compared to test form–function hypotheses of specific carpal, metacarpal, and phalangeal characters. Qualitative comparisons illustrate that several aspects of the tenrecoid carpus can be correlated with positional behaviour. Convergences within Tenrecoidea, and between tenrecoids and nontenrecoids with similar locomotor regimes, confirm a small number of carpal characters and a larger number of distal forearm, metacarpal, and phalangeal characters that reliably correspond with functional expectations. In addition, several features of the carpus appear to be phylogenetically constrained and indicate specific affinities within Tenrecoidea. Finally, there are a significant number of carpal features that vary among the studied taxa and remain ambiguous in terms of phylogenetic and/or functional significance.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 267–288.     

Phylogeny and taxonomic reassessment of pikas Ochotona pallasii and O. argentata (Mammalia,Lagomorpha)

We examined a cranial morphometric data set consisting of 186 specimens from the entire distribution range of Ochotona pallasii sensu lato and O. argentata, as well as 67 complete sequences of the COI gene and 28 sequences of the MGF and PRKCI introns from these and closely allied species. Our results show that the two allopatric morphologically similar taxa composing O. pallasii sensu lato – from Mongolia and adjacent territories and Kazakhstan – are paraphyletic relative to O. argentata. Genetic distances between these three taxa are larger than the intraspecific variation known for the subgenus Pika, in which the species under consideration belong; these distances are even larger than the interspecific differences among closely related species such as O. hyperborea, O. mantchurica and O. hoffmanni. Thus, the three focal taxa are recognized here as distinct species. Inspection of the type specimen of O. pallasii indicated that this specimen was not collected in Kazakhstan, has previously been theorized. The most probable place of the holotype's origin is Russian south‐eastern Altai (Chuyskaya Steppe); whatever its exact origin, it definitively originates from the ‘Mongolian’ taxon. Based on this evidence, the senior synonym for the Kazakh pika is O. opaca Argyropulo, 1930. Thus, we propose to recognize three separate species in the O. pallasii species group: O. pallasii (Mongolia and adjacent territories), O. opaca (eastern Kazakhstan) and O. argentata (Helan Shan Range, China).     

Evolutionary morphology of the Tenrecoidea (Mammalia) hindlimb skeleton

The tenrecs of Central Africa and Madagascar provide an excellent model for exploring adaptive radiation and functional aspects of mammalian hindlimb form. The pelvic girdle, femur, and crus of 13 tenrecoid species, and four species from the families Solenodontidae, Macroscelididae, and Erinaceidae, were examined and measured. Results from qualitative and quantitative analyses demonstrate remarkable diversity in several aspects of knee and hip joint skeletal form that are supportive of function‐based hypotheses, and consistent with studies on nontenrecoid eutherian postcranial adaptation. Locomotor specialists within Tenrecoidea exhibit suites of characteristics that are widespread among eutherians with similar locomotor behaviors. Furthermore, several characters that are constrained at the subfamily level were identified. Such characters are more indicative of postural behavior than locomotor behavior. J. Morphol., 2009. © 2008 Wiley‐Liss, Inc.     

Evolutionary trends in Hoplerythrinus unitaeniatus (Agassiz 1829) (Characiformes, Erythrinidae)

Hoplerythrinus unitaeniatus is a species of economic importance, widely distributed in the Brazilian territory. Many studies have reported the karyotypic variation that occurs in the species, which forms two groups according to the diploid number, one with 2n = 52 and the other with 2n = 48. In this work, three populations of H. unitaeniatus from different hydrographic basins were cytogenetically analyzed. The diploid number found was 2n = 48 for all populations, however, differences in karyotypic formula and fundamental number and heterochromatin distribution were also observed. Ag-NORs were multiple, GC-rich and when applied FISH with 18S rDNA bright signals in six chromosomes were observed. Data from this study confirm the occurrence of interpopulation variability in H. unitaeniatus. The basal number and the possible divergence among the different cytotypes analyzed for this species to date, as well as the forward the proposal of biogeographic and tectonic history for the continental margin are discussed.     

Characteristics of paravertebral muscles – fibre type distribution pattern in the pika, Ochotona rufescens (Mammalia: Lagomorpha)

The evolution of the locomotor apparatus in vertebrates is marked by major reorganizations in trunk's musculature. The hypothesized functions of mammalian back muscles in the literature are discussed under consideration of the distribution and proportion of oxidative, type‐I‐fibres, oxidative‐glycolytic, type‐IIa‐fibres and glycolytic, type‐IIb‐fibres in paravertebral muscles of a small mammal. The fibre type distribution was examined from a complete series of histological sections maintaining topographical relationships between the muscles as well as within the muscle, in order to establish the overall distribution pattern. The deep and short muscles showed the highest percentage of oxidative fibres. The larger, superficial paravertebral muscles contained the highest percentage of glycolytic fibres. Two muscles were intermediate in their proportion of fibre types. All epaxial muscles together can be interpreted as an antigravity muscle–complex counteracting enduringly against the rebound tendency caused by gravitation, comparable with antigravity muscles in limbs. A gradient from deep to superficial, or a clear regionalization of oxidative muscle fibres in central deep regions around a large intramuscular tendon was found in the m. spinalis and the m. quadratus lumborum, respectively. Concepts of the function of human back muscles as those of A. Bergmark (1989: Acta Orthop. Scand. 230 , 1) or S.G.T. Gibbons & M.J. Comerford (2001: Orthop. Division Rev. March/April, 21) were exposed to be more general within mammals. Functional specializations of different muscles and muscle parts are discussed under the consideration of evolutionary reorganization of the paravertebral musculature in tetrapods. Along the cranio‐caudal axis, the percentage of oxidative fibres was decreased in caudal direction within the same muscles, whereas the proportion of glycolytic fibres was increased. Therefore, classifications of muscles as ‘glycolytic’ or ‘oxidative’ based on biopsies or analyses of single cross‐sections may result in wrong interpretations. Changes in the proportions of the fibre type distribution pattern were mostly due to oxidative and glycolytic fibre types, whereas the percentage of oxidative‐glycolytic fibres had only minor influence. A significant positive correlation between the cross‐sectional area of the single fibre and its percentage in the area investigated were observed for oxidative fibres, whereby the size was positive correlated to the proportion of the oxidative fibres.