Canine carnassials: character displacement in the wolves, jackals and foxes of Israel

Previous research implies that competitive character displacement in felids and mustelids of Israel is expressed by canine size. Anatomy and observed killing behaviour of canids suggest that canines in this group are less adapted for the stylized role they play in felids and mustelids. Thus we hypothesized that character displacement, if it exists in canids, should not be manifested more clearly by canine size than by other traits. Five sympatric and at least partially syntopic canids occupy Israel, while in North Africa the largest (wolf) and smallest (Blanford's fox) are absent. Sexual size dimorphism in Israeli canids is generally less than in felids and mustelids (in which we analysed each sex as a separate ‘morphospecies’), so we used mixed-sex samples to represent each species. The three largest species (wolf, golden jackal and red fox) are also represented by Middle Palaeolithic samples in Israel, and all three had larger carnassial lengths then. Carnassial lengths, canine diameters and skull lengths are all remarkable evenly spaced among the five recent species in Israel. In Egypt, no trait manifests significant equality. Despite regional fluctuations in size, the carnassial length ratios of the three smaller species (foxes) are strikingly constant (1.18–1.21) throughout the region, while the ratios for the three larger species (wolf, jackal and red fox), sympatric only in Israel, are larger (1.33–1.34). Finally, mean carnassial length of jackals is constant across North Africa, while skull length and canine diameter both increase from Algeria through Egypt. All three traits are larger in Egypt than in Israel. We tentatively ascribe the equal ratios in Israel to competitive character displacement, though this hypothesis is speculative because of numerous lacunae in knowledge of diet, killing behaviour, available resources and extent of food limitation. Furthermore, humans have greatly affected range, density and ecology of wolves and jackals in the last century. Larger sizes in the Palaeolithic may well be manifestations of Bergmann's rule. The constancy of carnassial length in North African jackals, notwithstanding a longitudinal cline in CBLs of these populations, and the constant ratio between jackal and red fox carnassial length are both consistent with a hypothesis of character release in the absence of the wolf.     

Species co-existence and character divergence across carnivores

Co-occurring species might be morphologically similar because they are adapted to the same environment, or morphologically dissimilar to minimize competition. We use sister species comparisons to evaluate the relationship between morphological disparity and regional patterns of co-occurrence across carnivores. Up to 63% of the variation in range overlap can be explained by morphological divergence in dentition. Species that differ more in carnassial tooth length overlap more in their geographical range. Carnassials are the primary teeth associated with food processing, and hence difference in carnassial size may be a good indicator of difference in resource use. We suggest this pattern is consistent with competition in sympatry driving ecological character displacement, or competitive exclusion among ecologically similar species. Our study uses newly available data on global distributions, morphology and phylogeny, and is the first to demonstrate a close relationship between morphological disparity and co-occurrence at a regional scale encompassing multiple communities.     

The oldest Eurasian hominoid.

Engelswies is an early Miocene vertebrate locality in southern Germany with a rich assemblage of terrestrial mammals, invertebrates and fossil plants. It is dated to 16.5-17.0 Ma based on magnetostratigraphy, biostratigraphy and lithostratigraphy, and includes among the faunal remains a hominoid upper molar fragment, the oldest hominoid so far identified from Europe. The evidence from Engelswies suggests that hominoids arrived in Eurasia about 17 Ma, roughly contemporaneously with pliopithecoids and Deinotherium, and before the last marine transgression to isolate Eurasia from Africa. Thick enamel and low dentine penetrance may have been key adaptations that contributed to the success of hominoids of dentally modern aspect in western Eurasia and ultimately to their ability to spread to eastern Eurasia and Africa in the middle and late Miocene.     

The Leporid Datum: a late Miocene biotic marker

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Correlation of carnassial tooth size and body weight in recent carnivores (mammalia)

The correlation between the lower carnassial crown area and the body weight is examined for modern carnivores. It is very high, but the ursids and felids show a relationship different from the other carnivore families. Some exceptions are discussed. The possibility of predicting body weight for fossils is given.     

Miocene snakes from northeastern Kazakhstan: new data on the evolution of snake assemblages in Siberia

Abstract

The Neogene snake fauna from the central and eastern regions of Eurasia is still largely unknown. This paper reports on a unique snake fauna from the late middle Miocene of the Baikadam and Malyi Kalkaman 1 and 2 localities, northeastern Kazakhstan, which represents the best-documented Miocene snake assemblage in Central Asia. Previous studies admitted that snake fauna could be homogeneous over a large part of Eurasia during the Miocene, with the late middle to early late Miocene assemblages similar to snake assemblages that inhabited Europe in the late early and early middle Miocene. This assumption is partially supported by the presence of Texasophis bohemiacus and Coluber cf. hungaricus, as well as vipers of the ‘V. aspis’ complex. However, the presence of taxa which are (1) probably not related to European representatives (‘Colubrinae’ A and B), (2) probably never occurred in Central and Western Europe and (3) are closely related to species recently inhabiting southern Siberia (Elaphe aff. dione, Gloydius sp.) indicates that faunal dissimilarity was relatively high within Eurasia during the late middle Miocene. This assumption is in accordance with studies of small mammal assemblages which show a decreasing homogenity in the Eurasia in the course of the middle Miocene.     

Ontogeny of postcanine tooth form in the ferret,Mustela putorius (Carnivora: Mammalia), and the evolution of dental diversity within the mustelidae

This study describes dental development within the ferret, Mustela putorius, through study of the form of the carnassial teeth and the upper first molar at progressive growth stages. Primordial teeth were serially sectioned in sagittal and transverse planes and three-dimensional reconstructions of tooth primordia were generated using MacReco software. Regional growth of the crown and asynchronous maturation of the dental tissues were observed in each tooth. The upper carnassial blade develops early and the tooth increases in length rapidly. Lingual growth of the upper carnassial is less pronounced and the protocone and its surrounding region mature late. The lower carnassial blade develops early and the talonid is late to mature. Development of the upper first molar differs from carnassial development in the early emphasis upon transverse growth and reduced lengthwise expansion. The early development of the carnassial blades in the ferret is shared with other carnivores, and may reflect the functional significance of this feature. Later stages of tooth ontogeny differ among carnivoran taxa and the specialized morphology of ferret teeth results from an apparently truncated period of late tooth ontogeny. This suggests that carnivoran species may share a common path of early development that specifies the ontogeny of homologous tooth features and that in later stages developmental differences result in species-specific tooth forms. J. Morphol. 237:69–90, 1998. © 1998 Wiley-Liss, Inc.     

Convergence in morphological patterns and community organization between Old and New World rodent guilds

We studied morphological relationships within three guilds of gerbillid rodents in Israel. We found a nonrandom pattern of overdispersed means (community-wide character displacement) for upper incisor widths among the species in these three guilds. Upper tooth-row lengths, condylo-basal skull lengths, and tooth-row surfaces displayed similar patterns. We also studied seed-size selection by two well-studied gerbil species, which have previously been found to compete, in order to test whether specializing on husking seeds of different sizes as a mechanism of coexistence may underlie the morphological patterns. The seed-size selection experiments took place in two large aviaries with artificial lighting simulating full-moon nights, which is when predation risk is perceived as high. Seeds of different sizes (commercial seeds in one experiment and husked wheat particles in the other) mixed with sand were offered in trays. The larger Gerbillus pyramidum took significantly larger commercial seeds and marginally larger wheat particles than the smaller Gerbillus allenbyi. The patterns attest to ecomorphological convergence at the guild level; we previously demonstrated size structuring in several North American heteromyid rodent guilds, and we now report similar size structuring among Israeli gerbillid guilds. The occurrence of convergent community structure strongly indicates general rules governing ecological communities or guilds.     

Mio-pliocene faunal exchanges and african biogeography: the record of fossil bovids

The development of the Ethiopian biogeographic realm since the late Miocene is here explored with the presentation and review of fossil evidence from eastern Africa. Prostrepsiceros cf. vinayaki and an unknown species of possible caprin affinity are described from the hominid-bearing Asa Koma and Kuseralee Members (～5.7 and ～5.2 Ma) of the Middle Awash, Ethiopia. The Middle Awash Prostrepsiceros cf. vinayaki constitutes the first record of this taxon from Africa, previously known from the Siwaliks and Arabia. The possible caprin joins a number of isolated records of caprin or caprin-like taxa recorded, but poorly understood, from the late Neogene of Africa. The identification of these two taxa from the Middle Awash prompts an overdue review of fossil bovids from the sub-Saharan African record that demonstrate Eurasian affinities, including the reduncin Kobus porrecticornis, and species of Tragoportax. The fossil bovid record provides evidence for greater biological continuity between Africa and Eurasia in the late Miocene and earliest Pliocene than is found later in time. In contrast, the early Pliocene (after 5 Ma) saw the loss of any significant proportions of Eurasian-related taxa, and the continental dominance of African-endemic taxa and lineages, a pattern that continues today.     

Origines du peuplement mammalien en Afrique du Nord durant le miocène terminal

The late Miocene North African mammalianassemblage is considered here from three viewpoints: survivals, extinctions, and immigrations. The Eurasiatic affinities of the large mammals slightly prevail over the Ethiopian affinities. Amongst the North African large mammals, 4 to 8 taxa are Eurasiatic immigrants, while 4 to 6 are of Subsaharian origin. Contrarily, the micromammalian fauna is highly endemic, with only one species, a murid (Paraethomys miocaenicus), considered here as being related to an Asiatic form (Karnimata darwini). Our study of Eurasian and African Miocene faunas reveals that during the late Astaracian-early Turolian interval, the Saharo-Arabic belt permitted very little latitudinal faunal exchanges. However, during the middle and late Turolian such faunal exchanges became frequent. The micromammal record unequivocally indicates that a brief period of faunal exchange occurred between Africa and western Europe at the end of the Miocene, corresponding with the Messinian Salinity Crisis. The increased intercontinental faunal exchange between Africa and Eurasia during the late Miocene coincides with, and counterbalances the extinction of more than 10 taxa at the Mio/Pliocene boundary.     

Gradual evolution in the African hunting dog lineage Systematic implications

The hunting dog, Lycaon pictus, is one of the most representative species of the extant African fauna and one of the most efficient predators in the World. This species is actually a relict within Sub-Saharan Africa, but its lineage is well recorded in Eurasia during the Pliocene, and in Eurasia and Africa during the Early Pleistocene, while its record during the Middle Pleistocene is not well documented. Though postcranial skeletal remains dating from the Early Pleistocene show a tetradactyl forelimb, a characteristic feature of the extant lycaon, unique among canids, the upper and lower dentitions show gradual evolution from a primitive morphology in the Late Pliocene specimens to the highly specialized trenchant carnassials of the extant predatory species. We propose a new systematics for the lineage, grouping all the forms within the genus Lycaon. To cite this article: B. Martínez-Navarro, L. Rook, C. R. Palevol 2 (2003).     

Historical Biogeography and Body Form Evolution of Ground Squirrels (Sciuridae: Xerinae)

Xerinae is the most species-rich subfamily of the Sciuridae (Rodentia). This group of animals has a long complex evolutionary history, which witnessed severe environmental changes. In this paper, a comprehensive approach integrating information from fossil records, morphological, molecular and geographical data of extant species, and events of paleoclimate and paleogeography, were used to explore the evolutionary processes in the Xerinae. Xerinae probably originated in Eurasia around the early Oligocene, and dispersed to Africa via the Africa-Eurasia Land Bridge on two occasions during the Miocene, and subsequently evolved into the Protoxerini and African Xerini. The tribe Marmotini derived from a Eurasian ancestor and thrived in North America. Tamias re-occupied Eurasia in the early Miocene, while the distributions of Marmota and ‘Spermophilus’ genus-groups were restricted to North America at least until the late Miocene. Global cooling and the emergence of grass-dominated ecosystems from 15 Ma are likely to be the main causes for the radiation of Marmotini. The body form of Xerinae displays an allometric mode of evolution, with ground-living taxa, such as Marmota, Cynomys and Xerus notably enlarged, while Tamias has remained slim in body form. To cope with the global environmental changes, particularly the global cooling induced forest degradation and grassland expansion in the late Miocene, most Marmotini developed into true ground squirrels with short tails. The slim body adaptation in Tamias may be related to competition from tree squirrels, or their hoarding behavior, the latter helping them to cope with cold winter.     

First Finding of a Representative of Giant Mustelids of the Genus <Emphasis Type="Italic">Eomellivora</Emphasis> (Carnivora,Mustelidae) in Russia (Tuva,Upper Miocene)

In the late Miocene deposits in the Taralik–Cher locality (Tuva Republic), fossil remains of a giant mustelid of the genus Eomellivora Zdansky, 1924 have been found. This finding is the first reliable evidence of the Eomellivora inhabiting the modern Russia. Teeth of the Eomelliovra from Taralik–Cher are similar in size to those of E. wimani and E. piveteaui from the late Miocene in Eurasia. The morphology of teeth of the material from Tuva is most similar to that of E. ursogulo; therefore, it is considered a small form of Eomellivora ursogulo. The described finding expands the understanding of diversity, variability, and distribution of representatives of the genus Eomellivora in Asia during the late Miocene.     

The large carnivores from ‘Ubeidiya (early Pleistocene, Israel): biochronological and biogeographical implications

The archaeological site of ‘Ubeidiya provides a unique, rich faunal assemblage, as well as evidence for the presence of early hominins in the Levant. Paleontological analysis of the faunal community is pertinent to understanding the biochronology and the timing of hominin dispersal events from Africa to Eurasia. This study presents the large carnivores from the recent seasons of excavations (1989-1999) and previously published material. Based on biochronological correlations of the carnivore guild and paloemagnetism, the age of the site is 1.5-1.2 Ma. This date is 100-200,000 years earlier than previous estimates. Of specific importance is the presence of the African origin saber tooth Megantereon cf. M. whitei and the Eurasian origin canids Canis moschbachensis and Lycaon lycoanoides.     

Out of Africa: biogeographic history of the open‐habitat chats (Aves,Muscicapidae: Saxicolinae) across arid areas of the old world

Arid and semi‐arid areas constitute a prominent feature of the earth today, especially in Asia and Africa. Their formation started in the middle Miocene with increased stepwise aridification since the Pliocene. This aridification had strong ecological and evolutionary consequences and not only led to fragmentation of moist‐adapted biota, but also fostered the evolution of arid‐adapted taxa from mesic ancestors and triggered speciation within arid areas. The open‐habitat chats, a clade within Saxicolinae (Aves, Muscicapidae), constitute one of the most significant arid‐adapted passerine groups of Africa and Eurasia. Here, we present a temporal and spatial framework for the diversification of open‐habitat chats, using probabilistic approaches for the reconstruction of their biogeographic history based on a time‐calibrated multilocus molecular phylogenetic hypothesis. The diversification of open‐habitat chats was initiated in the late Miocene at around 7.4 Ma, most likely in sub‐Saharan Africa. Southern Africa and the Horn of Africa acted as centres of diversification and biogeographic expansion. From the latter area, the Arabo‐Sindic region and subsequently further parts of Eurasia and North Africa were colonized. The colonization history out of sub‐Saharan Africa contrasts with that of several other songbird clades, where a biogeographic expansion from Eurasia or northern Africa to southern Africa was prevalent. Habitat fragmentation through forest expansions during intermittent wetter periods in Africa influenced diversification in several clades. However, phases of increased aridity, with hyperarid regions acting as drivers of vicariance, seem to have also been important in radiations of the Arabo‐Sindic region and the Horn of Africa during the Pleistocene. Different processes such as colonization of new areas followed by vicariance or speciation across ecotones might have played a role throughout the radiation of open‐habitat chats.     

Evolution of skull shape in carnivores 2. Additional modern carnivores

Fifteen functionally significant aspects of skull morphology were measured on skulls of 36 additional species of carnivores to complete a survey of skull shape in modern fissiped (land) carnivores that includes most of the living genera. The measurements were transformed to dimensionless variables based on the residuals from allometric equations, and were analysed singly and in a 10 variable principal components analysis. An initial study of 62 species of viverrids, canids, mustelids and felids had shown those families to be distinguished from each other by the functionally significant measurements. However, among the additional 36 species, some procyonids, ursids and mustelids display a range of diversity of skull morphology that overlaps that of the other families and diminishes the potential value of the measurements as taxonomic characters. Intraspecific variation is presented for 12 species, and is low enough to allow use of some features as species level diagnostic characters. The lack of correlation between diet and functionally significant aspects of skull morphology among omnivorous carnivores, and the absence of certain skull shapes among carnivores are discussed.     

The evolution of cursorial carnivores in the Tertiary: implications of elbow-joint morphology

The evolution of cursorial adaptations in Tertiary (65-1.65 Myr ago) carnivores has been a contentious issue. Most such studies have focused on the relationship between hind limb proportions and running speed. Here, we show morphometrically that in extant carnivores, the elbow joint has evolved in two distinct directions with mutually exclusive implications for locomotor ability and prey procurement. Some carnivores retain supinatory ability, allowing them to manipulate prey and other items with the forepaws. Such carnivores can become very large. Other carnivores lose the ability to supinate and become cursors. This allows for only moderate size increase. Modern carnivores above ca. 20 kg body mass are committed to one or other of these strategies. This threshold coincides with a postulated threshold in carnivore physiology. The biaxial pattern mostly follows phylogenetic lines, but a strong selective regime can override this signal, as shown by the extant cheetah. Oligocene (33.7-23.8 Myr ago) and early-middle Miocene (23.8-11.2 Myr ago) carnivores follow the same pattern, though in the Miocene the pattern is shifted towards larger body mass, which may be owing to the extraordinary richness of browsing ungulates at this time.     

The generalized carnivore jaw

A model is presented of the jaw mechanism that relies on the geometrical similarities among mammalian carnivores with carnassial teeth. These similarities, together with estimates of the location of the resultant force of the jaw muscles, allow the model to predict that the mechanical advantage of the jaw lever system is the same in all carnivores with carnassials and, therefore, that the magnitude of the bite force is mainly determined by the absolute amount of jaw musculature.     

Ecological and reproductive character displacement on an environmental gradient

Character displacement, in which coevolution of similar species alters their phenotypes, can be difficult to identify on the basis of observational data alone. In two-species systems, the most commonly identified (i.e., classic) resulting pattern is greater phenotypic difference between species in sympatry than allopatry. We show that restricting studies to this pattern may exclude many instances of character displacement, particularly in the presence of spatial environmental gradients. We present four spatial models of character displacement in quantitative traits affecting competition and hybridization between the species. Our models highlight the connections between range limits and character displacement in continuous space. We conclude that the classic pattern is less likely to occur for a trait affecting resource acquisition than for a trait affecting mate choice. We also show that interspecific hybridization (when hybrids are inviable), even in very small amounts, has marked effects on the shape and stability of borders between species and the nature of character displacement. A survey of the empirical literature shows that character displacement studies often lack analysis of spatial phenotype and abundance data. We recommend more careful spatial sampling in character displacement studies, and we illustrate how comparison of clines in mean phenotype in sympatry and allopatry can be used to suggest the action of character displacement.     

Dispersal of African mammals in Eurasia during the Cenozoic: Ways and whys

Several groups of mammals originated in Africa and then immigrated to Eurasia during some intervals of the Cenozoic, thus greatly contributing to the mammalian biodiversity in Eurasia. Nevertheless, the African components of Eurasian mammalian faunas have had variable success in their diversification and survival. The Afro-Arabian plate remained separated from Eurasia by the Tethyan seaway, which was definitely closed in the Burdigalian, some 20 myr ago. Before its closure, the marine barrier between the Afro-Arabian and Eurasian plates did not totally prevent mammalian exchanges between these landmasses, as documented by the arrival of rodents and primates in Africa in the late Paleocene-early Eocene, the dispersal of embrithopods on both sides of the Tethyan seaway during the Eocene, and the immigration of elephantoids from Africa to Asia in late Oligocene. These events seem to be restricted to some groups of mammals, which apparently had abilities to use sweepstake dispersal routes. The massive mammalian dispersal from Africa to Eurasia started sometimes in the early Miocene, involving several groups of African mammals, in particular proboscideans, hyracoids, tubulidentates, and anthropoids. This contribution discusses the timing of these events under the light of recent discoveries of Africa-originated mammals in Eurasia. The impact of the evolving paleogeography of the area situated between the Afro-Arabian and Eurasian plates on the mammalian dispersal is reconsidered. The dispersal of land mammals from Africa to Eurasia is controlled not only by the paleogeographic changes (sea level changes, dispersal routes, terrestrial bridges, etc.), but also by climatic factors that modified the environments of terrestrial mammals, favoring or not the occurrence of dispersal routes and/or the enlargement or restriction of climatic belts and biogeographic provinces to which these mammals were adapted. These questions are discussed taking into account the present knowledge of the record of the Africa-originated mammals in Eurasia during the Cenozoic times.