Community-wide character displacement in Miocene hyaenas

Recent studies have found regularities in the pattern of distribution of dental parameters such as canine or carnassial length among sympatric carnivores. These regularities are taken to be indicative of community-wide character displacement. This study documents similar patterns in late Miocene and earliest Pliocene hyaenids from several localities in Eurasia and Africa. Statistical tests show ratios of lower carnassial total lengths and blade lengths between species to be suggestively equal among sympatric late Miocene hyaenas. Other measurements do not show this regular pattern. This finding mirrors that regarding modern canids in the Middle East, suggesting that a process leading to community-wide character displacement was in effect among these hyaenid taxa. Their response to this pattern suggests that they occupied a similar ecological role to modern canids. The causal basis for such a process is unknown but is suggested to lie in direct interspecific competition between carnivores rather than being a response to regularly spaced features of the environment. Character displacement, competition, Hyaenidae, Miocene, Eurasia, Africa.     

Comparative observations on some middle and upper miocenehyaenids,Genera: PercrocutaKretzoi, AllohyaenaKretzoi, AdcrocutaKretzoi (Mammalia,Carnivora, Hyaenidae)

This study deals with the «percrocutoid hyaenids,present in Eurasia and in Africa in faunas of late Middle Miocene (Astaracian) to terminal Miocene (Vallesian, Turolian) age. These hyaenid species are characterized by certain dental apomorphies (loss of M^2/2, specialization of the carnassials P⁴/M₁, tendency for the hypertrophy of the anterior premolars) which differentiate them from other hyaenids with which they are found associated. In recent literature, these hyaenids have been attributed to the genera PercrocutaKretzoi and AdcrocutaKretzoi, the latter of which is monospecific. In fact some should be removed from the genus Percrocuta and assigned to the genus AllohyaenaKretzoi, itself subdivided into the two subgenera, AllohyaenaKretzoi and DinocrocutaSchmidt-Kittler. The genus Adcrocuta is retained and the characters which distinguish this genus from HyaenictisGaudry are discussed.This study is based on revision of all, or nearly allsuch specimens, described and figured or in collections, of which certain among them have been hardly or poorly known. Particular attention has been given to their geologic age and their distribution in Neogene mammalian faunas (Mein's zones). Their apomorphies, considered in relation to their temporal distributions, permit an interpretation of the phylogenetic relationships of their several lineages which is consistant with the proposed systematic revision of the group.The radiation of «percrocutoid hyaenids would appear to have occurred in three phases. At the beginning of the Astaracian, after the dispersal of Anchitherium, the genus Percrocuta appeared with several species of relatively small size; in Eurasia this genus does not seem to have persisted after the middle Astaracian, although it did so in the sub-Himalayan Siwaliks. At the end of the Astaracian it was apparently succeeded by the genus Allohyaena, and resulting in gigantic forms (subgenus Dinocrocuta) coincident in Eurasia and in Africa with the extension of Hipparion. The third phase of the radiation is represented by the genus Adcrocuta, which unlike the others, was monospecific. This single species, A. eximia, occurs initially, and very rarely in the upper Vallesian; however, in the middle Turolian it is common and widespread throughout Eurasia where it, perhaps, limited the expansion of Dinocrocuta.     

A new species of Prosansanosmilus: implications for the systematic relationships of the family Barbourofelidae new rank (Carnivora, Mammalia)

A new barbourofelid species, Prosansanosmilus eggeri , is described from the Middle Miocene (MN 5) locality of Sandelzhausen, Germany. It differs from all other European barbourofelid species in being smaller and showing a more plesiomorphic morphology, especially in the relatively less developed sabretooth adaptations, low accessory cusps on the premolars, and the remnant of a very small talonid on the carnassial. The species is, however, stratigraphically later than the more apomorphic P. peregrinus, which is known from MN 4 of Germany and France. A phylogenetic analysis based on dental characters of early nimravids, barbourofelids and felids supports previous results on skull morphology of Barbourofelis that Barbourofelinae is not closely related to the Late Eocene and Oligocene Nimravinae. Instead, both subfamilies should be treated as separate families, with the Barbourofelidae closely related to the Felidae. The Barbourofelidae differ from the Felidae as well as from the Nimravidae s.s ., particularly in the unique morphology of their basicranium. They presumably originated in Africa; P. eggeri sp. nov. is interpreted as part of a Miocene immigration of African faunal elements into Europe that took place at the beginning of MN 5. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 43−61.     

Taxonomic affinities of the enigmatic Prionogale breviceps,early Miocene,Kenya

Prionogale breviceps is a tiny carnivorous mammal from the early Miocene of eastern Africa. Originally, specimens were interpreted as the adult morphology of the taxon. The dentition did not obviously align Prionogale with the carnivorous lineages present in Afro-Arabia during the early Miocene: Hyaenodonta and Carnivora. When Namasector was discovered in Namibia, the small taxa were placed together in Prionogalidae and aligned with Hyaenodonta. In this study, based on comparisons to hyaenodont specimens preserving deciduous dentition, the holotype of Prionogale is reinterpreted as preserving dP3 and dP4. Some of the lower dental specimens attributed to the taxon preserve dp4. The holotype of Namasector also preserves deciduous dental material. A phylogenetic analysis that includes deciduous dental characters for a broader sample of hyaenodonts resolved Prionogalidae as a clade. Understanding of the deciduous dentition of Prionogale allows future analyses to compare homologous morphology, and to explore the environmental factors that shaped carnivorous mammal evolution through the Miocene.     

Creodonts and carnivores from the Middle Miocene Muruyur Formation at Kipsaraman and Cheparawa,Baringo District,Kenya

The Middle Miocene Muruyur Formation (ca 14.5 Ma), Tugen Hills, Kenya, has yielded a huge creodont and a variety of carnivores ranging in size from mongoose-sized viverrids and herpestids to lion-sized amphicyonids. The fauna partly fills what used to be a major gap in our knowledge of Neogene African carnivores, spanning the period between the better known Early Miocene assemblages of western Kenya and eastern Uganda, and the Late Miocene and Plio–Pleistocene faunas of East Africa. Present in the deposits are Megistotherium, two species of Hecubides, one species of Agnotherium, Herpestes, Vishnuictis, and one or two undetermined felids.     

Deciduous dentition and eruption pattern in late Miocene Pachyrukhinae (Hegetotheriidae,Notoungulata) from La Pampa Province,Argentina

The study of juvenile remains of Paedotherium Burmeister from Cerro Azul Formation (La Pampa Province, Argentina; late Miocene) is presented. Upper and lower deciduous dentition (or permanent molars supposed to be associated with non-preserved deciduous teeth) are recognised. Several ontogenetic stages are distinguished among juveniles, according to the degree of wear and the replaced deciduous teeth. Besides, some morphological and metrical differences are observed along the crown height. Deciduous cheek teeth are high-crowned and placed covering the apex of the corresponding permanent tooth. The height of the crown and the degree of wear allow establishing the pattern of dental replacement of deciduous and permanent premolars in a posterior–anterior direction (DP/dp4–2 and P/p4–2), as well as the eruption of M/m3 before DP/dp4 is replaced. Some of the studied remains are recognised as young individuals of Tremacyllus Ameghino, but with complete permanent dentition, which leads to propose a different timing in the dental replacement with respect to Paedotherium; they also allow the establishment of an opposite premolar eruption pattern, from P/p2 to P/p4. This knowledge of the deciduous dentition of Paedotherium suggests the need of revising the morphological and metrical characters previously used for defining species within this taxon.     

Messelogale, eine neue Raubtiergattung aus dem Mitteleozän von Messel (Deutschland)

Another juvenile specimen of the small carnivoranMiacis?kessleri Springhorn 1982 from the lower Middle Eocene oilshales (MP 11) of Messel near Darmstadt (South Hesse, Germany) is described. The nearly complete skeleton displays characteristics in dentition, skull, and limbs showing additional anatomical details and suggesting a taxonomic revision. Especially the assumption that there exist two molars only in the lower and upper jaw, as well as the trenchant talonid of the lower carnassial contradict the allocation to the genusMiacis. The species consequently is assigned to a new genusMesselogale. The higher taxonomic status of miacids in relation to the system of Carnivora is discussed.Messelogale kessleri shows basicranial characters, which indicate a classification within the Caniformia. On the other hand the loss of M3 suggests a relationship to the Feliformia. The species has an almost completely reduced (cf. holotype) or moderately well-developed dP³-Parastyl (this specimen), but it is not as enlarged as in primitive Feliformia, nor as reduced as in some Caniformia. Moreover, a calcaneal fibular facet cannot be excluded. Both attributes are representing the primitive morphology of the species. That does not ally the species with the Feliformia, but makes it, according toFlynn &; Galiano (1982), a very primitive Caniformia.     

Carnivores from the early/middle Miocene locality of Antonios (Chalkidiki, Macedonia, Greece)

The mammal locality of Antonios is one of the few known early-middle Miocene ones of Greece with large mammals. It is situated in Chalkidiki Peninsula (Macedonia, Greece) and includes both small and large mammals. The study of the carnivores found in Antonios indicates the presence of the following taxa: Proputorius cf. P. sansaniensis, Protictitherium gaillardi, Protictitherium cf. P. crassum, Percrocuta sp., Percrocutidae indet. and Pseusaelurus romieviensis. The material of each taxon is described, compared with other material from various Eurasian localities and determined. The species P. cf. sansaniensis and P. romieviensis are traced for first time in Eastern Mediterranean, while Percrocuta for first time in Greece. The presence of the other carnivoran taxa in the Antonios fauna is also interesting as they are recognized in older stratigraphic levels enriching our knowledge about their stratigraphic distribution. The carnivores of Antonios cannot offer significant data for the dating of the fauna but its age is discussed using the data coming from the carnivores as well as those from older studies of the micromammals and suoids.     

Additional specimens of Hyracoidea (Mammalia) from the Early and Middle Miocene of Kenya

We describe unpublished material of the poorly known hyracoid, Brachyhyrax aequatorialis from the Early Miocene of Songhor and Koru, Kenya, on the basis of specimens stored in the Community Museums of Kenya, the National Museums of Kenya and the Natural History Museum, London. As a result, we added 17 specimens to the hypodigm of this species which was previously known from only seven specimens. In addition, we describe further material of another hyracoid, Afrohyrax championi, from the Early Miocene of Mfwangano and the Middle Miocene of Kipsaraman. The chronological distributions of the two hyracoids do not overlap. Brachyhyrax appears to have lived in forest environments, a suggestion supported by its brachyodont dentition and the associated molluscan fauna, whereas Afrohyrax seems to have lived in more open wooded habitats which agree with its slightly more hypsodont dentition and the cursorial postcranial skeleton as well as with the land snails that occur with it.     

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.     

Geographic Variation in Nonmetric Dental Traits of the Deciduous Molars of Pan and Gorilla

Physical anthropologists often use nonmetric dental traits to trace the movement of human populations, but similar analysis of the teeth of nonhuman primates or the deciduous teeth is rare. Because nonmetric dental characteristics are manifestations of genetic differences among groups, they vary among geographically distant members of the same species and subspecies. We use 28 nonmetric dental traits in the deciduous molars to compare genetically and geographically distinct groups of extant African apes (Gorilla and Pan). Previous researchers have studied these traits in the adult or juvenile teeth of great apes and humans, and we score our observations according to established standards for hominins. We observe marked differences in trait frequencies between Gorilla and Pan, Pan troglodytes and P. paniscus, and two P. troglodytes subspecies but we find no significant differences between geographically isolated groups within the subspecies. Trait frequencies differ from those found in previous studies that contained fewer individuals. We find that the deciduous molars show similar variation to adult premolars and molars within Pan and Gorilla. This suggests that the deciduous dentition of these and other apes may contain diagnostic traits that are not currently in use.     

On the Middle Miocene avifauna of Maboko Island,Kenya

The Middle Miocene sediments of Maboko Island (Lake Victoria) in western Kenya yielded numerous avian bones, which remained, however, little studied. The significance of this material is shown by the recent identification of an opisthocomiform bird. In the present study, further avian remains from Maboko Island are described. Most of the specimens belong to aquatic or semi-aquatic groups, of which some are closely related to taxa known from Early and Middle Miocene European avifaunas, that is, Nectornis cormorants (N. africanus nov. sp.) and Laricola-like Laromorphae. The fossil material also includes Ciconiidae (cf. Ciconia), Pelecanidae, Phoenicopteridae (Leakeyornis aethiopicus), Musophagidae, and a species of Ardeidae, which closely resembles the taxon Pikaihao from the Early Miocene of New Zealand. Some avian remains from Maboko Island belong to higher-level taxa unknown from the Middle Miocene of Europe. The occurrence of a giant Jacanidae (?Nupharanassa mabokoensis nov. sp.) is of particular interest, because these are globally absent in extant avifaunas and were previously only known from the Late Eocene/Early Oligocene of Egypt. Further unknown from contemporaneous European sites are small representatives of Jacanidae, Bucerotidae, and Alcedinidae, with the fossils of the latter two taxa being among the earliest published records of their respective groups. Several of the taxa that are common in contemporaneous European avifaunas have not been found in Maboko, and in spite of less pronounced climatic differences, Middle Miocene Afrotropical avifaunas already appear to have been distinct from contemporaneous European ones.     

Homologies of the anterior teeth in indriidae

Schwartz ('74) proposed revised homologies of the deciduous and permanent anterior teeth in living lemuriform primates of the family Indriidae. Gingerich ('77) described a juvenile specimen of Avahi and emphasized the importance of functional integrity in controlling the pattern of dental reduction in primates, neither of which supports Schwartz's interpretation. Schwartz ('78) recently reiterated his position without adequately discussing the Avahi evidence and the functional basis that probably controls dental reduction. Avahi has a deciduous dentition intermediate in morphology between that of Lemuridae and Indriidae, and similar to both. Thus the lower deciduous dental formula of Indriidae is probably 2.1.3, which is the typical and maximum deciduous complement known in living and fossil lemuriform primates. The formula of the lower permanent dentition in Indriidae is thus 2.0.2.3.     

Pollen analysis of coal-bearing Miocene sedimentary rocks from the Seyitömer Basin (Kütahya), Western Anatolia

The late Early-Middle Miocene sequences of the Seyitömer Basin (western Anatolia) were palynologically investigated. Fifty-five taxa belonging to seven gymnospermous and 48 angiospermous pollen genera were identified in the 19 productive samples. Two pollen zones were recognised based on the changing abundance of individual tree taxa. Zone 1 is characterized by predominance of Pinus and Cedrus. Zone 2 is characterized by predominance of deciduous Quercus and evergreen Quercus and a marked reduction in representation of Taxodiaceae. The differences in the pollen spectra between Zone 1 and Zone 2 may reflect the global Middle Miocene cooling. These results are largely comparable to pollen data derived from the neighbouring areas. The vegetation of the Seyitömer Basin was dominated by trees. This palynological analysis reveals the existence of a swamp-forest developed in a subtropical to warm-temperate humid climate.     

Sexual dimorphism in the canines and skulls of carnivores: effects of size, phylogency, and behavioural ecology

Sexual dimorphism in craniodental features is investigated in a sample of 45 carnivore species in relation to allometry, phylogeny, and behavioural ecology. Dimorphism is more pronounced in both upper and lower canine size and strength than in carnassial size, skull dimensions and biomechanical features, but all dimorphism indices covary. As with most morphological characters, differences in canine sexual dimorphism are significantly related to phylogeny, estimated from either taxonomic rankings or a limited matrix of molecular distances; in particular, mustelids, felids and procyonids are more dimorphic than other carnivore families. Thus, because of problems related to species dependence in comparative data, remaining analyses are based on phylogenetically transformed values using a spatial autoregressive method.
In contrast to other mammals, sexual dimorphism in carnivore canines is not correlated with differences in body weight, skull length or basicranial axis length. Nor is it correlated with categorical variables of activity pattern, habitat, or diet. In our Carnivore sample, canine dimorphism is related only to breeding system: uni-male, group-living (harem) species have significantly greater canine dimorphism than multi-male, multi-female groups and monogamous pair-bonding species. By contrast, dimorphism in carnassial size is related to dietary differences, specifically greater dimorphism in meat-eating species, and not breeding patterns. Dimorphism in estimates of jaw muscle size suggest functional demands from both diet and breeding type. It is concluded that, befitting patterns of heterodont dentition, sexual selection influences variation in canine dimorphism while feeding ecology is related to carnassial dimorphism.     

Un nouveau felide (Fissipeda,mammalia) du pleistocene moyen du Maroc: Lynx thomasi N. sp.

The site known as Thomas quarry III (Cassablanca, Morocco) yielded, together with a middle Pleistocene (Tensiftian) fauna, the lower jaw of a lynx, Lynx thomase n. sp., whose dentition is characterized by high and complex premolars, and a very long carnassial. This new species might be descended from post-villafranchian European lynxes.     

Dabbling ducks (Aves: Anatidae) from the Middle Miocene of Mongolia

A new genus and two new species of ducks (Aves: Anatidae) from the Middle Miocene Sharga locality are described. Mioquerquedula minutissima gen. et sp. nov. is a very small duck. Anas velox Milne-Edwards, 1868 from the Middle Miocene of France is transferred to the genus Mioquerquedula. Aix praeclara sp. nov. described here is the oldest record of the modern genus Aix. A revision of the previously described small duck Anas soporata Kurochkin, 1976 shows that only the specimens from the Sharga locality should be referred to this species. The status of other small ducks from the Neogene of Europe and North America is discussed. The diversity of herbivorous and diving ducks in the Sharga locality indicates that Miocene Shargyn Govi Lake was rich in food resources.     

Dentognathic remains of an Afropithecus individual from Kalodirr,Kenya

We describe here the well-preserved dentognathic remains of an Afropithecus individual from the early Miocene site of Kalodirr in northern Kenya. The specimen includes a nearly complete dentition in which most of the crowns are undamaged and unworn. The new information gleaned from this specimen adds to our knowledge of this genus in several ways. Afropithecus exhibits an atypical pattern of canine dimorphism, and is probably more easily sexed by the pattern of variation in its upper premolars. Both phenomena are likely related to the modification of its antemolar dentition for the purpose of sclerocarp harvesting. The new fossils clarify the role in this adaptation of the premolars, which appear specialized for initiating and propagating cracks in large food items bearing protective coats. Comparison with other Miocene apes suggests that the closest known relative of Afropithecus is Nacholapithecus, to which Equatorius is more distantly related. Morotopithecus shares some primitive traits with Afropithecus, but lacks the derived features shared by the latter and Nacholapithecus.     

Miocene spores and pollen from Pelitçik Basin,Turkey–environmental and climatic implications

The palynological investigation of the early Middle Miocene fluviolacustrine sedimentary rocks interfingering with volcanics of the Galatean Volcanic Province at Pelitçik Basin (Central Turkey) have yielded palynomorphs belonging to 51 spore and pollen taxa. The pollen record is dominated by Ulmus, Pinaceae, Quercus, Carpinus and Carya, and appears to reflect climatic conditions. Two pollen zones were established based on changing abundances of plant taxa. Zone 1 is characterized by dominance of Ulmus and Pinaceae. Zone 2 is differentiated and characterized by a dramatic increase in, and predominance of, Ulmus, Carya, deciduous Quercus, Carpinus, Salix and Pinaceae. Mixed mesophytic forests were widespread in the basin suggesting warm and temperate climate. The decreasing relative percentage of thermophilous taxa, with Engelhardia as the main component, and warm temperate taxa, such as Carya, at the upper part of Zone 2 generally indicates a climatic deterioration, probably related to the Middle Miocene cooling.     

A new duck from the Middle Miocene of Mongolia, with comments on Miocene evolution of ducks

A new relatively large duck, Chenoanas deserta gen. et sp. nov. from the Middle Miocene of the Sharga locality, which is morphologically intermediate between the extant genera Chenonetta and Tachyeres, is described. The diversity of Early and Middle Miocene ducks is discussed. It is noted that some Middle Miocene duck remains are incorrectly referred to the genus Mionetta. The distribution of morphological characters of the humerus in Neogene and extant ducks shows that the present day diversity of ducks apparently results from extinction of some taxa which were formed in the Oligocene-Early Miocene. The distribution of morphological characters in the evolution of diving ducks is evidence that not only the formation of different morphotypes but also so-called ??evolutionary maturation?? of taxa also explains the modern diversity.