Functional morphology and anatomy of cervical vertebrae in Nacholapithecus kerioi, a middle Miocene hominoid from Kenya

This paper describes the morphology of cervical vertebrae in Nacholapithecus kerioi, a middle Miocene primate species excavated from Nachola, Kenya in 1999-2002. The cervical vertebrae in Nacholapithecus are larger than those of Papio cynocephalus. They are more robust relative to more caudal vertebral bones. Since Nacholapithecus had large forelimbs, it is assumed that strong cervical vertebrae would have been required to resist muscle reaction forces during locomotion. On the other hand, the vertebral foramen of the lower cervical vertebrae in Nacholapithecus is almost the same size as or smaller than that of P. cynocephalus. Atlas specimens of Nacholapithecus resemble those of extant great apes with regard to the superior articular facet, and they have an anterior tubercle trait intermediate between that of extant apes and other primate species. Nacholapithecus has a relatively short and thick dens on the axis, similar to those of extant great apes and the axis body shape is intermediate between that of extant apes and other primates. Moreover, an intermediate trait between extant great apes and other primate species has been indicated with regard to the angle between the prezygapophyseal articular facets of the axis in Nacholapithecus. Although the atlas of Nacholapithecus is inferred as having a primitive morphology (i.e., possessing a lateral bridge), the shape of the atlas and axis leads to speculation that locomotion or posture in Nacholapithecus involved more orthograde behavior similar to that of extant apes, and, in so far as cervical vertebral morphology is concerned, it is thought that Nacholapithecus was incipiently specialized toward the characteristics of extant hominoids.     

New evidence for canine dietary function in Afropithecus turkanensis

Despite considerable post-cranial and cranial morphological overlap with Proconsul, Afropithecus turkanensis is distinguished from that taxon by a suite of anterior dental and gnathic characters shared in common with extant pitheciin monkeys (i.e. low crowned, robust and laterally splayed canines, procumbent incisors, prognathic premaxilla, powerful temporalis muscles, reduced or absent maxillary sinuses, and deep mandibular corpora). Pitheciins are unique among living anthropoids because their canines serve a habitual dietary function and are not strictly influenced by inter-male competition. Given the functional association between pitheciin canine morphological specializations and sclerocarp foraging, a feeding strategy where the hard pericarps of unripe fruit are mechanically deformed by the canines, it has been suggested that Afropithecus may also have used its canines in a dietary context. This is confirmed by quantitative morphometric analyses of Afropithecus canine curvature and basal dimensions demonstrating that Afropithecus and extant pitheciins (Chiropotes, Cacajao) are distinguished from all other anthropoids by pronounced and evenly distributed mesial canine crown contours as well as greater resistance to canine bending in both the mesiodistal and labiolingual axes. In addition, Afropithecus, Chiropotes and Cacajao are also shown to have significantly longer and more curved premaxillae with greater incisor procumbency that effectively isolates the incisor and canine functional complexes. These morphological similarities are a result of convergence and not a shared derived ancestry. Despite their considerable morphological overlap, it is unlikely that Afropithecus and extant pitheciin diets are identical given significant dissimilarities in their post-canine morphology, maximum angular gape and body size. Nevertheless, Afropithecus canine dietary function is unique among hominoids and may have been a key component for the expansion of hominoids into Eurasia at the end of the early Miocene.     

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.     

Early Miocene catarrhine dietary behaviour: the influence of the Red Queen Effect on incisor shape and curvature

The early Miocene catarrhine fossil record of East Africa represents a diverse and extensive adaptive radiation. It is well accepted that these taxa encompass a dietary range similar to extant hominoids, in addition to some potentially novel dietary behaviour. There have been numerous attempts to infer diet for these taxa from patterns of dental allometry and incisor and molar microwear, however, morphometric analyses until now have been restricted to the post-canine dentition. It has already been demonstrated that given the key functional role of the incisors in pre-processing food items prior to mastication, there is a positive correlation between diet and incisal curvature (Deane, A.S., Kremer, E.P., Begun, D.R., 2005. A new approach to quantifying anatomical curvatures using High Resolution Polynomial Curve Fitting (HR-PCF). Am. J. Phys. Anthropol. 128(3), 630-638.; Deane, A.S., 2007. Inferring dietary behaviour for Miocene hominoids: A high-resolution morphometric approach to incisal crown curvature. Ph.D. Dissertation. The University of Toronto.). This study seeks to re-examine existing dietary hypotheses for large-bodied early Miocene fossil catarrhines by contrasting the incisal curvature for these taxa with comparative models derived from prior studies of the correlation between extant hominoid incisor curvature and feeding behaviour. Incisor curvature was quantified for 78 fossil incisors representing seven genera, and the results confirm that early Miocene fossil catarrhines represent a dietary continuum ranging from more folivorous (i.e., Rangwapithecus) to more frugivorous (i.e., Proconsul) diets, as well as novel dietary behaviours that are potentially similar to extant ceboids (i.e., Afropithecus). Additionally, early Miocene fossil catarrhine incisors are less curved than extant hominoid incisors, indicating a general pattern of increasing mesio-distal and labial curvature through time. This pattern of morphological shifting is consistent with the Red Queen Effect (Van Valen, L., 1973. A new evolutionary law. Evol. Theory 1, 1-30), which predicts that taxa that are removed from one another by geological time, although potentially having similar diets, may exhibit differing degrees of a similar dietary adaptation (i.e., differing degrees of incisal curvature).     

Milk molars or extra premolars in Mesotheriinae (Mesotheriidae,Notoungulata): New insights into an old controversy

A recently recovered specimen of Mesotheriinae (Mesotheriidae, Notoungulata) from the late Miocene-early Pliocene of La Rioja Province (Argentina), CRILAR Pv 433, corresponds to an individual with three upper premolars, which appears to be an “anomaly” among mesotheriines. The detailed study of this specimen, however, brings up an old controversy on the interpretation of different mesotheriine specimens with three upper or two lower premolars. After being described as different taxa, these were later considered to be juvenile representatives of other known species. The three upper or two lower teeth were interpreted as the milk molars DP2–4 and dp3–4, respectively, which would be replaced in adult life by two upper (P3–4) and one lower (p4) permanent premolars. The new material leads us to set up a different interpretation. In our opinion, all these specimens actually preserve the permanent dentition, corresponding to different ontogenetic stages of more or less young individuals. This consideration implies the necessity of a deep systematic revision of the whole subfamily, keeping in mind this new point of view and the ontogenetic variation within a species. Therefore, the presence of P2/p3 is not enough to define a different taxon at this moment. Pending this taxonomic revision, the dental morphology of CRILAR Pv 433 resembles both Typotheriopsis (e.g., upper premolars with one labial sulcus) and Pseudotypotherium (e.g., P4 with lingual groove, wide median lobe of M3) as these two late Miocene genera are currently characterized. Furthermore, P2/p3 could be expelled soon in the lifetime of individuals, and even the presence of P2/p3 could be a variable character within the same taxon; if so, this might reflect an evolutionary trend to the loss of a dental element within mesotheriines, but the revision of a large sample is necessary to support or reject these hypotheses.     

Cladistic relationships of extant and fossil hominoids

There is general agreement that the hominoid primates form a monophyletic group, that the extant great apes and humans form a second clade within that group with the gibbons as the sister group, and that the African apes and humans form a third clade. Although it has recently been proposed that humans and orang utans are sister taxa and also that the great apes form a clade to the exclusion of humans, our analysis, particularly of the molecular evidence, supports the existence of an African ape and human clade. The major problem in hominoid phylogeny at present is the relationships of the species within this clade: morphological data generally support the existence of an African ape clade which is the sister group to humans; some molecular data also support this conclusion, but most molecular evidence indicates the existence of a chimpanzee/human clade. We have cladistically re-analysed the DNA and protein sequence data for which apomorphic character states can be assessed. It is clear that there is a high degree of homoplasy whichever branching pattern is produced, with some characters supporting the existence of a chimpanzee/human clade and others supporting an African ape clade. When the cladistic analyses of morphological and molecular data are combined we believe that the most parsimonious interpretation of the data is that the African apes form a clade which is the sister taxon of the human (i.e., Australopithecus, Homo and Paranthropus) clade.This paper is not intended as a survey of all hominoid fossils but as a study of branching points in hominoid evolution and fossils are included which are relevant to this branching pattern. The analysis of fossil taxa in this study leads us to conclude that Proconsul is the sister taxon to the later Hominoidea. A number of middle Miocene forms such as Dryopithecus, Kenyapithecus, Heliopithecus and Afropithecus are shown to share derived characters with great apes and humans and provide evidence for the divergence of that clade from the gibbon lineage prior to 18 Ma. The position that Sivapithecus represents the sister group of the orang utan clade is supported here and shows that the orang utan lineage had diverged from the African ape and human lineage prior to 11·5 Ma. There is unfortunately no definitive fossil cvidence on branching sequences within the African ape and human clade, although a new specimen from Samburu, Kenya may be related to the gorilla.     

First giraffid skulls (Bohlinia attica) from the late Miocene Maragheh fauna,Northwest Iran

An almost complete skull and a second partial skull of Bohlinia attica (Artiodactyla: Giraffidae) from the late Miocene of Maragheh in northwestern Iran is described along with a complete upper dentition from Samos in Greece. These specimens enrich what is known of this species. The skull has massive bent ossicones. The braincase is horizontal to the face in lateral view. The upper premolars have strongly curved styles. The metapodials of this species are elongate with a deep plantar trough. The braincase is short and the occipital does not extend caudally. The type material of Bohlinia is from Pikermi but this taxon is also known from localities in F.Y.R.O. Macedonia, Greece, Turkey, Bulgaria, Iraq, and Iran. The new specimens are similar to others except that the ossicone terminates in a knob and the palatine choanae are positioned caudally. This taxon along with Honanotherium, which is most similar but with shorter metapodials and simpler premolars, can be placed in the subfamily Bohlininae.     

First complete skull of an octodontoid (Rodentia, Caviomorpha) from the Early Miocene of South America and its bearing in the early evolution of Octodontoidea

Octodontoidea is the most diverse group of caviomorph rodents. The systematics of most of the fossil representatives has been essentially based upon dental characters. Described here is an almost complete skull with dentition assigned to Prospaniomys Ameghino based upon its dental morphology. The specimen comes from the Sarmiento Formation at Pampa de Gan Gan (central Patagonia, Argentina), assigned to the Colhuehuapian SALMA (early Miocene). The most remarkable features are in the posterior portion of the skull, some of them shared with the modern octodontids and interpreted as specialized by previous authors, which contrast with the generalized dental morphology. These combined features were not previously known in other octodontoids. The comparisons with other fossil and extant members of the superfamily suggest that the characters traditionally used to associate Prospaniomys with the echimyids are very probably plesiomorphies. Prospaniomys would represent an early diverging lineage more closely related to modern octodontids than to echimyids, in which cranial structures evolved more rapidly than dental and mandibular ones.     

Preliminary analysis of<Emphasis Type="Italic"> Nacholapithecus</Emphasis> scapula and clavicle from Nachola,Kenya

The Miocene ape Nacholapithecus is known from rather complete skeletons; some of them preserve the shoulder joint, identified by three scapulae and one clavicle. Comparisons made with other Miocene and living apes (Proconsul, Equatorius, Ugandapithecus) suggest that the mobility of the scapulohumeral joint was important, and scapular features such as the morphology and position of the spine and the morphology of the acromion and axillary border resemble those of climbing arboreal primates except for chimpanzees, gorillas, or orang-utans. From the size of the scapula (male Nasalis size), it is clear that the animal is smaller than an adult chimpanzee, but the clavicle is almost as relatively long as those of chimpanzees. Some features closer to colobine morphology reinforce the hypothesis that Nacholapithecus was probably a good climber and was definitely adapted for an arboreal life.     

Some observations on enamel thickness and enamel prism packing in the miocene hominoid Otavipithecus namibiensis

Otavipithecus namibiensis is currently the sole representative of a Miocene hominoid radiation in subequatorial Africa. Several nondestructive techniques, such as computed tomography (CT) and confocal microscopy (CFM), can provide useful information about dental characteristics in this southern African Miocene hominoid. Our studies suggest that the molars of Otavipithecus are characterized by (1) thin enamel and (2) a predominance of pattern 1 enamel prism. Together, these findings provide little support for the recent suggestion of an Afropithecini clade consisting of Otavipithecus, Heliopithecus, and Afropithecus. Instead, they lend some (though not conclusive) support to the suggestion of an Otavipithecus/African ape clade distinct from Afropithecus. © 1995 Wiley-Liss, Inc.     

Late Miocene Microstonyx remains (Suidae, Mammalia) from Northern China

Several large suid cranial remains attributed to Microstonyx major are part of a new Hipparion Fauna collection from the Hezheng area, Northern China. The new material confirms the presence of Microstonyx in the late Miocene of the area. The Chinese form belongs to a small-sized eastern population with reduced premolar row and clear sexual bimodality. Statistical comparison shows that Microstonyx major was a polymorphic species and reinforces recognition of Hippopotamodon as a separate genus, defined by relatively stout premolars resulting from a different underlying pattern of allometric growth. The presence of Microstonyx in North China and the distinct suid assemblage that lived there suggest biogeographic connections between Northern China and Western Eurasia in contrast to isolation from Southern China and the Indian subcontinent. The suid fauna of the late Miocene of Northern China seems to have been restricted to the later, more humid phase represented by the Red Clay faunas.     

A new Late Jurassic species of the rare synechodontiform shark, <Emphasis Type="Italic">Welcommia</Emphasis> (Chondrichthyes,Neoselachii)

A new Jurassic species of the very rare and incompletely known synechodontiform shark, Welcommia, is described. The new species, Welcommia cappettai, is represented only by a single tooth, precluding reconstruction of its dentition in detail. Nevertheless, this specimen provides sufficient information and characteristics to establish its taxonomic status. Welcommia cappettai n. sp. occurs in the middle Oxfordian (Upper Jurassic) of south-western Germany. This is the first unambiguous record and named species of Welcommia from the Late Jurassic, substantially reducing the rather large gap in the fossil record of this synechodontiform taxon. So far, two Welcommia species from the Lower Jurassic of Belgium and the Lower Cretaceous of southern France have been described. An additional, still unnamed species seemingly occurs in the Oxfordian of southern France. The new species has plesiomorphic and apomorphic characteristics and, probably, an intermediate dental pattern that tentatively enables reconstruction of evolutionary trends in the dentition of this shark from small and compact teeth with broad, almost triangular cusps, to mesio-distally lengthened teeth with elongated mesial heels resulting in an extremely extended mesial cutting edge in addition to more delicate cusp and cusplets in advanced forms. These differences might be related to improved feeding mechanisms. It is hypothesized that Welcommia was predominantly a component of the Mediterranean faunal province. The disappearance of Welcommia in the Early Cretaceous remains ambiguous and might be related to competition by other sharks, for example hexanchiforms, or might represent a collecting bias and/or taxonomic misidentification of isolated teeth.     

Evolutionary and Biological Implications of Dental Mesial Drift in Rodents: The Case of the Ctenodactylidae (Rodentia,Mammalia)

Dental characters are importantly used for reconstructing the evolutionary history of mammals, because teeth represent the most abundant material available for the fossil species. However, the characteristics of dental renewal are presently poorly used, probably because dental formulae are frequently not properly established, whereas they could be of high interest for evolutionary and developmental issues. One of the oldest rodent families, the Ctenodactylidae, is intriguing in having longstanding disputed dental formulae. Here, we investigated 70 skulls among all extant ctenodactylid genera (Ctenodactylus, Felovia, Massoutiera and Pectinator) by using X-ray conventional and synchrotron microtomography in order to solve and discuss these dental issues. Our study clearly indicates that Massoutiera, Felovia and Ctenodactylus differ from Pectinator not only by a more derived dentition, but also by a more derived eruptive sequence. In addition to molars, their dentition only includes the fourth deciduous premolars, and no longer bears permanent premolars, conversely to Pectinator. Moreover, we found that these premolars are lost during adulthood, because of mesial drift of molars. Mesial drift is a striking mechanism involving migration of teeth allowed by both bone remodeling and dental resorption. This dental innovation is to date poorly known in rodents, since it is only the second report described. Interestingly, we noted that dental drift in rodents is always associated with high-crowned teeth favoring molar size enlargement. It can thus represent another adaptation to withstand high wear, inasmuch as these rodents inhabit desert environments where dust is abundant. A more accurate study of mesial drift in rodents would be very promising from evolutionary, biological and orthodontic points of view.     

Polymorphism of dental formula and segregation of its variants in a pedigree of Kerry Blue Terrier dogs

Polymorphism of the dental formula was analyzed in a complex pedigree of Kerry Blue Terrier. A lack of one or more lower premolars was observed in some dogs. Two different patterns of missing teeth were identified. One pattern consisted in agenesis of a second premolar, often in combination with agenesis of neighbor teeth, including the fourth premolar. In the second pattern, agenesis of a fourth premolar was expressed as an isolated abnormality. It was shown previously that the first pattern is inherited as a recessive trait with near complete penetrance. In this work, the major-gene control was demonstrated for the second pattern. This abnormality develops in 70–80% of mutant homozygotes and in no more than 20% of heterozygotes and wild-type homozygotes. It was shown that the two dentition abnormalities are controlled by different genes, which were designated LPA2 and LPA4 (Lower Premolar Agenesis).     

First partial face and upper dentition of the Middle Miocene hominoid Dryopithecus fontani from Abocador de Can Mata (Vallès‐Penedès Basin,Catalonia, NE Spain): Taxonomic and phylogenetic implications

A well‐preserved 11.8‐million‐years‐old lower face attributed to the seminal taxon Dryopithecus fontani (Primates, Hominidae) from the Catalan site ACM/C3‐Ae of the Hostalets de Pierola area (Vallès‐Penedès Basin, Catalonia, NE Spain) is described. The new data indicate that D. fontani is distinct at the genus level from Late Miocene European taxa previously attributed to Dryopithecus, which are here reassigned to Hispanopithecus. The new facial specimen also suggests that D. fontani and the Middle Miocene Pierolapithecus catalaunicus are not synonymous. Anatomical and morphometric analyses further indicate that the new specimen shows a combination of lower facial features—hitherto unknown in Miocene hominoids—that resembles the facial pattern of Gorilla, thus providing the first nondental evidence of gorilla‐like lower facial morphology in the fossil record. Considering the current evidence, the gorilla‐like facial pattern of D. fontani is inferred to be derived relative to previously known stem hominids, and might indicate that this taxon is either an early member of the Homininae or, alternatively, a stem hominid convergent with the lower facial pattern of Gorilla. The biogeographic implications of both alternatives are discussed. This new finding in the Hostalets de Pierola section reinforces the importance of this area for understanding the elusive question of the Middle Miocene origin and early radiation of great apes. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.