Middle Miocene dispersals of apes

The earliest record of fossil apes outside Africa is in the latest early Miocene of Turkey and eastern Europe. There were at least 2, and perhaps 4, species of ape, which were found associated with subtropical mixed environments of forest and more open woodland. Postcranial morphology is similar to that of early Miocene primates and indicates mainly generalized arboreal quadrupedal behaviours similar to those of less specialized New World monkeys such as Cebus. Robust jaws and thick enamelled teeth indicate a hard fruit diet. The 2 best known species of fossil ape are known from the site of Pa?alar in Turkey. They have almost identical molar and jaw morphology. Molar morphology is also similar to that of specimens from Germany and Slovakia, but there are significant differences in the anterior teeth of the 2 Pa?alar species. The more common species, Griphopithecus alpani, shares mainly primitive characters with early and middle Miocene apes in Africa, and it is most similar phenetically to Equatorius africanus from Maboko Island and Kipsaramon. The second species is assigned to a new species of Kenyapithecus, an African genus from Fort Ternan in Kenya, on the basis of a number of shared derived characters of the anterior dentition, and it is considered likely that there is a phylogenetic link between them. The African sites all date from the middle Miocene, similar in age to the Turkish and European ones, and the earliest emigration of apes from Africa coincides with the closure of the Tethys Sea preceding the Langhian transgression. Environments indicated for the African sites are mixtures of seasonal woodlands with some forest vegetation. The postcrania of both African taxa again indicate generalized arboreal adaptation but lacking specialized arboreal function. This middle Miocene radiation of both African and non-African apes was preceded by a radiation of arboreal catarrhine primates in the early Miocene, among which were the earliest apes. The earliest Miocene apes in the genus Proconsul and Rangwapithecus were arboreal, and because of their association with the fruits of evergreen rain forest plants at Mfwangano Island, it would appear that they were forest adapted, i.e. were living in multi-storied evergreen forest. The same or similar species of the same genera from Rusinga Island, together with other genera such as Nyanzapithecus and the small ape Limnopithecus, were associated with plants and animals indicating seasonal woodland environments, probably with gallery forest forming corridors alongside rivers. While the stem ancestors of the Hominoidea were almost certainly forest adapted, the evidence of environments associated with apes in the later part of the early Miocene and the middle Miocene of East Africa indicates more seasonal woodlands, similar to those reconstructed for the middle Miocene of Pa?alar in Turkey. This environmental shift was probably a requisite for the successful emigration of apes out of Africa and made possible later movement between the continents for much of the middle Miocene, including possible re-entry of at least one ape lineage back into Africa.     

A reassessment of the taxonomic and phylogenetic affinities of the fossil catarrhines from fort Ternan,Kenya

This paper presents a reassessment of the taxonomic and phylogenetic affinities of the fossil catarrhine primates from the important middle Miocene site of Fort Ternan in Kenya. Although the sample of specimens is rather small, the material can be attributed to at least five different species, identified here asKenyapithecus wickeri, Proconsul sp., a large species of oreopithecid,Simiolus sp., and a small species of catarrhine of indeterminate status.Kenyapithecus wickeri probably represents a conservative sister-taxon of the extant large hominoids. It is more derived than“Sivapithecus” africanus from Maboko Island, from which it can be distinguished at the generic level. A small species of catarrhine from Fort Ternan can be attributed toSimiolus. It is probably a different species fromSimiolus enjiessi from the early Miocene of East Africa, but additional material is needed to confirm its taxonomic distinctiveness. The occurrence of at least five species of catarrhine primates at Fort Ternan confirms that species diversity levels were as high during the middle Miocene as they had been during the early Miocene. However, the overall taxonomic and ecological composition of the middle Miocene catarrhine community was quite different, evidently due to a significant change in the local ecological setting. Taxonomic differences between the catarrhine faunas at Fort Ternan and Maboko Island can probably be explained as a consequence of a chronological separation between the two sites, and, to a lesser degree, to paleoecological differences.     

An oreopithecid proximal humerus from the middle miocene of Maboko Island,Kenya

A proximal humerus, recently recovered from the middle Miocene of Maboko Island, Kenya, provides the earliest evidence of postcranial structure and adaptation of Oreopithecidae. Provisionally attributed toNyanzapithecus pickfordi (Harrison, 1986), the specimen manifests a globose head, subequally large tuberosities, and a board, shallow bicipital groove. Although readily distinguished from the fundamentally cercopithecoid proximal humeral morphology ofVictoriapithecus (Senut, 1986), the Maboko Island oreopithecid, shows none of the derived features that are characteristic of the proximal humeri of extant hominoids. It is inferred from proximal humeral anatomy that the Maboko Island oreopithecid was an active arboreal scansor with moderate mobility at the shoulder but lacking adaptations for circumduction of the arm. In combination with craniodental evidence, proximal humeral morphology indicates that Oreopithecidae was a clade of hominoids which originated before the last common ancestor of extant apes and went extinct, without issue, in the later Miocene.     

New species of bushbaby from the middle Miocene of Maboko Island, Kenya.

A mandible recovered from ca. 15 million year old deposits of Maboko Island, Kenya, represents the first bushbaby known from the middle Miocene. The specimen is from a new species of Komba, a genus previously known from early Miocene occurrences in western Kenya and northeastern Uganda. Komba is revised, with emended diagnoses proposed for the genus, type-species, and referred species. Komba sp. nov. is distinguished by its larger size and differences of molar cusp acuity, buccal cingulum expression, and mental foramen configuration. Contrary to previous opinion, species of Komba probably diverged prior to the last common ancestor of extant Galaginae, and it is unlikely that they represent early stages of living bushbaby species lineages. Although contemporary Progalago is widely regarded as a galagine, aspects of upper molar, lower premolar, and mandibular corpus morphology indicate that it is more closely related to lorisines. Unlike the greater success currently enjoyed by bushbabies, lorisines were more diverse and almost as abundant as galagines in the early Miocene of eastern Africa.     

Faunal correlation of Wadi Moghara, Egypt: implications for the age of Prohylobates tandyi

Wadi Moghara, Egypt, is an early Miocene fossil locality with a mammalian fauna that includes Prohylobates tandyi, one of the earliest known representatives of the Cercopithecoidea. Faunal correlations were conducted between Moghara, Gebel Zelten (Libya) and a series of East African fossil sites with established radiometric dates in order to estimate the age of the Moghara mammals, including P. tandyi. Results confirm hypotheses proposed in some previous studies that: (1) Moghara is about 18-17 Ma, approximately the same age as the Hiwegi fauna from Rusinga Island (Kenya); (2) Moghara is slightly older than Gebel Zelten (Libya) (17-15 Ma); and (3) the cercopithecoid tooth presently identified as Victoriapithecus sp. from Napak V (ca. 19 Ma) is currently the oldest known record of a fossil Old World monkey, followed by P. tandyi from Moghara (Egypt) (18-17 Ma) and Prohylobates sp. from Buluk (Kenya) (>17.2 Ma), P. simonsi from Gebel Zelten (Libya) (ca. 17-15 Ma), and V. macinnesi from Maboko (Kenya) (ca. 16-14.7 Ma).     

A New Rhinoceros, Victoriaceros kenyensis gen. et sp. nov., and Other Perissodactyla from the Middle Miocene of Maboko, Kenya

The middle Miocene site of Maboko (Lake Victoria, Kenya), dated to ca. 15 Ma, has yielded one of the best collection of rhinos in Africa. The most common taxon, Victoriaceros kenyensis n.gen., n.sp., is represented by an almost perfect skull (whose main features are the large nasal horn, an orbit located very anteriorly and with a prominent border, and very broad zygomatic arches) and numerous limb bones, probably belonging to only a few individuals. Characters of the teeth and skull support an assignment to the subfamily Elasmotheriinae, a group best known in the middle and upper Miocene, but whose monophyly is disputable, as some of their tooth characters could be adaptations to a grazing diet (in agreement with their distribution in the Maboko beds). In any case, Victoriaceros clearly differs from other East African middle Miocene rhinos, whose diversity is far greater than currently assumed. A few other specimens attest to the occurrence at Maboko of at least one other species, perhaps close to the brachypotheres; a single calcaneum is tentatively assigned to the Chalicotheriidae.     

First fossil record of an East Asian endemic genus Sladenia (Sladeniaceae) from its modern range: Implications for floristic evolution and conservation biology

Fossil records of endemic plants play an important role in recognizing the floristic history of East Asia and thereby facilitate the conservation of plant diversity in the region. However, the fossil record of many extant East Asian endemic genera remains poorly documented thus far. Here, we report an infructescence fossil of an East Asian endemic genus, Sladenia (Sladeniaceae), from the early Miocene of southeastern Yunnan, China. The fossil is characterized by: (i) dichasial cymes; and (ii) flask‐shaped ovary with dense subparallel ribs on the surface extending from the base to the distal end of the united style. It represents the first fossil record of Sladenia in Asia, showing that the genus was established in the region at least by the early Miocene. Given that a much older fossil record of Sladeniaceae has been reported from Africa and the sister group of Sladenia is distributed only in Africa, Sladenia is not likely of East Asian origin. The present endemic status of Sladenia was possibly achieved by regional extirpation in Africa and taking refuge in East Asia. This case thus supports the “Museum” rather than “Cradle” hypothesis for the genesis of high plant species in the flora of East Asia. A comparison of the present fossil with extant Sladenia infructescence shows morphological stasis from the early Miocene to present. Such evolutionary tardiness might have resulted in the reduced fitness of the genus, which further caused its current endangered situation.     

5. JIMMY THE CROW

Harrison, C. J. O. 1980. Fossil birds from Afrotropical Africa in the collection of the British Museum (Natural History). Ostrich 51:92-98.

Although it has a rich avifauna, few fossil birds are know from Africa south of the Sahara. In the present paper 22 species are identified from three Miocene and two Pleistocene localities, 20 being additions to the existing list. A new touraco Apopempsis africanus sp. nov. is described from the Lower Miocene of Songhor, Kenya; and from the same period on Rusinga Island, Kenya, a new stork Ciconia minor sp. nov., in addition to a small flamingo, a hawk and a francol-in. There is evidence of a bustard and a Ciconia stork from the Middle Miocene of Maboko Island, Kenya. In the Pleistocene nine species are listed for Olduvai, Tanzania, and seven for Broken Hill, Zambia. There is some evidence of a shared Afro-tropicai/Palaearctic fuana in the Miocene, and of a larger inland lake bird fauna in the Early Pleistocene of Olduvai, but the Broken Hill material is purely Afrotropical.     

Morphology of the hallucial phalanges in extant anthropoids and fossil hominoids

Pedal phalanges of living anthropoids and several Miocene fossil hominoid taxa were studied to reveal functional adaptations of living anthropoid feet and to infer positional behavior of fossil hominoids. Among the examined living anthropoids, Pan has a very developed (long and robust) hallux. Proconsul and Nacholapithecus, a large hominoid from Nachola, northern Kenya, display a moderately long hallux like Alouatta and Cebus, suggesting the well-developed capability of a hallux-assisted power grip. Allometric analyses revealed that the Miocene hominoids examined (mainly from East Africa) as a whole displayed a different scaling pattern about the width of the proximal articular surface of the hallucial terminal phalanx from that of living anthropoids. Larger-sized hominoids display a wider articular surface than comparable-sized living anthropoids while smaller-sized fossil hominoids do the reverse. Such a difference was less marked for the height of the articular surface. These results may suggest that positional adaptations of Miocene hominoids are not merely resultants of a common body size function that is observed in living anthropods. The wide articular surface of fossil hominoid hallucial terminal phalanges suggests an adaptation for vertical climbing and clinging, in which the hallux is kept perpendicularly to the long axis of the vertical support.     

3. NOTES ON OSPREY,EGYPTIAN COOSE AND SPECKLED PICEON

Rich, P. V. &; Walker, C. A. 1983. A new genus of Miocene flamingo from East Africa. Ostrich 54:95-104.

Study of fossil flamingo (Phoenicopteridae) remains from the Miocene of East Africa suggests that a new genus, Leakeyornis, should be recognized to include Phoenicopterus aethiopicus Harrison &; Walker 1976. Osteologically this new genus is a mosaic between the three genera of living flamingoes, clearly distinct from all fossil forms, and possibly most closely related to Phoeniconaias, the Lesser Flamingo of East Africa.     

The fossil vertebrate locality Kossom Bougoudi,Djurab desert,Chad: A window in the distribution of the carnivoran faunas at the Mio–Pliocene boundary in Africa

The fossil vertebrate bearing locality Kossom Bougoudi (KB) is situated in the Djurab desert (Chad, Africa), 600 km north-east of N’djamena. It has yielded about 1250 specimens with many mammalian remains, principally artiodactyls. Its geologic age has been estimated to be about 5 Ma by biochronologic estimation and about 5.3 Ma by radiometric studies on cosmogenic nuclides of beryllium (authigenic ¹⁰B/⁹B). The carnivoran fauna contains few specimens which belong to five different families. All the taxa were unknown in central Africa. A large lutrine is close to Sivaonyx but different from known species of the genus. Another large lutrine is similar by its size to a species described from the Middle Pliocene of Uganda. An edentulous mandible of a small machairodont cat resembles a small species of Dinofelis, while a distal humerus indicates the presence of a larger member of the same genus. A hunting hyaenid is also much like the European species. An unidentified canid reaches the size of the recent Canis aureus and an isolated calcaneum matches that of the large extant viverrid. This small fauna allows a first look at the guild of the carnivorans at the Latest Miocene–Pliocene boundary in Central Africa and is a milestone between North African, East African and South African carnivore faunas.     

First fossil record of the catfish genus Sorubim (Siluriformes,Pimelodidae), from the Late Miocene of Paraná, Northeastern Argentina

The aim of the present contribution is to describe a partial fossil skull belonging to the catfish genus Sorubim. The specimen originates from the Late Miocene Ituzaingó Formation, at Paraná locality, Entre Ríos Province, Argentina. The material described here (MACN Pv-14224) was determined just to genus because the living species of Sorubim are morphologically similar. Presence of Sorubim is in concordance with the hypothesis indicating that most extant pimelodid genera were already present by the Late Miocene. Present finding constitutes the first fossil record for the genus.     

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.     

Long bones of the primate upper limb: Monomorphic or dimorphic?

An important debate has been taking place during the last few years concerningAustralopithecus afarensis: can the Hadar sample be ascribed to one highly dimorphic species or should it be separated into two distinct taxa? A similar problem occurs with the Middle Miocene cercopithecoids from East Africa: does this material belong to one dimorphic group or can we recognize two different taxa? The study of the long bones of the upper limb of many extant primates suggests that the extremities in different taxa are very distinctive but that within taxa the joints are weakly or are not morphologically dimorphic although they can be markedly size dimorphic. The main shape and size differences which can be ascribed to sexual dimorphism occur in the shafts of the long bones. Examinations have been made inHomo, Pan, Gorilla, Pongo, Hylobates, Alouatta, Cebus, Saimiri, Ateles, Nasalis, Presbytis and some Cercopithecinae. It appears, then, that the extremities of the bones are shape monomorphic. If the same relationships occurred in the fossil record, then the differences observed in the hominid fossil elbow joints at Hadar suggest that at least two different taxa are represented in the collection. In addition, among the cercopithecoid material assigned toVictoriapithecus from Maboko and Nyakach in East Africa, we recognize two distinct elbow morphologies indicating that two different taxa occur in the localities.     

Micropithecus clarki, a small ape from the Miocene of Uganda.

Micropithecus clarki, from Miocene sediments of Napak, Uganda, is the smallest known hominoid primate, living or fossil. In facial morphology it is very similar to extant gibbons. Dentally, it is most similar to the small apes from the Miocene of Kenya, Dendropithecus and Limnopithecus. All of the apes from the early Miocene of East Africa seem to represent a single phyletic group that could be easily derived from the Oligocene apes known from the Fayum of Egypt. Pliopithecus from the Miocene of Europe is more closely allied with the Oligocene radiation than with the later East African radiation.     

Anthropoid versus strepsirhine status of the African Eocene primates Algeripithecus and Azibius: craniodental evidence

Recent fossil discoveries have demonstrated that Africa and Asia were epicentres for the origin and/or early diversification of the major living primate lineages, including both anthropoids (monkeys, apes and humans) and crown strepsirhine primates (lemurs, lorises and galagos). Competing hypotheses favouring either an African or Asian origin for anthropoids rank among the most hotly contested issues in paleoprimatology. The Afrocentric model for anthropoid origins rests heavily on the >45 Myr old fossil Algeripithecus minutus from Algeria, which is widely acknowledged to be one of the oldest known anthropoids. However, the phylogenetic position of Algeripithecus with respect to other primates has been tenuous because of the highly fragmentary fossils that have documented this primate until now. Recently recovered and more nearly complete fossils of Algeripithecus and contemporaneous relatives reveal that they are not anthropoids. New data support the idea that Algeripithecus and its sister genus Azibius are the earliest offshoots of an Afro–Arabian strepsirhine clade that embraces extant toothcombed primates and their fossil relatives. Azibius exhibits anatomical evidence for nocturnality. Algeripithecus has a long, thin and forwardly inclined lower canine alveolus, a feature that is entirely compatible with the long and procumbent lower canine included in the toothcomb of crown strepsirhines. These results strengthen an ancient African origin for crown strepsirhines and, in turn, strongly challenge the role of Africa as the ancestral homeland for anthropoids.     

Neogene paleobiogeography and East African paleoenvironments: contributions from the Tugen Hills rodents and lagomorphs.

A minimum of 28 genera of rodents and one genus of lagomorph were recovered from the Tugen Hills, Baringo District, Kenya, from localities dating from over 15.5 to about 4.4 Ma. The middle Miocene (sites dated between 15.8 and 15.3 Ma) rodent fauna recovered primarily from the Kipsaramon site complex, Muruyur Formation, includes a mixture of characteristically early Miocene taxa, and more derived forms. Composition of the African rodent fauna changes dramatically with the introduction of myocricetodontines, democricetodontines, and dendromurines, immigrants primarily from southern Asia. In the Tugen Hills, these taxa are first found in the Kabasero localities, Ngorora Formation, at sites dating from 12.5-12.33 Ma. A second major change in the African rodent fauna reflects the introduction of murines, immigrants from southern Asia. In the Tugen Hills murines are first encountered at Kapcheberek, Lukeino Formation, dated to 5.9-5.7 Ma. One rodent genus from the Lukeino Formation (Arvicanthis), and two from the Tabarin locality, Chemeron Formation (Heliosciurus, Paraxerus; 4.5-4.4 Ma), represent the earliest records of these extant African genera. A cricetomyine from the Ngorora Formation (12.5 Ma) is likely the earliest report of this exclusively African group. One of the earliest African records of porcupines (Hystricide) is from the Lukeino Formation. Lagomorphs are poorly represented, but include one of the earliest African occurrences of the family Leporidae from the Mpesida Beds (bracketed by dates of 7-6.2 Ma), and possibly a new genus of leporid from the Kapcheberek locality. Analysis of the Tugen Hills small mammals in association with other African records suggests several episodes of dispersal between Africa and Eurasia during the middle and late Miocene. Rodents from Kipsaramon are indicative of forests in conjunction with more open habitats. Those from the Kapcheberek locality are suggestive of a savanna habitat. The rodents from the Tabarin locality suggest a woodland environment.     

Victoriapithecus: The key to Old World monkey and catarrhine origins

The past ten years have witnessed major changes in reconstructions of the history of Old World monkeys, most of them driven by new material of the Miocene monkey Victoriapithecus from Maboko Island, Kenya. Before the mid-1980s, predictions about the morphological and ecological adaptations of the earliest cercopithecoids relied heavily on evidence from extant colobine and cercopithecine monkeys. It was argued that the earliest cercopithecoids were largely or at least partly folivorous, had short colobine-like faces, and were arboreal. The only studies suggesting that some of these arguments were not true were based on limited knowledge of the anatomy of Victoriapithecus. The presence of semi-terrestrial adaptations in middle Miocene monkeys hinted to some that early monkeys may not have been arboreal. Others attempted to cope with the discrepancy between neontological predictions and the fossil evidence by proposing that limb bones with stronger terrestrial adaptations within the Maboko sample were derived cercopithecine remains, while those with more arboreal features belonged in the subfamily Colobinae and should be regarded as primitive.     

A taxonomic revision of the small catarrhine primates from the early Miocene of East Africa

This paper presents a detailed systematic revision of the small catarrhine primates from the early Miocene of East Africa, recovered from sites in Western Kenya and in Uganda dated at between 22 and 17 m.y. Revised diagnoses and amended hypodigms for each of the species are presented. In addition to the currently identified taxa, Limnopithecus legetet Hopwood, 1933, Dendropithecus macinnesi (Le Gros Clark and Leakey, 1950) and Micropithecus clarki Fleagle and Simons, 1978, two further species are recognized. Limnopithecus evansi (MacInnes, 1943) is resurrected as a valid species, based primarily on previously described material from Songhor, and a new genus, Kalepithecus, is described here for the first time, in order to accommodate distinctive material from Songhor and Koru. The distribution of each species in time and space, and the phylogenetic relationships, are discussed in the light of this taxonomic revision.     

Geology and fauna of the middle Miocene hominoid site at Muruyur,Baringo district,Kenya

The middle Miocene sediments assigned to the Muruyur Beds have yielded abundant faunal remains which indicate an age somewhere near the early part of the middle Miocene, perhaps being earlier in time than Fort Ternan but probably coeval or slightly later than Maboko. Available radioisotopic age determinations suggest that the beds are between 13.5 and 14 m.y. old, which seems to be too young when compared with the biostratigraphic estimate. The importance of Muruyur Beds lies in their rich fossil content which includes hominoids of an age which is in general poorly represented in East Africa’s fossil record. This article places the fossil discoveries on record, and discusses their geological context.