New fossil anthropoids from the middle Miocene of East Africa and their bearing on the origin of the oreopithecidae

Recent paleontological collections at the middle Miocene locality of Maboko Island in Kenya, dated at 15-16 million years, have yielded numerous new specimens belonging to at least five species of fossil anthropoids. The most common species of ape at the site, a medium-sized primate with a very distinctive dental morphology, clearly represents a previously undescribed taxon. When compared with other Miocene anthropoids from East Africa, it has its closest affinities with the poorly known species Rangwapithecus vancouveringi from the early Miocene locality of Rusinga Island. The species from Maboko Island is described here as belonging to a new genus of fossil anthropoid, to which "Rangwapithecus" vancouveringi is also referred. The new genus has a highly distinctive suite of derived characters of its molars and premolars, which it shares with Oreopithecus bambolii from the late Miocene of Europe. These synapomorphies indicate a close phyletic relationship between the East African species and Oreopithecus and form the basis for the inclusion of these taxa in a single family, the Oreopithecidae Schwalbe, 1915. In many respects, however, the East African forms are more conservative than Oreopithecus, and in a general sense they can be regarded as an intermediate grade between Oreopithecus and the more generalized early Miocene catarrhines, the proconsuloids. There is, therefore, good fossil evidence to indicate that the origins of the Oreopithecidae can be traced back to the early Miocene of Africa.     

Paleosols and paleoenvironments of the middle Miocene,Maboko Formation,Kenya

The middle Miocene (15 Ma) Maboko Formation of Maboko Island and Majiwa Bluffs, southwestern Kenya, has yielded abundant fossils of the earliest known cercopithecoid monkey (Victoriapithecus macinnesi), and of a kenyapithecine hominoid (Kenyapithecus africanus), as well as rare proconsuline (Simiolus leakeyorum, cf. Limnopithecus evansi) and oreopithecine apes (Mabokopithecus clarki, M. pickfordi), and galagids (Komba winamensis). Specific habitat preferences can be interpreted from large collections of primate fossils in different kinds of paleosols (pedotypes). Fossiliferous drab-colored paleosols with iron-manganese nodules (Yom pedotype) are like modern soils of seasonally waterlogged depressions (dambo). Their crumb structure and abundant fine root-traces, as well as scattered large calcareous rhizoconcretions indicate former vegetation of seasonally wet, wooded grassland. Other fossiliferous paleosols are evidence of nyika bushland (Ratong), and early-successional riparian woodland (Dhero). No fossils were found in Mogo paleosols interpreted as saline scrub soils. Very shallow calcic horizons (in Yom, Ratong, and Mogo paleosols) and Na-montmorillonite (in Mogo) are evidence of dry paleoclimate (300-500 mm MAP=mean annual precipitation). This is the driest paleoclimate and most open vegetation yet inferred as a habitat for any Kenyan Miocene apes or monkeys. Victoriapithecus was abundant in dambo wooded grassland (Yom) and riparian woodland (Dhero), a distribution like that of modern vervet monkeys. Kenyapithecus ranged through all these paleosols, but was the most common primate in nyika bushland paleosols (Ratong), comparable to baboons and macaques today. Mabokopithecus was virtually restricted to riparian woodland paleosols (Dhero), and Simiolus had a similar, but marginally wider, distribution. Habitat preferences of Mabokopithecus and Simiolus were like those of modern colobus monkeys and mangabeys. A single specimen of Komba was found in dambo wooded grassland paleosol (Yom), a habitat more like that of the living Senegal bushbaby than of rainforest galagids. A shift to non-forest habitats may explain the terrestrial adaptations of Victoriapithecus, basal to the cercopithecid radiation, and of Kenyapithecus, basal to the hominoid radiation. Both taxa are distinct from earlier Miocene arboreal proconsulines, oreopithecines and galagids.     

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.     

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.     

New platyrrhine monkeys from the Solimões Formation (late Miocene, Acre State, Brazil)

We report here a new fossil primate from the late Miocene of Brazil. The material consists of a lower first molar and a maxilla with P3-4. The fossils were collected in the Solim?es Formation at the locality of Patos, upper Acre River, Acre State, Brazil. The locality is assigned to the Huayquerian South American Land Mammal Age based on faunal content (late Miocene; dated to between 9 and 6 Ma). The new material is the oldest known occurrence of fossil primates in Brazil and is recognized as a new genus and species, Solimoea acrensis. Solimoea is the oldest known member of the ateline subfamily, which includes the living genera Ateles, Lagothrix, and Brachyteles. By analogy with the molar structures and diets of extant platyrrhines, Solimoea primarily had a diet of fruit, perhaps similar to that of the spider monkey, Ateles. Two other primate teeth described previously from the same formation in Bolivia document the occurrence of alouattines and cebines. One of those specimens is a late Miocene representative of the middle Miocene Colombian genus Stirtonia. The other represents one of the largest known platyrrhine primates, for which is erected a new primate genus, Acrecebus fraileyi.     

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.     

Early evolution and biogeography of lorisiform strepsirrhines

This brief review summarizes new paleontological and molecular data that together support a late middle Eocene Afro-Arabian origin for crown Lorisiformes. Phylogenetic analysis indicates that late Eocene Karanisia is a possible stem lorisiform, late Eocene Saharagalago and Wadilemur and Miocene Komba are stem galagids, and early Miocene Mioeuoticus and Progalago may be crown lorisids. Character evolution along the lorisid and galagid stem lineages is reconstructed as having occurred primarily in postcranial and dental morphology, respectively. These patterns have important implications for interpreting an early lorisiform fossil record that is still composed primarily of jaws and isolated teeth.     

A new species of Gaimanophis (Serpentes,Boidae) from the Miocene of northwestern Argentina with remarks on the Neogene boids of South America

Gaimanophis is an extinct boid genus represented so far by a single species (Gaimanophis tenuis) known by isolated vertebrae from the early Miocene of Patagonia. In this paper, a new species of Gaimanophis is described from the India Muerta Formation (late Miocene) of Tucumán province (Argentina). Gaimanophis powelli sp. nov. distinguishes itself from G. tenuis mainly in its larger size, prezygapophyses less slanting, neural spine shorter dorsally than ventrally, and zygosphene straight bearing an anteriorly protruding tongue. This record indicates a wider temporal and geographical distribution of the genus from the early Miocene of Patagonia to the late Miocene of northwestern Argentina. The recognition of a new species of boid in South America increases the known diversity of this group. Although boids have inhabited in this territory since the Paleocene, fossils belonging to the group only show a glimpse of the real past diversity in the continent.     

New or little known epiphyllous liverworts,XX Cololejeunea nosykombae A.Szabó & Pócs sp.nov. from Madagascar

A new epiphyllous Cololejeunea species is described from Nosy Komba Island of northwest Madagascar. Two varieties are distinguished, var. nosykombae and var. laevis. The species is distinguished by its acute, swallow-tail shaped perianth wings, unique in the genus, and by the special leaf dentition in its typical variety. The new species seems to be endemic to the small Nosy Komba Island. Conservation aspects are discussed and the new species is considered to be critically endangered.     

Afrochoerodon nov. gen. kisumuensis (MacInnes) (Proboscidea,Mammalia) from Cheparawa,Middle Miocene,Kenya

A proboscidean skull from Cheparawa, (Muruyur Formation, Kenya), differs markedly from those of Eurasian Choerolophodon (C. pentelici, C. dhokpathanensis). It is morphologically and metrically close to the holotype of Choerolophodon kisumuensis (MacInnes) a partial skull from Maboko, much of which has been reconstructed in plaster of Paris. The more complete remains of this species now available indicate that it should be placed in a genus separate from Choerolophodon. The new genus Afrochoerodon is erected for it. Choerolophodon ngorora from Ngorora and Fort Ternan (Kenya), Choerolophodon zaltaniensis from Gebel Zelten (Libya) and Choerolophodon chioticus from Chios, Greece, should be transferred to the genus Afrochoerodon. Late Miocene specimens from Nakali, Kenya are probably referrable to the genus Choerolophodon. Fossils from Burji-Soyama (Ethiopia) hitherto assigned to Choerolophodon sp. are excluded from the subfamily Choerolophodontinae.     

铲齿象一新种在甘肃省党河地区下中新统的发现

记述了甘肃省党河地区的铲齿象化石一新种———党河铲齿象 (Platybelodondangheen sis)。其主要特征是 :下颌联合和下门齿构成宽短的铲形 ;下门齿薄 ,由单层稀少的齿柱组成 ;具较大的p3 ,p4前窄后宽 ,前齿带和跟座较小 ,m1构造简单 ,齿冠较低 ,第一齿脊无前斜脊 ,前齿带和后跟座较小。它是欧亚大陆目前所知时代最早 (早中新世 )、构造最原始的一种铲齿象。     

NEW AUSTRALIAN OLIGOCENE TO MIOCENE RINGTAIL POSSUMS (PSEUDOCHEIRIDAE) AND REVISION OF THE GENUS MARLU

Abstract:  Four genera of extinct ringtail possums have been reported from Australian Oligocene to Miocene sediments since 1987. The genus Marlu was described from two species, M. kutjamarpensis and M. praecursor (Woodburne, Tedford and Archer), from the Miocene Kutjamarpu Local Fauna (Leaf Locality) and late Oligocene Wadikali Local Fauna respectively, of northern South Australia. New fossil material referable to this genus has been collected from the Leaf Locality and the Oligocene to Miocene Riversleigh World Heritage Area in northwest Queensland. Three new species, Marlu karya sp. nov. from middle Miocene Riversleigh local faunas and Marlu syke sp. nov. and Marlu ampelos sp. nov. from the Leaf Locality as well as early to middle Miocene Riversleigh local faunas are described. A revision and rediagnosis of the genus and published species are made following the re-examination of referred material in conjunction with the new material. Marlu is characterized by simple dentition and synapomorphies with extant pseudocheirids such as a conjoined postmetacristid and preentocristid on m1 and the loss of the entostylid ridge. New material from middle Miocene Riversleigh deposits has been referred to M. kutjamarpensis , extending the known distribution and age of that species. Re-examination of M. praecursor has revealed the presence of a small m1 entostylid ridge, contributing further to differences between M. praecursor and all other Marlu species and raising the possibility that Marlu is paraphyletic. The new material does not contradict a sister group relationship between ' Marlu ' (excluding M. praecursor ) and the Pliocene–Pleistocene genus Pseudokoala. Homology of the pseudocheirid m1 protostylid identified in species of Paljara , Pildra and Marlu with that observed in extant species is reconsidered. The 'protostylid' of the extinct genera is herein described as the buccal stylid.     

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.     

Systematics of early and middle Miocene Old World monkeys

New information about the early cercopithecoids Prohylobates tandyi (Wadi Moghra, Egypt) and Prohylobates sp. indet. (Buluk and Nabwal, Kenya) is presented. Comparisons are made among all major collections of Early and Middle Miocene catarrhine monkeys, and a systematic revision of the early Old World monkeys is provided. Previous work involving the systematics of early Old World monkeys (Victoriapithecidae; Cercopithecoidea) has been hampered by a number of factors, including the poor preservation of Prohylobates material from North Africa and lack of comparable anatomical parts across collections. However, it is now shown that basal cercopithecoid species from both northern and eastern Africa can be distinguished from one another on the basis of degree of lower molar bilophodonty, relative lower molar size, occlusal details, symphyseal construction, and mandibular shape. Results of particular interest include: 1) the first identification of features that unambiguously define Prohylobates relative to Victoriapithecus; 2) confirmation that P. tandyi is incompletely bilophodont; and 3) recognition of additional victoriapithecid species.     

青海柴达木盆地晚中新世深沟小哺乳动物群

描述了2002-2005年间在青海德令哈深沟上油砂山组采集的小哺乳动物化石。化石共有16种,隶属食虫目、啮齿目和兔形目的12科,代表了迄今青藏高原发现种类最多、材料最丰富的小哺乳动物群。深沟动物群的组成与陕西蓝田灞河组下部层位发现的小哺乳动物组合最为接近,两者有相似的群体结构,共有Sciurotamias,Lophocricetus,Protalactaga,Myocricetodon,Nannocricetua,Pararhizomys和Ochotona 7属。但由于深沟动物群含有略进步的属、种,其时代似乎稍晚,很可能属晚中新世早期,或者中国陆生哺乳动物时代的保德期早期,与欧洲MN10上部或MN11下部的时代相当。深沟动物群指示了一个温带开阔干旱草原为主体的景观。晚中新世柴达木盆地的气候似乎没有现在这样干燥,在草原中尚有一些灌丛甚至林地。动物群的组成及所指示的生态环境都表明,青藏高原在中新世以后有过较大幅度的上升。深沟动物群中含有一鼠科新种——细弱许氏鼠(Huerzelerimys extguus sp.nov.)。Huerzele—rimys属过去只发现于欧洲,新种为该属在亚洲的首次发现。其特征为:与现知最小种H.minor相比,其臼齿更小,M1中t1的位置相对靠后,c6和t9间有超过50%的标本具一弱脊连接,m1和m2的唇侧齿带狭窄、附尖弱小。Ochotonoma primitiva(Zheng & Li,1982)是动物群中较为多见的一种鼠兔。该种最先发现于甘肃天祝,并作为Ochotona属描述。正型地点的标本不多,但尺寸和形态完全落入深沟标本的变异范围,因此认为同属一种。该种的特征增订为:中等大小的鼠兔。P2冠面长三角形,舌侧长度明显大。p3的下前边尖宽大,一般具有两个前褶或凹槽,而且至少有一褶具水泥质充填物;连接前边尖和后边尖间的齿桥(dentine isthmus)宽;前褶(paraflexis)比O.anatolica的短而狭窄,向后延伸没有O.csarnotana的那样明显。     

Cautious climbing and folivory: a model of hominoid differentation

Despite the large and growing number of Miocene fossil catarrhine taxa, suitable common ancestors of great apes and humans have yet to be agreed upon. Considering a) the conservative and primitive nature of the hominoid molar cusp pattern, and b) the variability of secondary dental features, it is difficult to discern whether a hominoid dentition is primitive, secondarily simplified to the primitive condition or too far derived to be ancestral to any of the living forms. Nonetheless, the inability to recognize a common ancestor is primarly due to the absence of a model of hominoid differentiation that provides a basis for its recognition. Vertical climbing as the limiting component of cautious climbing, explains all of the locomotor anatomy shared by living hominoids. Comparison of the shared derived characters of hominoids to those of forms which have converged on hominoidsi.e colobines, atelines, lorisines, paleopropithecines and sloths suggest that early hominoids were probably folivores. In arboreal forms there is a strong link between a large body size, folivory and cautious climbing. Comparison of craniodental characters of committed folivores to committed frugivores from among each of the compared groups with the exception of lorisines, indicates that many of the distinguishing craniodental characters of humans and great apes are adaptations to folivory. Many of these characters, however, are also present in Jolly's seed eating complex. As such folivory may be the heritage factor which Jolly hypothesized to account for differential reduction of canines in fossilTheropithecus and hominids.     

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.     

A new species of Ameripodius (Aves: Galliformes: Quercymegapodiidae) from the Lower Miocene of France

The Quercymegapodiidae, primitive galliforms resembling recent megapodes, have been described from the Upper Eocene of Quercy, France. They have also been identified in the Upper Oligocene–Lower Miocene of Brazil, where they are represented by the genus Ameripodius Alvarenga. A new species of this genus, Ameripodius alexis sp. nov., from the Lower Miocene of France, is described here. The occurrence of the same genus on both sides of the Atlantic Ocean emphasizes the similarities between South American and European avifaunas during the early Tertiary. New discoveries indicate that a similar avifauna was also present in North America, and that a characteristic association of taxa can be defined for the group that includes South America, North America and Eurasia. However, so far as is known, the same avifauna does not occur in contemporaneous African avifaunas.     

A new pliopithecoid genus from the early Miocene of Uganda

A partial face and mandible from the early Miocene site of Napak IX in Uganda are described here as a new genus and species of catarrhine primate, Lomorupithecus harrisoni gen. et sp. nov. The face is among the most complete specimens known for a Miocene small-bodied catarrhine. Several aspects of its anatomy indicate that the new species is a stem catarrhine, and as such, it may provide valuable information pertaining to the primitive catarrhine cranial morphotype. Lomorupithecus is most similar in its facial anatomy to members of the Pliopithecoidea, and these similarities could be interpreted in three ways. They could be symplesiomorphies, which would support the traditional view of the primitive catarrhine cranial morphotype; they could be synapomorphies reflecting a phylogenetic position of Lomorupithecus within Pliopithecoidea; or they could represent convergence. Phylogenetic analysis of Lomorupithecus along with 35 other primates indicates that it is a pliopithecoid. As such, it would be the oldest and only Afro-Arabian member of this otherwise Eurasian clade.