A Review of the Fossil Record of the Genus Itea (Iteaceae, Saxifragales) with Comments on its Historical Biogeography

Itea is a genus of about 20 species of trees and shrubs that are today native to southeastern North America, eastern Asia, and eastern Africa. In this paper, I review the fossil record of Itea, which is based on four types of fossils: diporate, psilate pollen attributed to Itea or the dispersed pollen genus Iteapollis; carpofossils representing fruits and seeds attributed to Itea europaea; flowers preserved in amber and assigned to Adenanthemum iteoides; and leaf impressions attributed to Itea. The distributions of these fossils indicate that Itea was present in western North America from the early Eocene to Miocene, in eastern North America beginning no later than the early Miocene, and in western Eurasia from the late Eocene to Pliocene. Only one datapoint is known from eastern Asia; it is early Miocene in age. Based on the fossil record, it can be inferred that Itea crossed between continents over both the Bering Land Bridge and North American Land Bridge, and that it reached Africa from Europe via Anatolia. Thus, it is predicted that the sole extant North American species, I. virginica, may be most closely related to the sole extant African species, I. rhamnoides. The potential application of Itea fossils to calibrating phylogenetic trees generated from molecular sequence data is also discussed.     

Exploring the Formation of a Disjunctive Pattern between Eastern Asia and North America Based on Fossil Evidence from Thuja (Cupressaceae)

Thuja, a genus of Cupressaceae comprising five extant species, presently occurs in both East Asia (3 species) and North America (2 species) and has a long fossil record from Paleocene to Pleistocene in the Northern Hemisphere. Two distinct hypotheses have been proposed to account for the origin and present distribution of this genus. Here we recognize and describe T. sutchuenensis Franch., a new fossil Thuja from the late Pliocene sediments of Zhangcun, Shanxi, North China, based on detailed comparisons with all living species and other fossil ones, integrate the global fossil records of this genus plotted in a set of paleomaps from different time intervals, which show that Thuja probably first appeared at high latitudes of North America in or before the Paleocene. This genus reached Greenland in the Paleocene, then arrived in eastern Asia in the Miocene via the land connection between East Asia and western North America. In the late Pliocene, it migrated into the interior of China. With the Quaternary cooling and drying, Thuja gradually retreated southwards to form today’s disjunctive distribution between East Asia and North America.     

Evolution of the second orangutan: phylogeny and biogeography of hominid origins

Aim To resolve the phylogeny of humans and their fossil relatives (collectively, hominids), orangutans (Pongo) and various Miocene great apes and to present a biogeographical model for their differentiation in space and time. Location Africa, northern Mediterranean, Asia. Methods Maximum parsimony analysis was used to assess phylogenetic relationships among living large‐bodied hominoids (= humans, chimpanzees, bonobos, gorillas, orangutans), and various related African, Asian and European ape fossils. Biogeographical characteristics were analysed for vicariant replacement, main massings and nodes. A geomorphological correlation was identified for a clade we refer to as the ‘dental hominoids’, and this correlation was used to reconstruct their historical geography. Results Our analyses support the following hypotheses: (1) the living large‐bodied hominoids represent a monophyletic group comprising two sister clades: humans + orangutans, and chimpanzees (including bonobos) + gorillas (collectively, the African apes); and (2) the human–orangutan clade (dental hominoids) includes fossil hominids (Homo, australopiths, Orrorin) and the Miocene‐age apes Hispanopithecus, Ouranopithecus, Ankarapithecus, Sivapithecus, Lufengpithecus, Khoratpithecus and Gigantopithecus (also Plio‐Pleistocene of eastern Asia). We also demonstrate that the distributions of living and fossil genera are largely vicariant, with nodes of geographical overlap or proximity between Gigantopithecus and Sivapithecus in Central Asia, and between Pongo, Gigantopithecus, Lufengpithecus and Khoratpithecus in East Asia. The main massing is represented by five genera and eight species in East Asia. The dental hominoid track is spatially correlated with the East African Rift System (EARS) and the Tethys Orogenic Collage (TOC). Main conclusions Humans and orangutans share a common ancestor that excludes the extant African apes. Molecular analyses are compromised by phenetic procedures such as alignment and are probably based on primitive retentions. We infer that the human–orangutan common ancestor had established a widespread distribution by at least 13 Ma. Vicariant differentiation resulted in the ancestors of hominids in East Africa and various primarily Miocene apes distributed between Spain and Southeast Asia (and possibly also parts of East Africa). The geographical disjunction between early hominids and Asian Pongo is attributed to local extinctions between Europe and Central Asia. The EARS and TOC correlations suggest that these geomorphological features mediated establishment of the ancestral range.     

Phylogeny and biogeography of tropical carnivorous land‐snails (Pulmonata: Streptaxoidea) with particular reference to East Africa and the Indian Ocean

Rowson, B., Tattersfield, P. & Symondson, W. O. C. (2010). Phylogeny and biogeography of tropical carnivorous land‐snails (Pulmonata: Streptaxoidea) with particular reference to East Africa and the Indian Ocean. —Zoologica Scripta, 40, 85–98. A phylogeny is presented for the speciose, near pan‐tropical, carnivorous achatinoid land‐snail superfamily Streptaxoidea inferred from DNA sequences (two nuclear and two mitochondrial regions) from 114 taxa from Africa, the Indian Ocean, Asia, South America and Europe. In all analyses, Streptaxidae are monophyletic, while the (two to six) previously recognised subfamilies are polyphyletic, as are several genus‐level taxa including the most speciose genus Gulella, necessitating major taxonomic review. The Asian Diapheridae are sister to Streptaxidae, which forms several well‐supported clades originating in a persistent basal polytomy. Divergence dating estimates, historical biogeography, and the fossil context suggest a Cretaceous origin of these families, but suggest Gondwanan vicariance predated most radiation. The basal polytomy dates to the Paleogene and may correspond to a rapid radiation in Africa. There is evidence for multiple Cenozoic dispersals followed by radiation, including at least two from Africa to South America, at least two from Africa to Asia and at least two from Africa to Madagascar, indicating Cenozoic turnover in tropical snail faunas. The endemic Seychelles and Mascarene streptaxid faunas each are composites of early Cenozoic lineages and more recent dispersals from Africa, with no direct evidence for an Asian origin as currently proposed. Peak streptaxid diversity in East Africa is explained by Neogene speciation among a large number of coexisting ancient lineages, a phenomenon most pronounced in the Eastern Arc‐Coastal Forests centre of endemism. This includes Miocene diversification in Gulella, a primarily East and South‐East African group which remains strikingly diverse even after unrelated ‘Gulella’ species are reclassified.     

Hagenia from the early Miocene of Ethiopia: Evidence for possible niche evolution?

Fossil pollen believed to be related to extant Hagenia abyssinica were discovered in the early Miocene (21.73 Ma) Mush Valley paleoflora, Ethiopia, Africa. Both the fossil and extant pollen grains of H. abyssinica were examined with combined light microscopy, scanning electron microscopy, and transmission electron microscopy to compare the pollen and establish their relationships. Based on this, the fossil pollen grains were attributed to Hagenia. The presence of Hagenia in the fossil assemblage raises the questions if its habitat has changed over time, and if the plants are/were wind pollinated. To shed light on these questions, the morphology of extant anthers was also studied, revealing specialized hairs inside the anthers, believed to aid in insect pollination. Pollen and anther morphology are discussed in relation to the age and origin of the genus within a molecular dated phylogenetic framework, the establishment of complex topography in East Africa, other evidence regarding pollination modes, and the palynological record. The evidence presented herein, and compiled from the literature, suggests that Hagenia was an insect‐pollinated lowland rainforest element during the early Miocene of the Mush Valley. The current Afromontane habitat and ambophilous (insect and wind) pollination must have evolved in post‐mid‐Miocene times.     

The first Loranthaceae fossils from Africa

An ongoing re-investigation of the early Miocene Saldanha Bay (South Africa) palynoflora, using combined light and scanning electron microscopy (single grain method), is revealing several pollen types new to the African fossil record. One of the elements identified is Loranthaceae pollen. These grains represent the first and only fossil record of Loranthaceae in Africa. The fossil pollen grains resemble those produced by the core Lorantheae and are comparable to recent Asian as well as some African taxa/lineages. Molecular and fossil signals indicate that Loranthaceae dispersed into Africa via Asia sometime during the Eocene. The present host range of African Loranthaceae and the composition of the palynoflora suggest that the fossil had a range of potential host taxa to parasitise during the early Miocene in the Saldanha Bay region.     

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.     

Fossil Ptelea samaras (Rutaceae) in North America

Fossil samaras referred to the extant rutaceous genus Ptelea L. comprise an artificial collection of five distinct taxa with convergent fruit morphologies. Critical reexamination of these fossils revealed that only one of the five samara types can be referred to Ptelea. Two of the forms can be assigned without reservation to Tiliaceae (Pteleaecarpum = Craigia) or Caprifoliaceae (“Ptelea” cassioides [part], “Ptelea” miocenica [part]). Taxonomic affinities of the two remaining fruit types (“Ptelea“ eocenica, “Ptelea“ cassioides [part]) are undetermined. Genuine fossil Ptelea samaras are restricted to Miocene floras in the western United States, including the Hog Creek (Weiser) and Succor Creek floras near the Oregon–Idaho border and the Stinking Water flora of east-central Oregon. These samaras bear the names P. miocenica Berry and P. enervosa H. V. Smith. The holotype of P. miocenica is a caprifoliaceous infructescence similar to that of Dipelta Maximowicz, so P. enervosa, previously considered a synonym of P. miocenica, is the valid combination for fossil Ptelea samaras. P. enervosa samaras are the oldest reliably determined fossil Ptelea fruits. These samaras indicate that the Toddalieae-Pteleinae lineage of subfamily Toddalioideae has been present in North America since the Miocene, rather than the Eocene as suggested by previously misidentified specimens.     

Inflorescence bracts of fossil and extant Tilia in North America,Europe, and Asia: patterns of morphologic divergence and biogeographic history

The enlarged inflorescence bract diagnostic of extant Tilia has an extensive Tertiary fossil record in the Northern Hemisphere. Diversity of bract morphology, and the extent of adnation between peduncle and bract, is reviewed for fossil and extant species of Tilia. An extinct type of bract with an orbicular outline and palmate venation is documented by the fossil species Tilia circularis (Chaney) comb. nov. from the early Oligocene of Oregon and is designated Type A. Living species of the genus have elongate bracts with predominately pinnate venation that are borne in two basic configurations: Type B, with the peduncle fused only to the extreme base of the bract lamina, as in extant Tilia endochrysea Hand.-Mzt. of southern China; and Type C with the peduncle fused medially along the basal one-third of the bract lamina, as in most extant species. Bracts of Type B were widely distributed in the Tertiary of western North America (late Eocene to Miocene) and Europe (early Miocene to Pliocene), while those of Type C are known in the fossil condition only from the middle and late Tertiary of Asia and Pliocene of Europe. The bracts of T. circularis, like those of type B, are borne on relatively long stalks and have the peduncle fused only at the extreme base. The fossil record supports recognition of the following characters as apomorphic in Tilia bract evolution: bracts sessile, peduncle adnate to the upper surface of the bract, and pinnate bract venation.     

Phytogeographical Implication of Bridelia Will. (Phyllanthaceae) Fossil Leaf from the Late Oligocene of India

Background

The family Phyllanthaceae has a predominantly pantropical distribution. Of its several genera, Bridelia Willd. is of a special interest because it has disjunct equally distributed species in Africa and tropical Asia i.e. 18–20 species in Africa-Madagascar (all endemic) and 18 species in tropical Asia (some shared with Australia). On the basis of molecular phylogenetic study on Bridelia, it has been suggested that the genus evolved in Southeast Asia around 33±5 Ma, while speciation and migration to other parts of the world occurred at 10±2 Ma. Fossil records of Bridelia are equally important to support the molecular phylogenetic studies and plate tectonic models.

Results

We describe a new fossil leaf of Bridelia from the late Oligocene (Chattian, 28.4–23 Ma) sediments of Assam, India. The detailed venation pattern of the fossil suggests its affinities with the extant B. ovata, B. retusa and B. stipularis. Based on the present fossil evidence and the known fossil records of Bridelia from the Tertiary sediments of Nepal and India, we infer that the genus evolved in India during the late Oligocene (Chattian, 28.4–23 Ma) and speciation occurred during the Miocene. The stem lineage of the genus migrated to Africa via “Iranian route” and again speciosed in Africa-Madagascar during the late Neogene resulting in the emergence of African endemic clades. Similarly, the genus also migrated to Southeast Asia via Myanmar after the complete suturing of Indian and Eurasian plates. The emergence and speciation of the genus in Asia and Africa is the result of climate change during the Cenozoic.

Conclusions

On the basis of present and known fossil records of Bridelia, we have concluded that the genus evolved during the late Oligocene in northeast India. During the Neogene, the genus diversified and migrated to Southeast Asia via Myanmar and Africa via “Iranian Route”.     

A South American snake lineage from the Eocene Greenhouse of North America and a reappraisal of the fossil record of “anilioid” snakes

“Anilioidea” is a likely paraphyletic assemblage of pipe snakes that includes extant Aniliidae from equatorial South America, Uropeltoidea from South and Southeast Asia, and a fossil record that consists primarily of isolated precloacal vertebrae ranging from the earliest Late Cretaceous and includes geographic distributions in North America, South America, Europe, and Africa. Articulated precloacal vertebrae from the middle Eocene Bridger Formation of Wyoming, attributed to Borealilysia nov. gen., represent an unambiguous North American aniliid record and prompts a reconsideration of described pipe snakes and their resultant biogeographic histories. On the basis of vertebral apomorphies, the vast majority of reported fossils cannot be assigned to “Anilioidea”. Instead, most records represent stem taxa and macrostomatans erroneously assigned to anilioids on the basis of generalized features associated with fossoriality. A revised fossil record demonstrates that the only extralimital distributions of fossil “anilioids” consist of the North American aniliid record, and there is no unambiguous fossil record of Old World taxa. The occurrence of aniliids in the mid-high latitudes of the late early Eocene of North America is consistent with histories of northward shifts in equatorial ecosystems during the early Paleogene Greenhouse.     

<Emphasis Type="Italic">Altingioxylon hainanensis</Emphasis> sp. nov.: earliest fossil wood record of the family Altingiaceae in Eastern Asia and its implications for historical biogeography

A new species, Altingioxylon hainanensis, is described from the Eocene Changchang Formation of the Changchang Basin on Hainan Island, South China. It is the first record of a fossil wood assigned to Altingiaceae found in China, and the most ancient evidence of wood for this family in eastern Asia. The new species is similar to A. rhodoleioides, known since the Miocene in India and Java Island, and to Altingia hisauchii from the Miocene to Pliocene of Japan. The close resemblance between these species and Liquidambar sp., known from the Middle Miocene of western North America, provides additional evidence for the migration of their ancestors from Asia to North America across the Bering land bridge during the Miocene. Distinctions in ray sizes between the eastern Asian specimens and their contemporaries from Europe to Kazakhstan is suggested as a result of the divergence between the large eastern Asian clade and the North American–west Asian clade within Altingiaceae during the Eocene–Oligocene. The presence of crystals in ray cells may be considered an ancestral condition that persists in the eastern Asian lineages up to the extant Altingia and Semiliquidambar, but which was lost in other Altingiaceae in the course of evolution.     

福建漳浦中中新世几种藓类植物研究北大核心

亚洲新生代藓类植物化石非常稀少,漳浦琥珀是目前已知的我国低纬度地区唯一含苔藓植物化石的新生代琥珀矿床。漳浦琥珀源自中中新世气候适宜期(~14-17 Ma)的龙脑香树脂,温暖潮湿的热带季雨林气候和广泛分布的龙脑香科植物,为藓类植物的生长、多样性演化和三维立体保存提供了条件。本研究共报道了漳浦县中中新世藓类植物化石3个属种,包括:白发藓属Leucobryum Hampe、牛舌藓属Anomodon Hook.et Taylor和扁枝藓属Homalia(Brid.)B.S.G.。首次利用三维X射线断层扫描技术(Micro-CT)研究了白发藓属化石,发现其与现生种短枝白发藓Leucobryum humillimum Cardot非常相似,因此将其命名为短枝白发藓(相似种)Leucobryum cf.humillimum。此外,当前牛舌藓属Anomodon和扁枝藓属Homalia化石是东亚地区的首例记录。漳浦琥珀中的藓类植物化石揭示了中中新世时期苔藓植物的多样性,丰富了东亚低纬度地区新生代苔藓植物的化石记录。     

Diverse fossil Onagraceae pollen from a Miocene palynoflora of north-east China: early steps in resolving the phytogeographic history of the family

The origin and evolution of angiosperms can be unravelled by using fossil records to determine first occurrences and phytogeographic histories of plant families and genera. Many angiosperm families, for example the Onagraceae, have a poor macrofossil record, but are more common in palynological records. Modern Onagraceae produce pollen clearly distinct from that of other angiosperms. Combined morphological features obtained by use of light and scanning electron microscopy have enabled assignment of fossil Onagraceae pollen to extant genera, and therefore tracing of the origin and past distributions of extant Onagraceae lineages. We studied a Miocene palynoflora from the Daotaiqiao Formation of north-east China. Using the single-grain technique, we examined individual Onagraceae pollen/tetrads using both light and scanning electron microscopy. Fossil Onagraceae pollen is more frequent than macrofossil remains, but is still rare, and usually represented by a single taxon in palynological samples. Remarkably, samples from the Miocene of north-east China contain five different species: two of Circaea, one of Epilobium, and two of Ludwigia. Such a large number of Onagraceae taxa from a single palynoflora is unknown elsewhere. Whereas Ludwigia pollen is known from Cenozoic sediments of the northern hemisphere, the Circaea pollen is the first fossil pollen assignable to this extant genus. This is also the first fossil record of Epilobium from China. Although the young geological age of this sample does not enable consideration of time of origin for the genera encountered, the co-occurrence of Circaea, Epilobium, and Ludwigia in the mid to late-Miocene of East Asia sheds some light on their phytogeographic histories.     

India at the cross-roads of human evolution

The Indian palaeoanthropological record, although patchy at the moment, is improving rapidly with every new find. This broad review attempts to provide an account of (a) the Late Miocene fossil apes and their gradual disappearance due to ecological shift from forest dominated to grassland dominated ecosystem around 9-8 Ma ago, (b) the Pliocene immigration/evolution of possible hominids and associated fauna, (c) the Pleistocene record of fossil hominins, associated fauna and artifacts, and (d) the Holocene time of permanent settlements and the genetic data from various human cultural groups within India. Around 13 Ma ago (late Middle Miocene) Siwalik forests saw the emergence of an orangutan-like primate Sivapithecus. By 8 Ma, this genus disappeared from the Siwalik region as its habitat started shrinking due to increased aridity influenced by global cooling and monsoon intensification. A contemporary and a close relative of Sivapithecus, Gigantopithecus (Indopithecus), the largest ape that ever-lived, made its first appearance at around 9 Ma. Other smaller primates that were pene-contemporaneous with these apes were Pliopithecus (Dendropithecus), Indraloris, Sivaladapis and Palaeotupia. The Late Pliocene and Early Pleistocene witnessed northern hemisphere glaciations, followed by the spread of arid conditions on a global scale, setting the stage for hominids to explore “Savanahastan”. With the prominent expansion of grassland environments from East Africa to China and Indonesia in the Pliocene, monkeys and baboons dispersed into the Indian subcontinent from Africa along with other mammals. Though debated, there are several claims of the presence of early hominins in this part of the world during the Late Pliocene, based primarily on the recovery of Palaeolithic tools. Fossils of our own ancestor and one of the first globe-trotters, early Homo erectus, has been documented from the Early Pleistocene of East Africa, Western Asia and Southeast Asia, thus indirectly pointing towards Indian subcontinent as a possible migration corridor between these regions. The only definite pre-Homo sapiens fossil hominin remains come from the Central Narmada Valley and are thought to be of Middle to late Pleistocene age, and the cranium has been shown to be closely linked to archaic Homo sapiens/H. heidelbergensis of Europe. Around ∼74,000 yrs ago, a super volcanic eruption in Sumatra caused the deposition of Youngest Toba Tephra, that covered large parts of the Indian peninsula. Just around this time anatomically-and-behaviorally modern humans or Homo sapiens possibly arrived into India as evidenced by the so called Middle and Upper Palaeolithic assemblages and associated symbolic evidence. The available genetic data reveals that the gene pool to which modern Indians races belong was extremely diverse and had variable mixed links with both European and Asian populations.     

A new species of Macaranga from the middle Miocene of Fujian,China and its significance

A well-preserved Macaranga leaf fossil from the middle Miocene Fotan Group of Zhangpu County, Fujian, South-eastern China is examined and described as a new species, Macaranga zhangpuensis Z.X. Wang et B.N. Sun sp. nov. The present fossil species represents the highest latitudinal distribution of a reliable Macaranga fossil in the world, and we present the first Macaranga fossil described with detailed cuticular characteristics from China. Based on the global palaeogeographic distribution of Macaranga, we infer that the genus probably originated during the Oligocene in Africa and spread from Africa to India and then to South Fujian, China, further into the Mariana Islands and finally into the Philippine Islands and Malaysia, leading to its present distribution. In addition, the new leaf material described herein is the first Macaranga fossil record with three leaf tips globally. This finding demonstrates that Macaranga with three leaf tips already existed during the Miocene in Fujian, South China, and it provides new information for understanding climatic changes between the Miocene and the present -day.     

Rudabánya: A late miocene subtropical swamp deposit with evidence of the origin of the African apes and humans

Rudabánya, a rich late Miocene fossil site in northern central Hungary, has yielded an abundant record of fossil primates, including the primitive catarrhine Anapithecus and the early great ape Dryopithecus. While the affinities of Anapithecus are not clear, Dryopithecus is clearly a great ape sharing numerous characteristics of its dental, cranial and postcranial anatomy with living great apes. Like all Miocene hominids (great apes and humans), Dryopithecus is more primitive in a number of ways than any living hominid, which is probably related to the passage of time since the divergence of the various lineages of living hominids, allowing for similar refinements in morphology and adaptation to take place independently. On the other hand, Dryopithecus (and Ouranopithecus) share derived characters with hominines (African apes and humans), and Sivapithecus (and Ankarapithecus) share derived characters with orangutans, thus dating the split between pongines and hominines to a time before the evolution of these fossil great apes. Pongines and hominines follow similar fates in the late Miocene, the pongines moving south into Southeast Asia from southern or eastern Asia and the hominines moving south into East Africa from the Mediterranean region, between 6 to 9 Ma.     

Oldest known Varanus (Squamata: Varanidae) from the Upper Eocene and Lower Oligocene of Egypt: support for an African origin of the genus

Abstract: A large collection of lizard vertebrae from northern Africa represents the oldest unambiguous occurrence of the genus Varanus. The fossils come from late Eocene and early Oligocene freshwater deposits of the Fayum, Egypt, an area noted for many significant primate finds. The recovery and identification of this material indicate that the genus Varanus arose in Africa, before dispersing to Australia and Asia. This dispersal occurred prior to the early to mid‐Miocene, by which time fossil Varanus are known from Australia and Eurasia. Although the dispersal route remains unknown, the lizard material reported here supports the hypothesis that a corridor existed allowing freshwater and terrestrial organisms to cross from Africa to Asia.     

The Hand of Cercopithecoides williamsi (Mammalia,Primates): Earliest Evidence for Thumb Reduction among Colobine Monkeys

Thumb reduction is among the most important features distinguishing the African and Asian colobines from each other and from other Old World monkeys. In this study we demonstrate that the partial skeleton KNM-ER 4420 from Koobi Fora, Kenya, dated to 1.9 Ma and assigned to the Plio-Pleistocene colobine species Cercopithecoides williamsi, shows marked reduction of its first metacarpal relative to the medial metacarpals. Thus, KNM-ER 4420 is the first documented occurrence of cercopithecid pollical reduction in the fossil record. In the size of its first metacarpal relative to the medial metacarpals, C. williamsi is similar to extant African colobines, but different from cercopithecines, extant Asian colobines and the Late Miocene colobines Microcolobus and Mesopithecus. This feature clearly links the genus Cercopithecoides with the extant African colobine clade and makes it the first definitive African colobine in the fossil record. The postcranial adaptations to terrestriality in Cercopithecoides are most likely secondary, while ancestral colobinans (and colobines) were arboreal. Finally, the absence of any evidence for pollical reduction in Mesopithecus implies either independent thumb reduction in African and Asian colobines or multiple colobine dispersal events out of Africa. Based on the available evidence, we consider the first scenario more likely.