Early anthropoid primates: New data and new questions.

Although the evolutionary history of anthropoid primates (monkeys, apes, and humans) appears relatively well-documented, there is limited data available regarding their origins and early evolution. We review and discuss here the earliest records of anthropoid primates from Asia, Africa, and South America. New fossils provide strong support for the Asian origin of anthropoid primates. However, the earliest recorded anthropoids from Africa and South America are still subject to debate, and the early evolution and dispersal of platyrhines to South America remain unclear. Because of the rarity and incomplete nature of many stem anthropoid taxa, establishing the phylogenetic relationships among the earliest anthropoids remains challenging. Nonetheless, by examining evidence from anthropoids and other mammalian groups, we demonstrate that several dispersal events occurred between South Asia and Afro-Arabia during the middle Eocene to the early Oligocene. It is possible that a microplate situated in the middle of the Neotethys Ocean significantly reduced the distance of overseas dispersal.

An extinct deep-snouted Alligator species from the Quaternary of Thailand and comments on the evolution of crushing dentition in alligatorids.

Fossil Alligator remains from Asia are critical for tracing the enigmatic evolutionary origin of the Chinese alligator, Alligator sinensis, the only living representative of Alligatoridae outside the New World. The Asian fossil record is extremely scarce and it remains unknown whether A. sinensis is an anagenetic lineage or alternatively, extinct divergent species were once present. We provide a detailed comparative description of a morphologically highly distinct Alligator skull from the Quaternary of Thailand. Several autapomorphic characters warrant the designation of a new species. Alligator munensis sp. nov. shares obvious derived features with A. sinensis but autapomorphies imply a cladogenetic split, possibly driven by the uplift of the southeastern Tibetan plateau. The presence of enlarged posterior alveoli in Alligator munensis is most consistent with a reversal to the alligatorine ancestral condition of having crushing dentition, a morphology strikingly absent among living alligatorids. Crushing dentition has been previously considered to indicate an ecological specialisation in early alligatorines that was subsequently lost in Alligator spp. However, we argue that there is yet no evidence for crushing dentition reflecting an adaptation for a narrower niche, while opportunistic feeding, including seasonal utilisation of hard-shelled preys, is a reasonable alternative interpretation of its function.

A new parapithecine (Primates: Anthropoidea) from the early Oligocene of Libya supports parallel evolution of large body size among parapithecids.

Parapithecines are an extinct subfamily of stem anthropoid primates previously known only from the Jebel Qatrani Formation in Egypt. Here, we describe isolated teeth pertaining to Simonsius harujensis sp. nov., a relatively small-bodied parapithecine from strata near Zallah Oasis in the Sirt Basin of central Libya that is estimated to date to ∼31 Ma on the basis of mammalian biostratigraphy. The dental morphology of S. harujensis sp. nov. is generally intermediate between that of the closely related parapithecines Parapithecus fraasi and Simonsius grangeri, highlighting some of the anatomical features distinguishing the latter taxa and providing further support for their generic separation. A phylogenetic analysis using parsimony methods was performed on a character-taxon matrix incorporating data from the new Libyan parapithecine, virtually all other parapithecids and the proteopithecid Proteopithecus sylviae. Results of this analysis suggest that parapithecids comprise a basal clade consisting of three species of Biretia and a more derived clade including Parapithecinae (Parapithecus and Simonsius) and Qatraniinae (Qatrania, Ucayalipithecus, and Apidium). Body mass estimates for parapithecids were calculated on the basis of regression equations generated to predict body mass from the occlusal area of upper and lower cheek teeth in extant anthropoids. The relatively small body mass of S. harujensis sp. nov. and its reconstructed phylogenetic position as the sister group of S. grangeri, which is the largest known parapithecid, support the convergent acquisition of body mass larger than 500 g among multiple clades of early Oligocene African anthropoids. The new Libyan parapithecine augments previously reported evidence supporting a substantial degree of faunal provincialism across northern Africa/Arabia during the early Oligocene.

A new primate from the late Eocene of Vietnam illuminates unexpected strepsirrhine diversity and evolution in Southeast Asia.

Sivaladapidae is a poorly known Asian strepsirrhine family originally discovered in Miocene sediments of the Indian subcontinent. Subsequent research has considerably increased the diversity, temporal range, and geographical distribution of this group, now documented from China, Thailand, Myanmar, Pakistan, and India and whose earliest representatives date back to the Middle Eocene. We present here a new taxon of sivaladapid from the Na Duong coal mine in the Latest Middle Eocene-Late Eocene of Vietnam. It represents the first Eocene primate from Vietnam and the first medium-sized mammal recovered from this locality, thus documenting a completely new part of the Na Duong paleobiodiversity. This taxon is the largest sivaladapid ever found with an estimated body weight of 5.3 kg and it represents a new subfamily of sivaladapids in exhibiting a very peculiar combination of dental features yet unknown in the fossil record of the family (e.g., retention of four premolars, high-crowned molars with accentuated bunodonty and extreme crest reduction). Besides documenting a complete new part of sivaladapid evolution, its primitive dental formula and derived features shared with the Early Eocene Asiadapidae reinforce the hypothesis of a basal branching of sivaladapids among strepsirrhines.

First maxilla of a late Miocene hominid from Thailand and the evolution of pongine derived characters.

The evolutionary history of Asian Miocene hominids (great apes and humans) remains poorly documented, obscuring the ancestry of orangutan (Pongo). Khoratpithecus from the middle and late Miocene of Thailand and Myanmar was previously documented only by mandibles and isolated teeth. It has been interpreted as the closest relative of Pongo based on shared derived mandible characters such as symphyseal morphology and the lack of anterior digastric muscle scars. Here we describe a new maxilla, MFT-K176, which originates from the same sedimentary unit as the holotype mandible of Khoratpithecus piriyai from the late Miocene in Nakhon Ratchasima province, Northeastern Thailand. The new maxilla displays a unique subnasal morphology with several derived characters being shared only with Sivapithecus and Pongo, confirming its attribution to the pongine clade. However, it differs from other known Asian hominids by it subnasal and dental morphology, showing more similar to Khoratpithecus chiangmuanensis teeth. Metric similarities with the mandible of K. piriyai corroborate the referral of MFT-K176 to Khoratpithecus. However, as associated upper and lower teeth would be required to evaluate the latter assertion more conclusively, we attribute it provisionally to cf. Khoratpithecus sp. Other anatomical characters from the clivus, the palate and the molars are peculiar for this hominid but do not exclude it from a sister group relationship with Pongo. This new maxilla plays a pivotal role in understanding that Ankarapithecus occupies a more basal position within the pongine clade and supports the exclusion of Lufengpithecus from this clade.

New Eocene primate from Myanmar shares dental characters with African Eocene crown anthropoids.

Recent discoveries of older and phylogenetically more primitive basal anthropoids in China and Myanmar, the eosimiiforms, support the hypothesis that Asia was the place of origins of anthropoids, rather than Africa. Similar taxa of eosimiiforms have been discovered in the late middle Eocene of Myanmar and North Africa, reflecting a colonization event that occurred during the middle Eocene. However, these eosimiiforms were probably not the closest ancestors of the African crown anthropoids. Here we describe a new primate from the middle Eocene of Myanmar that documents a new clade of Asian anthropoids. It possesses several dental characters found only among the African crown anthropoids and their nearest relatives, indicating that several of these characters have appeared within Asian clades before being recorded in Africa. This reinforces the hypothesis that the African colonization of anthropoids was the result of several dispersal events, and that it involved more derived taxa than eosimiiforms.

The Middle Pleistocene vertebrate fauna from Khok Sung (Nakhon Ratchasima, Thailand): biochronological and paleobiogeographical implications.

The fluviatile terrace deposits of Khok Sung, Nakhon Ratchasima province, have yielded more than one thousand fossils, making this the richest Pleistocene vertebrate fauna of Thailand. The excellent preservation of the specimens allows precise characterization of the faunal composition. The mammalian fauna consists of fifteen species in thirteen genera, including a primate, a canid, a hyaenid, proboscideans, rhinoceroses, a suid, cervids, and bovids. Most species correspond to living taxa but globally (Stegodon cf. orientalis) and locally (Crocuta crocuta ultima, Rhinoceros unicornis, Sus barbatus, and Axis axis) extinct taxa were also present. The identification of Axis axis in Khok Sung, a chital currently restricted to the Indian Subcontinent, represents the first record of the species in Southeast Asia. Three reptilian taxa: Crocodylus cf. siamensis, Python sp., and Varanus sp., are also identified. Faunal correlations with other Southeast Asian sites suggest a late Middle to early Late Pleistocene age for the Khok Sung assemblage. However, the Khok Sung mammalian fauna is most similar to that of Thum Wiman Nakin, dated to older than 169 ka. The Khok Sung large mammal assemblage mostly comprises mainland Southeast Asian taxa that migrated to Java during the latest Middle Pleistocene, supporting the hypothesis that Thailand was a biogeographic pathway for the Sino-Malayan migration event from South China to Java.

A new species of Apidium (Anthropoidea, Parapithecidae) from the Sirt Basin, central Libya: First record of Oligocene primates from Libya.

A new species of Apidium is the most common primate currently known from a newly discovered site near Zallah Oasis in the Sirt Basin of central Libya. Based on current knowledge of the associated fauna, this new species of Apidium is early Oligocene in age, being roughly contemporaneous with faunas from Quarries G and V in the upper part of the Jebel Qatrani Formation in Egypt that also contain species of Apidium. A phylogenetic analysis based on dental characters indicates that the new species of Apidium from Libya is the sister group of Apidium phiomense. Apidium bowni and Apidium moustafai from the Jebel Qatrani Formation in the Fayum are similar in age to the new species of Apidium from Libya, but both of these Egyptian species are more distantly related to A. phiomense from younger stratigraphic levels in the Fayum. This phylogenetic pattern underscores the benefit of enhanced geographic sampling of the fossil record, even in cases where local records are thought to be reasonably comprehensive and well documented. Oligocene parapithecids can be partitioned into two clades corresponding to the subfamilies Parapithecinae (containing Parapithecus and Simonsius) and Qatraniinae (including Qatrania and Apidium). Climatic deterioration during the early Oligocene may have impacted the macroevolutionary dynamics of early Afro-Arabian anthropoids by fostering the fragmentation of forest habitats, thereby promoting allopatric speciation among widespread populations of Apidium and other arboreal taxa.

A new small pliopithecoid primate from the Middle Miocene of Thailand.

Pliopithecoids represent a monophyletic group of putative stem catarrhines whose evolutionary history is incompletely known. They have been recorded from Europe and Asia, between the late Early Miocene and the Late Miocene. Asian pliopithecoids are less well documented than their European counterparts, often being represented by a fragmentary fossil record. New discoveries are therefore critical to reconstruct the evolutionary history of the whole group. Here, we describe two isolated molars from Ban San Klang, a late Middle Miocene locality in northern Thailand, which confirms the presence of pliopithecoids in Southeast Asia. The lower molar had originally been described as being that of a dendropithecoid, but it was later recognized as pertaining to a pliopithecoid. The discovery, in the same locality, of an additional upper molar attributed to the same species confirms the pliopithecoid status of this taxon and highlights its distinctiveness with respect to other known Asian pliopithecoids. However, the mosaic of primitive and autapomorphic features characterizing this Thai fossil, as well as its limited anatomical representation, preclude us from assigning it to either of the known pliopithecid subfamilies. Nevertheless, it represents the only pliopithecoid in Southeast Asia and displays a mosaic of unique characters which emphasizes the peculiarity of that province, as suggested previously with respect to its hominoid primate.

A new Late Eocene primate from the Krabi Basin (Thailand) and the diversity of Palaeogene anthropoids in southeast Asia.

According to the most recent discoveries from the Middle Eocene of Myanmar and China, anthropoid primates originated in Asia rather than in Africa, as was previously considered. But the Asian Palaeogene anthropoid community remains poorly known and inadequately sampled, being represented only from China, Myanmar, Pakistan and Thailand. Asian Eocene anthropoids can be divided into two distinct groups, the stem group eosimiiforms and the possible crown group amphipithecids, but the phylogenetic relationships between these two groups are not well understood. Therefore, it is critical to understand their evolutionary history and relationships by finding additional fossil taxa. Here, we describe a new small-sized fossil anthropoid primate from the Late Eocene Krabi locality in Thailand, Krabia minuta, which shares several derived characters with the amphipithecids. It displays several unique dental characters, such as extreme bunodonty and reduced trigon surface area, that have never been observed in other Eocene Asian anthropoids. These features indicate that morphological adaptations were more diversified among amphipithecids than was previously expected, and raises the problem of the phylogenetic relations between the crown anthropoids and their stem group eosimiiforms, on one side, and the modern anthropoids, on the other side.

Uniquely derived upper molar morphology of Eocene Amphipithecidae (Primates: Anthropoidea): homology and phylogeny.

The extinct Southeast Asian primate family Amphipithecidae is regularly cited in discussions of anthropoid origins, but its phylogenetic position remains controversial. In part, the lack of consensus regarding amphipithecid relationships can be attributed to uncertainty regarding the homology of upper molar structures in this group. Here, we describe a virtually pristine upper molar of Pondaungia cotteri from the late middle Eocene Pondaung Formation of Myanmar, which is the first example of a relatively unworn and well-preserved amphipithecid upper molar ever recovered. The distolingual upper molar cusp in this new specimen of Pondaungia appears to be a lingually displaced and enlarged metaconule, rather than a hypocone or pseudohypocone as previous workers have thought. Reassessment of the upper molar morphology of other amphipithecids and putative amphipithecids reveals a very similar pattern in Siamopithecus, Myanmarpithecus and Ganlea, all of which are interpreted as having upper molars showing many of the same derived features apparent in Pondaungia. In contrast, the upper molar morphology of Bugtipithecus diverges radically from that of undoubted amphipithecids, and the latter taxon is excluded from Amphipithecidae on this basis. Phylogenetic analyses of several character-taxon matrices culled from the recent literature and updated to reflect the new information on amphipithecid upper molar morphology yield similar results. Consensus tree topologies derived from these analyses support amphipithecid monophyly and stable relationships within Amphipithecidae. Amphipithecids appear to be stem members of the anthropoid clade.

New Apterodontinae (Hyaenodontida) from the Eocene locality of Dur At-Talah (Libya): systematic, paleoecological and phylogenetical implications.

The African Hyaenodontida, mainly known from the Late Eocene and Early Oligocene Fayum depression in Egypt, show a very poor diversity in oldest Paleogene localities. Here we report new hyaenodontidans found in the late Middle Eocene deposits of Dur At-Talah (Central Libya), known to have recorded the earliest radiation of African anthropoids. The new hyaenodontidan remains are represented by dental and postcranial specimens comprising the historical material discovered by R.J.G. Savage in the last century and that of the recent Franco-Libyan campaigns. This material includes two apterodontines, in particular a subcomplete skeleton of Apterodon langebadreae nov. sp., bringing new postcranial elements to the fossil record of the genus Apterodon. Anatomical analysis of the postcranial remains of Dur At-Talah suggests a semi-aquatic lifestyle for Apterodon, a completely unusual locomotion pattern among hyaenodontidans. We also perform the first cladistic analysis of hyaenodontidans including apterodontines: Apterodon and Quasiapterodon appear close relatives to "hyainailourines", in particular to the African Oligo-Miocene Metasinopa species. Apterodon langebadreae nov. sp. could be the most primitive species of the genus, confirming an African origin of the Apterodontinae and a further dispersion event to Europe before the early Oligocene. These data enhance our knowledge of early hyaenodontidan diversification into Africa and underline how crucial is the understanding of their evolutionary history for the improvement of Paleogene paleobiogeographic scenarii.

Late Middle Eocene primate from Myanmar and the initial anthropoid colonization of Africa.

Reconstructing the origin and early evolutionary history of anthropoid primates (monkeys, apes, and humans) is a current focus of paleoprimatology. Although earlier hypotheses frequently supported an African origin for anthropoids, recent discoveries of older and phylogenetically more basal fossils in China and Myanmar indicate that the group originated in Asia. Given the Oligocene-Recent history of African anthropoids, the colonization of Africa by early anthropoids hailing from Asia was a decisive event in primate evolution. However, the fossil record has so far failed to constrain the nature and timing of this pivotal event. Here we describe a fossil primate from the late middle Eocene Pondaung Formation of Myanmar, Afrasia djijidae gen. et sp. nov., that is remarkably similar to, yet dentally more primitive than, the roughly contemporaneous North African anthropoid Afrotarsius. Phylogenetic analysis suggests that Afrasia and Afrotarsius are sister taxa within a basal anthropoid clade designated as the infraorder Eosimiiformes. Current knowledge of eosimiiform relationships and their distribution through space and time suggests that members of this clade dispersed from Asia to Africa sometime during the middle Eocene, shortly before their first appearance in the African fossil record. Crown anthropoids and their nearest fossil relatives do not appear to be specially related to Afrotarsius, suggesting one or more additional episodes of dispersal from Asia to Africa. Hystricognathous rodents, anthracotheres, and possibly other Asian mammal groups seem to have colonized Africa at roughly the same time or shortly after anthropoids gained their first toehold there.

Analysis of dental root apical morphology: a new method for dietary reconstructions in primates.

The reconstruction of paleo-diets is an important task in the study of fossil primates. Previously, paleo-diet reconstructions were performed using different methods based on extant primate models. In particular, dental microwear or isotopic analyses provided accurate reconstructions for some fossil primates. However, there is sometimes difficult or impossible to apply these methods to fossil material. Therefore, the development of new, independent methods of diet reconstructions is crucial to improve our knowledge of primates paleobiology and paleoecology. This study aims to investigate the correlation between tooth root apical morphology and diet in primates, and its potential for paleo-diet reconstructions. Dental roots are composed of two portions: the eruptive portion with a smooth and regular surface, and the apical penetrative portion which displays an irregular and corrugated surface. Here, the angle formed by these two portions (aPE), and the ratio of penetrative portion over total root length (PPI), are calculated for each mandibular tooth root. A strong correlation between these two variables and the proportion of some food types (fruits, leaves, seeds, animal matter, and vertebrates) in diet is found, allowing the use of tooth root apical morphology as a tool for dietary reconstructions in primates. The method was then applied to the fossil hominoid Khoratpithecus piriyai, from the Late Miocene of Thailand. The paleo-diet deduced from aPE and PPI is dominated by fruits (>50%), associated with animal matter (1-25%). Leaves, vertebrates and most probably seeds were excluded from the diet of Khoratpithecus, which is consistent with previous studies.

Three-dimensional analysis of mandibular dental root morphology in hominoids.

Although often preserved in the fossil record, mandibular dental roots are rarely used for evolutionary studies. This study qualitatively and quantitatively characterizes the three-dimensional morphology of hominoid dental roots. The sample comprises extant apes as well as two fossil species, Khoratpithecus piriyai and Ouranopithecus macedoniensis. The morphological differences between extant genera are observed, quantified and tested for their potential in systematics. Dental roots are imaged using X-ray computerized tomography, conventional microtomography and synchrotron microtomography. Resulting data attest to the high association between taxonomy and tooth root morphology, both qualitatively and quantitatively. A cladistic analysis based on the dental root characters resulted in a tree topology congruent with the consensus phylogeny of hominoids, suggesting that tooth roots might provide useful information in reconstructing hominoid phylogeny. Finally, the evolution of the dental root morphology in apes is discussed.

First hominoid from the Late Miocene of the Irrawaddy Formation (Myanmar).

For over a century, a Neogene fossil mammal fauna has been known in the Irrawaddy Formation in central Myanmar. Unfortunately, the lack of accurately located fossiliferous sites and the absence of hominoid fossils have impeded paleontological studies. Here we describe the first hominoid found in Myanmar together with a Hipparion (s.l.) associated mammal fauna from Irrawaddy Formation deposits dated between 10.4 and 8.8 Ma by biochronology and magnetostratigraphy. This hominoid documents a new species of Khoratpithecus, increasing thereby the Miocene diversity of southern Asian hominoids. The composition of the associated fauna as well as stable isotope data on Hipparion (s.l.) indicate that it inhabited an evergreen forest in a C3-plant environment. Our results enlighten that late Miocene hominoids were more regionally diversified than other large mammals, pointing towards regionally-bounded evolution of the representatives of this group in Southeast Asia. The Irrawaddy Formation, with its extensive outcrops and long temporal range, has a great potential for improving our knowledge of hominoid evolution in Asia.

First castorid (mammalia, rodentia) from the middle miocene of southeast Asia.

Today and in the Tertiary, the geographical distribution of castorids is limited throughout all of the northern continents. Fossils of the Castoridae genus Steneofiber are abundant in many localities of Eurasia from the late Oligocene to Pliocene period. Recently, Steneofiber fossils were discovered in two localities of northern Thailand, Mae Moh and Chiang Muan coal mines, in layers of late middle Miocene age. These discoveries represent the first records of castorids from Southeast Asia and correspond to their southernmost known range. The focus of this study is to describe this new Thai species of Steneofiber and to define its wear stages from the molar occlusal surfaces by using micro-CT scan analysis. The CT scan technique permits the analysis of the virtual occlusal surface changes from wear, allowing easier comparison to related species of Steneofiber cheek teeth without destroying the teeth. The new species, Steneofiber siamensis n. sp., can be distinguished from the other species of Steneofiber by several distinct characters, longer mesostriid on p4, presence of premesostria and metastria on P4, which are smaller than most of the other known species. The occurrence of this new castorid also supports a subtropical to tropical paleoclimate for these two localities of northern Thailand.

A new Middle Miocene tarsier from Thailand and the reconstruction of its orbital morphology using a geometric-morphometric method.

Tarsius is an extant genus of primates endemic to the islands of Southeast Asia that is characterized by enormously enlarged orbits reflecting its nocturnal activity pattern. Tarsiers play a pivotal role in reconstructing primate phylogeny, because they appear to comprise, along with Anthropoidea, one of only two extant haplorhine clades. Their fossils are extremely rare. Here, we describe a new species of Tarsius from the Middle Miocene of Thailand. We reconstructed aspects of its orbital morphology using a geometric-morphometric method. The result shows that the new species of Tarsius had a very large orbit (falling within the range of variation of modern Tarsius) with a high degree of frontation and a low degree of convergence. Its relatively divergent lower premolar roots suggest a longer mesial tooth row and therefore a longer muzzle than in extant species. The new species documents a previous unknown Miocene group of Tarsius, indicating greater taxonomic diversity and morphological complexity during tarsier evolution. The current restriction of tarsiers to offshore islands in Southeast Asia appears to be a relatively recent phenomenon.

Late middle Eocene epoch of Libya yields earliest known radiation of African anthropoids.

Reconstructing the early evolutionary history of anthropoid primates is hindered by a lack of consensus on both the timing and biogeography of anthropoid origins. Some prefer an ancient (Cretaceous) origin for anthropoids in Africa or some other Gondwanan landmass, whereas others advocate a more recent (early Cenozoic) origin for anthropoids in Asia, with subsequent dispersal of one or more early anthropoid taxa to Africa. The oldest undoubted African anthropoid primates described so far are three species of the parapithecid Biretia from the late middle Eocene Bir El Ater locality of Algeria and the late Eocene BQ-2 site in the Fayum region of northern Egypt. Here we report the discovery of the oldest known diverse assemblage of African anthropoids from the late middle Eocene Dur At-Talah escarpment in central Libya. The primate assemblage from Dur At-Talah includes diminutive species pertaining to three higher-level anthropoid clades (Afrotarsiidae, Parapithecidae and Oligopithecidae) as well as a small species of the early strepsirhine primate Karanisia. The high taxonomic diversity of anthropoids at Dur At-Talah indicates either a much longer interval of anthropoid evolution in Africa than is currently documented in the fossil record or the nearly synchronous colonization of Africa by multiple anthropoid clades at some time during the middle Eocene epoch.

New data on Mustelidae (Carnivora) from Southeast Asia: Siamogale thailandica, a peculiar otter-like mustelid from the late middle Miocene Mae Moh Basin, northern Thailand.

We report new dental remains of Mustelidae from the late middle Miocene of Mae Moh Basin, northern Thailand, improving the poor fossil record of the family in Southeast Asia. Siamogale thailandica is a poorly known mustelid, previously recorded from just a single tooth. Here we present over a hundred new specimens attributable to this species. S. thailandica shows a combination of primitive and convergent features of the dentition that makes its original subfamilial assignment to Lutrinae doubtful. Evidence from the dental morphology suggests that it belongs to a bunodont otter-like mustelid that evolved in convergence with "true" otters (Lutrinae) toward a semi-aquatic way of life. Autapomorphic features such as the height and the position of the m1 metaconid and the shape of the P4 lingual shelf make S. thailandica unique among Mustelidae. The morphology of this species is mostly similar to Mionictis species and Lartetictis dubia, reported in the Miocene of North America and Europe, respectively. These similarities could imply immigration events to Thailand in the early or middle Miocene. Alternately, the lineage leading to Siamogale might have deeper origins from an endemic early Miocene Southeast Asian mustelid.