The oldest African bat from the early Eocene of El Kohol (Algeria)

The Afro-Arabian Paleogene fossil record of Chiroptera is very poor. In North Africa and Arabia, this record is limited, thus far, to a few localities mainly in Tunisia (Chambi, late early Eocene), Egypt (Fayum, late Eocene to early Oligocene), and Sultanate of Oman (Taqah, early Oligocene). It consists primarily of isolated teeth or mandible fragments. Interestingly, these African fossil bats document two modern groups (Vespertilionoidea and Rhinolophoidea) from the early Eocene, while the bat fossil record of the same epoch of North America, Eurasia, and Australia principally includes members of the ??Eochiroptera.?? This paraphyletic group contains all primitive microbats excluding modern families. In Algeria, the region of Brezina, southeast of the Atlas Mountains, is famous for the early Eocene El Kohol Formation, which has yielded one of the earliest mammalian faunas of the African landmass. Recent fieldwork in the same area has led to the discovery of a new vertebrate locality, including isolated teeth of Chiroptera. These fossils represent the oldest occurrence of Chiroptera in Africa, thus extending back the record of the group to the middle early Eocene (Ypresian) on that continent. The material consists of an upper molar and two fragments of lower molars. The dental character association matches that of ??Eochiroptera.?? As such, although very fragmentary, the material testifies to the first occurrence of ??Eochiroptera?? in Algeria, and by extension in Africa. This discovery demonstrates that this basal group of Chiroptera had a worldwide distribution during the early Paleogene.     

New remains of the enigmatic cetartiodactyl Bugtitherium grandincisivum Pilgrim, 1908, from the upper Oligocene of the Bugti Hills (Balochistan, Pakistan)

Newly discovered fossil material of the enigmatic cetartiodactyl Bugtitherium grandincisivum from the upper Oligocene of the Bugti Member of the Chitarwata Formation in the Bugti Hills (Balochistan, Pakistan) is reported. These new specimens consist of two fragmentary muzzles (one preserving the first incisors and belonging to a juvenile) and a fragmentary right mandible with m3. The morphologies of the anterior dentition and m3 provided by these new specimens confirm the validity of the genus Bugtitherium and advocate probable anthracotheriid affinity for the genus rather than entelodontid or suoid affinities, but do not definitively close the debate about Bugtitherium’s familial affinities within Cetartiodactyla. Although still poorly documented, this large-sized anthracotheriid-like cetartiodactyl is a possible key form for understanding the early evolution of hippos, and, in turn, the ancestry of whales, because of both its morphological similarities with hippos and primitive Paleogene whales and its Tethysian distribution.     

A Phororhacoid bird from the Eocene of Africa

The bird fossil record is globally scarce in Africa. The early Tertiary evolution of terrestrial birds is virtually unknown in that continent. Here, we report on a femur of a large terrestrial new genus discovered from the early or early middle Eocene (between ～52 and 46?Ma) of south-western Algeria. This femur shows all the morphological features of the Phororhacoidea, the so-called Terror Birds. Most of the phororhacoids were indeed large, or even gigantic, flightless predators or scavengers with no close modern analogs. It is likely that this extinct group originated in South America, where they are known from the late Paleocene to the late Pleistocene (～59 to 0.01?Ma). The presence of a phororhacoid bird in Africa cannot be explained by a vicariant mechanism because these birds first appeared in South America well after the onset of the mid-Cretaceous Gondwana break up (～100?million years old). Here, we propose two hypotheses to account for this occurrence, either an early dispersal of small members of this group, which were still able of a limited flight, or a transoceanic migration of flightless birds from South America to Africa during the Paleocene or earliest Eocene. Paleogeographic reconstructions of the South Atlantic Ocean suggest the existence of several islands of considerable size between South America and Africa during the early Tertiary, which could have helped a transatlantic dispersal of phororhacoids.     

Artiodactyls from the Pondaung Formation (Myanmar): new data and reevaluation of the South Asian Faunal Province during the Middle Eocene

Although Asia is thought to have played a critical role in the radiation of artiodactyls, the fossil record of stem selenodonts (“dichobunoids”) remains dramatically poor in tropical Asian regions. In this study, we report a new dichobunid genus and species Cadutherium kyaukmagyii and a new basal ruminant genus and species Irrawadymeryx pondaungi, from the late Middle Eocene Pondaung Formation, Central Myanmar. Although the scarcity of the present material prevents any attempts to investigate the phylogenetic relationships of Cadutherium with contemporaneous forms from other Holarctic landmasses, this new form shed new light on the diversity of these small rabbit-like ungulates during a key period of their evolutionary history. Reexamination of the small-bodied artiodactyls from Pondaung leads us to propose new identifications of certain published specimens and, in turn, to investigate the temporal and geographic distribution of taxa recognized in the Pondaung Formation. Although fragmentary, these potential new taxa reveal an unsuspected diversity of small forms among artiodactyls of Pondaung. This addition to the Eocene record of dichobunoids and early ruminants provides further insight in the diversity of dental patterns among small artiodactyls from the Pondaung Formation and attests to the antiquity of these groups in Southeast Asia.     

Skeletal remains of a diminutive primate from the Paleocene of Germany

Most living mammal orders, including our own, started their career during the first 10 million years of the Cenozoic, the Age of Mammals. The fossil record documents that early Paleogene adaptive radiations of various clades included tiny species of the size of living shrews. Remains of particularly diminutive limb bones are described from the late Paleocene site of Walbeck, Sachsen-Anhalt. Discovered in 1939, it has remained the only known Paleocene mammal-bearing locality from Germany. The remains are referred to the family Adapisoriculidae, which is considered on the basis of the present postcranial evidence to represent plesiadapiform primates rather than alleged lipotyphlan insectivores as previously proposed. The Walbeck fossils compete with the Early Eocene species Toliapina vinealis from Europe and Picromomys petersonorum from North America for the status of the smallest known primate, fossil and living. Their estimated body weights are as small as 10 g. The limb bones show features related to enhanced flexion at the elbow and hip joint, suggesting arboreal habits and environments such as terminal branches. The diminutive size and tooth morphology suggest feeding on small insects and other invertebrates. Postcranials are important to assess early radiations, such tiny specimens as the present ones are extremely scarce in the fossil record, however.     

Bizarre tubercles on the vertebrae of Eocene fossil birds indicate an avian disease without modern counterpart

Remains of fossil birds with numerous bony tubercles on the cervical vertebrae are reported from the Middle Eocene of Messel in Germany and the Late Eocene of the Quercy fissure fillings in France. These structures, which are unknown from extant birds and other vertebrates, were previously described for an avian skeleton from Messel but considered a singular feature of this specimen. The new fossils are from a different species of uncertain phylogenetic affinities and show that tuberculated vertebrae have a wider taxonomic, temporal, and geographic distribution. In contrast to previous assumptions, they are no ontogenetic feature and arise from the vertebral surface. It is concluded that they are most likely of pathologic origin and the first record of a Paleogene avian disease. Their regular and symmetrical arrangement over most of the external vertebral surface indicates a systemic disorder caused by factors that do not affect extant birds, such as especially high-dosed phytohormones or extinct pathogens.     

The oldest European fossil songbird from the early Oligocene of Germany

We report on the oldest European songbird (Passeriformes), from the early Oligocene (30–34 million years ago) of Frauenweiler in Germany. The specimen represents the earliest associated remains of an early Tertiary passerine described so far. It ties the first appearance of Passeriformes in Europe to a minimum age of 30 million years. Passeriform birds are absent in Eocene deposits that yielded abundant remains of small land birds and apparently dispersed into Europe around the Eocene/Oligocene boundary (about 34 million years ago), not at the Oligocene/Miocene boundary (about 24 mya) as hitherto thought. This possibly relates the appearance of songbirds in Europe to a well-known major faunistic break at the Eocene/Oligocene boundary, called the grande coupure. The Frauenweiler songbird most notably differs from extant Passeriformes in having a larger processus procoracoideus on the coracoid and appears to be outside Eupasseres, the taxon which includes Oscines (all modern European and most Old World songbirds) and Suboscines (most South and Central American songbirds). It shows that there were earlier dispersal events of non-oscine songbirds into Europe before the arrival of Oscines from the Australian continental plate towards the late Oligocene.     

Enigmatic ungulate-like mammals from the Eocene of Central Asia

A new mammal family, Olseniidae, is proposed based on a complete foot skeleton of cf. Olsenia sp. from the Eocene Toru Ajgyr locality in Kyrgyzstan and an astragalus of Olsenia mira from the Eocene Shara Murun locality in northern China. The new form cf. Olsenia sp. is an early ungulate that combines characteristics of mesonychids, perissodactyls and artiodactyls: tetradactyl and paraxonic foot, terminal phalanges claw-like but not fissured, astragalus with shallow proximal caput and without distal trochlea. This unique character set fills a gap in the fossil record and gives insights into the ungulate phylogeny, which is still not completely understood.     

A chicken-sized crane precursor from the early Oligocene of France

A skeleton of a chicken-sized crane precursor is described from the Lower Oligocene of the Lubéron in Southern France. Parvigrus pohli gen. et sp. nov. is the most substantial Paleogene fossil record of the Grues (Aramidae [limpkin] + Gruidae [cranes]), and among its oldest representatives. The fossil species is classified in the new taxon Parvigruidae, which is shown to be the sister group of extant Grues. It is the first fossil record of a stem lineage representative of the Grues and, among others, differs from modern Grues in its smaller size, shorter beak, and rail-like limb proportions. Size increase in the stem lineage of the Gruidae may be related to the spread of grasslands during the Oligocene and Miocene. Occurrence of stem lineage Grues in the Lower Oligocene of Europe is in concordance with the fact that there is no evidence for the presence of crown group members of modern avian “families” in pre-Oligocene fossil deposits. Electronic Supplementary Material  Supplementary material is available for this article at     

A new fossil thryonomyid from the Late Miocene of the United Arab Emirates and the origin of African cane rats

Cane rats (Thryonomyidae) are represented today by two species inhabiting sub-Saharan Africa. Their fossil record is predominately African, but includes several Miocene species from Arabia and continental Asia that represent dispersal events from Africa. For example, Paraulacodus indicus, known from the Miocene of Pakistan, is closely related to living Thryonomys. Here we describe a new thryonomyid, Protohummus dango, gen. et sp. nov., from the late Miocene Baynunah Formation of the United Arab Emirates. The new thryonomyid is less derived than “Thryonomys” asakomae from the latest Miocene of Ethiopia and clarifies the origin of crown Thryonomys and the evolutionary transition from Paraulacodus. A phylogenetic analysis shows Protohummus dango to be morphologically intermediate between Paraulacodus spp. and extinct and living Thryonomys spp. The morphological grade and phylogenetic position of Protohummus dango further supports previous biochronological estimates of the age of the Baynunah Formation (ca. 6–8 Ma).     

Spine anatomy reveals the diversity of catfish through time: a case study of Synodontis (Siluriformes)

Synodontis (Mochokidae, Siluriformes) is a freshwater catfish endemic to Africa. The 118 extant species are present in almost all hydrographic basins. Some species are restricted to a single stream, whereas others have a vast distribution. Synodontis is known in the fossil record since the Miocene, and its history depends on the connections among African basins through time. The identification of species in the fossil record is essential to reconstruct this historical pattern. Catfish pectoral and dorsal spines are robust, they preserve well and they form most of the fossil remains for the genus Synodontis. Unfortunately, the criteria for the identification of extant Synodontis species are not applicable to fossil specimens. Here, we define 11 original morphological characters that permit to discriminate four extant species from the Chad-Chari hydrographic system. Six of these characters are defined on pectoral spines and five on dorsal spines. We then show that these characters can be used successfully for identifying fossil specimens. In particular, we present a case study in which we identify Synodontis cf. schall and Brachysynodontis cf. batensoda in the hominid-bearing sector Toros-Menalla (Late Miocene, northern Chad). We show that spine anatomy can be a powerful tool to recognise catfish species through time and thus to identify historical diversity pattern.     

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.     

Survival of Theriosuchus (Mesoeucrocodylia: Atoposauridae) in a Late Cretaceous archipelago: a new species from the Maastrichtian of Romania

Small terrestrial non-eusuchian mesoeucrocodylians are common components of Cretaceous assemblages of Gondwanan provinces with notosuchians and araripesuchids as flagship taxa in South America, Africa and Madagascar, well into the Late Cretaceous. On the other hand, these are exceedingly rare in Laurasian landmasses during the Late Cretaceous. Small terrestrial mesoeucrocodylians from Europe were often referred to the genus Theriosuchus, a taxon with stratigraphic range extending from the Late Jurassic to the late Early Cretaceous. Theriosuchus is abundantly reported from various European localities, although Asiatic and possibly North American members are also known. It has often been closely associated with the first modern crocodilians, members of the Eusuchia, because of the presence of procoelous vertebrae, a widespread key character diagnosing the Eusuchia. Nevertheless, the relationships of Theriosuchus have not been explored in detail although one species, Theriosuchus pusillus, has been extensively described and referred in numerous works. Here, we describe a new basal mesoeucrocodylian, Theriosuchus sympiestodon sp. nov. from the Maastrichtian of the Haţeg Basin, Romania, suggesting a large temporal gap (about 58 myr) in the fossil record of the genus. Inclusion of the new taxon, along with Theriosuchus guimarotae, in a phylogenetic analysis confirms its referral to the genus Theriosuchus, within a monophyletic atoposaurid clade. Although phylogenetic resolution within this clade is still poor, the new taxon appears, on morphological grounds, to be most closely related to T. pusillus. The relationships of Atoposauridae within Mesoeucrocodylia and especially to Neosuchia are discussed in light of the results of the present contribution as well as from recent work. Our results raise the possibility that Atoposauridae might not be regarded as a derived neosuchian clade anymore, although further investigation of the neosuchian interrelationships is needed. Reports of isolated teeth referable to a closely related taxon from the Upper Cretaceous of Romania and France, together with the presence of Doratodon and Ischyrochampsa, indicate a previously unsuspected diverse assemblage of non-eusuchian mesoeucrocodylians in the Late Cretaceous European archipelago.     

Earliest and first Northern Hemispheric hoatzin fossils substantiate Old World origin of a “Neotropic endemic”

The recent identification of hoatzins (Opisthocomiformes) in the Miocene of Africa showed part of the evolution of these birds, which are now only found in South America, to have taken place outside the Neotropic region. Here, we describe a new fossil species from the late Eocene of France, which constitutes the earliest fossil record of hoatzins and the first one from the Northern Hemisphere. Protoazin parisiensis gen. et sp. nov. is more closely related to South American Opisthocomiformes than the African taxon Namibiavis and substantiates an Old World origin of hoatzins, as well as a relictual distribution of the single extant species. Although recognition of hoatzins in Europe may challenge their presumed transatlantic dispersal, there are still no North American fossils in support of an alternative, Northern Hemispheric, dispersal route. In addition to Opisthocomiformes, other avian taxa are known from the Cenozoic of Europe, the extant representatives of which are only found in South America. Recognition of hoatzins in the early Cenozoic of Europe is of particular significance because Opisthocomiformes have a fossil record in sub-Saharan Africa, which supports the hypothesis that extinction of at least some of these “South American” groups outside the Neotropic region was not primarily due to climatic factors.     

First evidence of poisonous shrews with an envenomation apparatus

Herein, we report evidence of an envenomation apparatus (EA) in two different species of extinct “giant” shrews, Beremendia and an indeterminate soricine (Mammalia, Eulipotyphla, Soricidae), documented by very well preserved fossil specimens recovered from two Early Pleistocene cave deposits of the Sierra de Atapuerca in Burgos, Spain. The two soricine taxa from Atapuerca have evolved specialized teeth as EAs, which differ from those of recently reported mammals of the Paleocene age, being more similar to the ones described in the modern Solenodon. This discovery reveals the first instance of shrews possessing what appears to be an EA, an evolutionary adaptation that, in these species, was probably related to an increase in body mass and hunting of a larger-sized prey. The Atapuerca specimens would have a highly specialized EA, one of the very few reported for an extinct or living mammal of any time. In addition to the presence of a gutter-like groove along the medial side of the crown of the lower incisors, these two species also present stout jaws and a modified mandibular symphysis with a conspicuous cavity, which in life would likely contain large amounts of connective tissue. The strong mandible architecture of these large shrews would be, in this way, reinforced by a more immovable symphysis, increasing the bite force exerted over a potential prey. This adaptation, together with the grooved incisors, would ensure a rapid and efficient transmission of the poisonous saliva to paralyze relatively large-sized prey.     

An opilioacarid mite in Cretaceous Burmese amber

A fossil opilioacarid mite (Parasitiformes: Opilioacarida) in Burmese amber is described as ?Opilioacarus groehni sp. nov. This ca. 99 Ma record (Upper Cretaceous: Cenomanian) represents only the third fossil example of this putatively basal mite lineage, the others originating from Eocene Baltic amber (ca. 44–49 Ma). Our new record is not only the oldest record of Opilioacarida, but it is also one of the oldest examples of the entire Parasitiformes clade. The presence of Opilioacarida—potentially Opiloacarus—in the Cretaceous of SE Asia suggests that some modern genus groups were formerly more widely distributed across the northern hemisphere, raising questions about previously suggested Gondwanan origins for these mites.     

Does leaf flushing in the dry season affect leaf traits and herbivory in a tropical dry forest?

We report a small hawk-like diurnal bird from the early Oligocene (30–31 million years ago) of Poland. Aviraptor longicrus, n. gen. et sp. is of a size comparable with the smallest extant Accipitridae. The new species is characterized by very long legs, which, together with the small size, suggest an avivorous (bird-eating) feeding behavior. Overall, the new species resembles extant sparrowhawks (Accipiter spp.) in the length proportions of the major limb bones, even though some features indicate that it convergently acquired an Accipiter-like morphology. Most specialized avivores amongst extant accipitrids belong to the taxon Accipiter and predominantly predate small forest passerines; the smallest Accipiter species also hunts hummingbirds. Occurrence of a possibly avivorous raptor in the early Oligocene of Europe is particularly notable because A. longicrus coexisted with the earliest Northern Hemispheric passerines and modern-type hummingbirds. We therefore hypothesize that the diversification of these birds towards the early Oligocene may have triggered the evolution of small-sized avivorous raptors, and the new fossil may exemplify one of the earliest examples of avian predator/prey coevolution.

     

A revised stratigraphic framework for Later Cenozoic sequences in the northeastern Mediterranean region

This study describes the lithostratigraphic character of mid-Cenozoic (Oligocene–Pliocene) sequences in different parts of the northeastern Mediterranean area and offers a detailed stratigraphic correlation for this region. The sequences concerned are drawn from the Camardi area (south–central Anatolia), the Adana Basin, the Misis Mountains and the Kyrenia Range (northern Cyprus) and the submerged Florence Rise (west of Cyprus). The stratigraphic relationships identified here indicate the following: (a) Following the middle Eocene (Lutetian) regression there was uplift throughout the entire region; (b) Episodes of fluvial and lacustrine deposition in intramontane settings ensued in most of this region during the late Eocene/early Miocene interval; (c) Following a regionally extensive phase of tectonic compression, major marine transgression commenced in the late Oligocene in northern Cyprus and in the early Miocene in adjacent southern Turkey, with the exception of the Ecemis Fault Zone where continental deposition continued; (d) These Oligo-Miocene transgressive sequences comprise a broadly diachronous complex of both shallow and deeper marine facies, including reefal carbonates, littoral clastics, basinal shales and fan-turbidites; (e) Deeper marine Miocene facies persisted longer in the Misis area and in northern Cyprus; (f) A regional regression occurred throughout most of the area during the late Serravallian to Tortonian interval and is marked by the abrupt, locally discordant appearance of extensive shallow marine, deltaic and fluvial deposits; (g) Continued regression in the Messinian led to the formation of significant evaporite deposits in the western and southern parts of the region, but localized uplift of the Misis area is attested by the initial deformation of the Neogene rocks there and the absence of Messinian sediments from this area; (h) In the Pliocene there was extensive emergence of the northern parts of the region interrupted by brief marine incursions. The present-day drainage pattern was established at this time; (i) Marine conditions persisted longer in northern Cyprus, where emergence occurred only in the latest Pliocene.     

Middle-Eocene artiodactyls from Shanghuang (Jiangsu Province,Coastal China) and the diversity of basal dichobunoids in Asia

A new assemblage of basal dichobunoid artiodactyls from the middle-Eocene Shanghuang fissure fillings includes the diacodexeid Jiangsudon shanghuangensis gen. and sp. nov., a new species of the lantianine dichobunoid Elaschitotherium, Elaschitotherium crepaturus sp. nov., and an indeterminate suoid which is presently the earliest record of this clade. Diacodexeids are also represented by two forms provisionally referred to cf. Diacodexis sp. and to an indeterminate Diacodexeidae, respectively. The occurrence of diacodexeids in Shanghuang contrasts with the early and earliest middle-Eocene chronological range of the family in Europe and North America and suggests that the stratigraphic range of the family in Asia extends up to the middle Eocene. This may reflect particular habitats in coastal China that may have been relatively stable during the early and middle Eocene, thus preserving forest-dwelling artiodactyls that became extinct in the other Holarctic regions. Compared to other supposedly coeval North American, European, and Asian faunas, the Shanghuang mammalian assemblage is most similar to early Uintan faunas of North America but is also remarkable in recording forms close to taxa that are characteristic of the Wasatchian and Bridgerian North American Land Mammal Ages. The Irdinmanhan age of the Shanghuang fauna is supported by the mammalian assemblage recovered from the fissure D, but an Arshantan age cannot be completely ruled out at this point. Although the Shanghuang assemblage is biased towards preservation of small components of the mammalian fauna, the Shanghuang fauna provide an important and unique window into the Eocene diversity and early evolution of cetartiodactyls in eastern Asia.     

Euarchontan affinity of Paleocene Afro-European adapisoriculid mammals and their origin in the late Cretaceous Deccan Traps of India

The controversial family Adapisoriculidae, a group of shrew-sized Paleocene mammals, had proposed relationships with insectivores, marsupials and more recently to plesiadapiforms. Adapisoriculid remains are numerous in the early Paleocene locality of Hainin, Belgium, and allow us a test of these different phylogenetic hypotheses. Here, we identify the first tarsal bones of adapisoriculid mammals. The highly specialised bones indicate an arboreal mode of life with euarchontan affinity. Moreover, the tarsal bones are morphologically very close to those of the late Cretaceous Deccanolestes from the Deccan intertrappean beds of India, and also share several characters with the Paleocene plesiadapiforms and the extant cynocephalid dermopterans. The adapisoriculid affinities of Deccanolestes are also confirmed by tooth morphology, indicating that Deccanolestes is a primitive member of this family. These phylogenetic affinities suggest a paleobiogeographic scenario for the family with dispersal either via East Africa or across the Tethys area.