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The bat genus Myotis is represented by 120+ living species and 40+ extinct species and is found on every continent except Antarctica. The time of divergence of Myotis has been contentious as has the time and place of origin of its encompassing group the Vespertilionidae, the most diverse (450+ species) and widely distributed extant bat family. Fossil Myotis species are common, especially in Europe, beginning in the Miocene but earlier records are poor. Recent study of new specimens from the Belgian early Oligocene locality of Boutersem reveals the presence of a relatively large vespertilionid. Morphological comparison and phylogenetic analysis confirms that the new, large form can be confidently assigned to the genus Myotis, making this record the earliest known for that taxon and extending the temporal range of this extant genus to over 33 million years. This suggests that previously published molecular divergence dates for crown myotines (Myotis) are too young by at least 7 million years. Additionally, examination of first fossil appearance data of 1,011 extant placental mammal genera indicates that only 13 first occurred in the middle to late Paleogene (48 to 33 million years ago) and of these, six represent bats, including Myotis. Paleogene members of both major suborders of Chiroptera (Yangochiroptera and Yinpterochiroptera) include extant genera indicating early establishment of successful and long-term adaptive strategies as bats underwent an explosive radiation near the beginning of the Early Eocene Climatic Optimum in the Old World. A second bat adaptive radiation in the New World began coincident with the Mid-Miocene Climatic Optimum.

In 1967 G.G. Simpson described three partial mandibles from early Miocene deposits in Kenya that he interpreted as belonging to a new strepsirrhine primate, Propotto . This interpretation was quickly challenged, with the assertion that Propotto was not a primate, but rather a pteropodid fruit bat. The latter interpretation has not been questioned for almost half a century. Here we re-evaluate the affinities of Propotto , drawing upon diverse lines of evidence to establish that this strange mammal is a strepsirrhine primate as originally suggested by Simpson. Moreover, our phylogenetic analyses support the recognition of Propotto , together with late Eocene Plesiopithecus from Egypt, as African stem chiromyiform lemurs that are exclusively related to the extant aye-aye ( Daubentonia ) from Madagascar. Our results challenge the long-held view that all lemurs are descended from a single ancient colonization of Madagascar, and present an intriguing alternative scenario in which two lemur lineages dispersed from Africa to Madagascar independently, possibly during the later Cenozoic. The fossil taxon Propotto was originally identified as a primate, but is currently widely interpreted as a bat. Here, the authors identify Propotto as a stem chiromyiform lemur and, based on phylogenetic analysis, suggest two independent lemur colonizations of Madagascar.

Flight first The Green River formation in Wyoming has produced many important fossils, including Icaronycteris index , which for over 40 years has been regarded as the oldest known bat. Its cranial features suggest that it could locate its insect prey by echolocation. This fuelled a spirited debate between proponents of the 'flight-first', 'echolocation-first' and 'tandem-development' hypotheses of bat evolution. New Green River bat fossils — including two near-complete skeletons, a cast of one of which is shown on the cover — looks to have settled the matter in favour of flight first. The new species is the most primitive bat known. It had fully developed wings and was clearly capable of powered flight, but the morphology of the ear region suggests that it could not echolocate, making it a possible intermediate link between bats and their non-flying, non-echolocating mammalian ancestors. Limb characteristics, including robust hind legs and retention of tiny claws on all of its elongate fingers, indicate that the new bat may have been an agile climber. Bats (Chiroptera) represent one of the largest and most diverse radiations of mammals, accounting for one-fifth of extant species 1 . Although recent studies unambiguously support bat monophyly 2 , 3 , 4 and consensus is rapidly emerging about evolutionary relationships among extant lineages 5 , 6 , 7 , 8 , the fossil record of bats extends over 50 million years, and early evolution of the group remains poorly understood 5 , 7 , 8 , 9 . Here we describe a new bat from the Early Eocene Green River Formation of Wyoming, USA, with features that are more primitive than seen in any previously known bat. The evolutionary pathways that led to flapping flight and echolocation in bats have been in dispute 7 , 8 , 9 , 10 , 11 , 12 , 13 , 14 , 15 , 16 , 17 , 18 , and until now fossils have been of limited use in documenting transitions involved in this marked change in lifestyle. Phylogenetically informed comparisons of the new taxon with other bats and non-flying mammals reveal that critical morphological and functional changes evolved incrementally. Forelimb anatomy indicates that the new bat was capable of powered flight like other Eocene bats, but ear morphology suggests that it lacked their echolocation abilities, supporting a ‘flight first’ hypothesis for chiropteran evolution. The shape of the wings suggests that an undulating gliding–fluttering flight style may be primitive for bats, and the presence of a long calcar indicates that a broad tail membrane evolved early in Chiroptera, probably functioning as an additional airfoil rather than as a prey-capture device. Limb proportions and retention of claws on all digits indicate that the new bat may have been an agile climber that employed quadrupedal locomotion and under-branch hanging behaviour.

Myzopodidae is a family of bats today represented by two extant species of the genus Myzopoda that are restricted to the island of Madagascar. These bats possess uniquely derived adhesive pads on their thumbs and ankles that they use for clinging to smooth roosting surfaces. Only one fossil myzopodid has been reported previously, a humerus from Pleistocene deposits at Olduvai Gorge in Tanzania that was tentatively referred to the genus Myzopoda. Here we describe a new genus and two new species of myzopodids based on dental remains from Paleogene deposits in the Fayum Depression in Egypt, and provide an emended diagnosis for the family Myzopodidae. Phasmatonycteris phiomensis n. sp. is represented by four specimens from the early Oligocene Jebel Qatrani Formation and P. butleri n. sp. is known from a single specimen from the late Eocene Birket Qarun Formation. Together these specimens extend the temporal range of Myzopodidae by 36+ million years, and the geographic range by nearly 4000 kilometers. The new myzopodids, along with previously described bats from the Fayum and Australia, suggest that eastern Gondwana played a critical role in the origin and diversification of several bats clades notably including the superfamily Noctilionoidea, the majority of which live in the Neotropics today.

Borings in fossil turtle shells collected from the lowermost beds of the early Eocene Cathedral Bluffs Tongue of the Wasatch Formation in the northwestern part of the Green River Basin near South Pass, Wyoming, are herein described. Individual turtle shells in the study area are characterized by as few as one or two and as many as >100 borings. The borings include both non-penetrative forms (those which do not pass fully though the shell) as well as penetrative forms (those which pass fully from the exterior to the interior surface of the shell). All non-penetrative forms occur on external surfaces of the carapace and plastron (i.e. those that would have been accessible while the host taxon was alive). Two new ichnogenera and four new ichnospecies are established to describe these borings. Karethraichnus (new ichnogenus) includes three ichnospecies: K. lakkos (new ichnospecies), K. kulindros (new ichnospecies), and K. fiale (new ichnospecies). Karethraichnus lakkos are shallow (non-penetrating), hemispherical pits with rounded, to flattened bases. Karethraichnus kulindros are deep, non-penetrative traces with a cylindrical profile, an axis approximately perpendicular to the substrate surface and with rounded to flattened, hemispherical termini. Karethraichnus fiale are penetrative traces with a cylindrical to bi-convex or flask-shaped profile, and an axis approximately perpendicular to the substrate surface. Thatchtelithichnus (new ichnogenus) Thatchtelithichnus holmani (new ichnospecies) consist of non-penetrative borings into a bone substrate. They consist of a ring-shaped trace, with a central pedestal or platform. The position of the borings on the shells, and evidence of syn-emplacement healing of the borings in several of the turtles, indicates that these borings were emplacement by ectoparasites/mesoparasites while the animals were living. Similar traces in modern emydid turtles are attributed to ticks, leeches, or spirorchid liver flukes.

The modern effect of climate on plants and animals is well documented. Some have cautioned against assigning climate a direct role in Cenozoic land mammal faunal changes. We illustrate 3 episodes of significant mammalian reorganization in the Eocene of North America that are considered direct responses to dramatic climatic events. The first episode occurred during the Paleocene-Eocene Thermal Maximum (PETM), beginning the Eocene (55.8 Ma), and earliest Wasatchian North American Land Mammal Age (NALMA). The PETM documents a short (<170 k.y.) global temperature increase of [almost equal to]5 °C and a substantial increase in first appearances of mammals traced to climate-induced immigration. A 4-m.y. period of climatic and evolutionary stasis then ensued. The second climate episode, the late early Eocene Climatic Optimum (EECO, 53-50 Ma), is marked by a temperature increase to the highest prolonged Cenozoic ocean temperature and a similarly distinctive continental interior mean annual temperature (MAT) of 23 °C. This MAT increase [and of mean annual precipitation (MAP) to 150 cm/y) promoted a major increase in floral diversity and habitat complexity under temporally unique, moist, paratropical conditions. Subsequent climatic deterioration in a third interval, from 50 to 47 Ma, resulted in major faunal diversity loss at both continental and local scales. In this Bridgerian Crash, relative abundance shifted from very diverse, evenly represented, communities to those dominated by the condylarth HYOPSODUS: Rather than being \"optimum,\" the EECO began the greatest episode of faunal turnover of the first 15 m.y. of the Cenozoic.

Mammalian extinction worldwide during the Late Pleistocene has been a major focus for Quaternary biochronology and paleoecology. These extinctions have been variably attributed to the impacts of climate change and human interference. However, until relatively recently, research has been largely restricted to the Americas, Europe, and Australasia. We present the oldest Middle–Late Pleistocene stratified and numerically dated faunal succession for the Indian subcontinent from the Billasurgam cave complex. Our data demonstrate continuity of 20 of 21 identified mammalian taxa from at least 100,000 y ago to the present, and in some cases up to 200,000 y ago. Comparison of this fossil record to contemporary faunal ranges indicates some geographical redistribution of mammalian taxa within India. We suggest that, although local extirpations occurred, the majority of taxa survived or adapted to substantial ecological pressures in fragmented habitats. Comparison of the Indian record with faunal records from Southeast and Southwest Asia demonstrates the importance of interconnected mosaic habitats to long-term faunal persistence across the Asian tropics. The data presented here have implications for mammalian conservation in India today, where increasing ecological circumscription may leave certain taxa increasingly endangered in the most densely populated region of the world.

A new genus and species of fossil bat is described from New Zealand’s only pre-Pleistocene Cenozoic terrestrial fauna, the early Miocene St Bathans Fauna of Central Otago, South Island. Bayesian total evidence phylogenetic analysis places this new Southern Hemisphere taxon among the burrowing bats (mystacinids) of New Zealand and Australia, although its lower dentition also resembles Africa’s endemic sucker-footed bats (myzopodids). As the first new bat genus to be added to New Zealand’s fauna in more than 150 years, it provides new insight into the original diversity of chiropterans in Australasia. It also underscores the significant decline in morphological diversity that has taken place in the highly distinctive, semi-terrestrial bat family Mystacinidae since the Miocene. This bat was relatively large, with an estimated body mass of ~40 g, and its dentition suggests it had an omnivorous diet. Its striking dental autapomorphies, including development of a large hypocone, signal a shift of diet compared with other mystacinids, and may provide evidence of an adaptive radiation in feeding strategy in this group of noctilionoid bats.

Primates have long been a test case for the development of phylogenetic methods for divergence time estimation. Despite a large number of studies, however, the timing of origination of crown Primates relative to the Cretaceous–Paleogene (K–Pg) boundary and the timing of diversification of the main crown groups remain controversial. Here, we analysed a data set of 372 taxa (367 Primates and 5 outgroups, 3.4 million aligned base pairs) that includes nine primate genomes. We systematically explore the effect of different interpretations of fossil calibrations and molecular clock models on primate divergence time estimates. We find that even small differences in the construction of fossil calibrations can have a noticeable impact on estimated divergence times, especially for the oldest nodes in the tree. Notably, choice of molecular rate model (autocorrelated or independently distributed rates) has an especially strong effect on estimated times, with the independent rates model producing considerably more ancient age estimates for the deeper nodes in the phylogeny. We implement thermodynamic integration, combined with Gaussian quadrature, in the program MCMCTree, and use it to calculate Bayes factors for clock models. Bayesian model selection indicates that the autocorrelated rates model fits the primate data substantially better, and we conclude that time estimates under this model should be preferred. We show that for eight core nodes in the phylogeny, uncertainty in time estimates is close to the theoretical limit imposed by fossil uncertainties. Thus, these estimates are unlikely to be improved by collecting additional molecular sequence data. All analyses place the origin of Primates close to the K–Pg boundary, either in the Cretaceous or straddling the boundary into the Palaeogene.