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The Tibetan Plateau is the youngest and highest plateau on Earth, and its elevation reaches one-third of the height of the troposphere, with profound dynamic and thermal effects on atmospheric circulation and climate. The uplift of the Tibetan Plateau was an important factor of global climate change during the late Cenozoic and strongly influenced the development of the Asian monsoon system. However, there have been heated debates about the history and process of Tibetan Plateau uplift, especially the paleo-altimetry in different geological ages. Here we report a well-preserved skeleton of a 4.6 million-y-old three-toed horse (Hipparion zandaense) from the Zanda Basin, southwestern Tibet. Morphological features indicate that H. zandaense was a cursorial horse that lived in alpine steppe habitats. Because this open landscape would be situated above the timberline on the steep southern margin of the Tibetan Plateau, the elevation of the Zanda Basin at 4.6 Ma was estimated to be ∼4,000 m above sea level using an adjustment to the paleo-temperature in the middle Pliocene, as well as comparison with modern vegetation vertical zones. Thus, we conclude that the southwestern Tibetan Plateau achieved the present-day elevation in the mid-Pliocene.

The ‘third pole’ of the world is a fitting metaphor for the Himalayan–Tibetan Plateau, in allusion to its vast frozen terrain, rivalling the Arctic and Antarctic, at high altitude but low latitude. Living Tibetan and arctic mammals share adaptations to freezing temperatures such as long and thick winter fur in arctic muskox and Tibetan yak, and for carnivorans, a more predatory niche. Here, we report, to our knowledge, the first evolutionary link between an Early Pliocene (3.60–5.08 Myr ago) fox, Vulpes qiuzhudingi new species, from the Himalaya (Zanda Basin) and Kunlun Mountain (Kunlun Pass Basin) and the modern arctic fox Vulpes lagopus in the polar region. A highly hypercarnivorous dentition of the new fox bears a striking resemblance to that of V. lagopus and substantially predates the previous oldest records of the arctic fox by 3–4 Myr. The low latitude, high-altitude Tibetan Plateau is separated from the nearest modern arctic fox geographical range by at least 2000 km. The apparent connection between an ancestral high-elevation species and its modern polar descendant is consistent with our ‘Out-of-Tibet’ hypothesis postulating that high-altitude Tibet was a training ground for cold-environment adaptations well before the start of the Ice Age.

Pantherine felids (‘big cats’) include the largest living cats, apex predators in their respective ecosystems. They are also the earliest diverging living cat lineage, and thus are important for understanding the evolution of all subsequent felid groups. Although the oldest pantherine fossils occur in Africa, molecular phylogenies point to Asia as their region of origin. This paradox cannot be reconciled using current knowledge, mainly because early big cat fossils are exceedingly rare and fragmentary. Here, we report the discovery of a fossil pantherine from the Tibetan Himalaya, with an age of Late Miocene–Early Pliocene, replacing African records as the oldest pantherine. A ‘total evidence’ phylogenetic analysis of pantherines indicates that the new cat is closely related to the snow leopard and exhibits intermediate characteristics on the evolutionary line to the largest cats. Historical biogeographic models provide robust support for the Asian origin of pantherines. The combined analyses indicate that 75% of the divergence events in the pantherine lineage extended back to the Miocene, up to 7 Myr earlier than previously estimated. The deeper evolutionary origin of big cats revealed by the new fossils and analyses indicate a close association between Tibetan Plateau uplift and diversification of the earliest living cats.

Ice Age megafauna have long been known to be associated with global cooling during the Pleistocene, and their adaptations to cold environments, such as large body size, long hair, and snow-sweeping structures, are best exemplified by the woolly mammoths and woolly rhinos. These traits were assumed to have evolved as a response to the ice sheet expansion. We report a new Pliocene mammal assemblage from a high-altitude basin in the western Himalayas, including a primitive woolly rhino. These new Tibetan fossils suggest that some megaherbivores first evolved in Tibet before the beginning of the Ice Age. The cold winters in high Tibet served as a habituation ground for the megaherbivores, which became preadapted for the Ice Age, successfully expanding to the Eurasian mammoth steppe.

All previous records of the lamniform shark, Archaeolamna kopingensis, are based on isolated teeth. Here we describe a partial skeleton from the Sharon Springs Formation of the Pierre Shale Group of western Kansas, U.S.A. The specimen includes portions of the upper and lower jaws with articulated teeth. The dentition consists of two files of upper and lower anterior teeth that, together with a single file of intrabullar intermediate teeth, are housed in a dental bulla, as well as multiple files of lateral teeth, along with at least two files of lower symphysial teeth and a single file of upper symphysial teeth. The intrabullar intermediate tooth is slightly shorter than the other anterior teeth and has a median cusp with distinctive distal curvature. The dental sequence of A. kopingensis is unique among both extinct and extant lamniforms. Associated with the jaws are fragments of the neurocranium and multiple vertebral centra. A sagittal section through a centrum shows that this shark deposited 18 annual marker bands after its birth and adult size was attained by the 10th band. The robust but penetrating tooth morphology and large jaw circumference suggest that A. kopingensis likely fed upon large prey items.

Modern wild sheep, Ovis, is widespread in the mountain ranges of the Caucasus through Himalaya, Tibetan Plateau, Tianshan-Altai, eastern Siberia, and the Rocky Mountains in North America. In Eurasia, fossil sheep are known at a few Pleistocene sites in North China, eastern Siberia, and western Europe, but are so far absent from the Tibetan Plateau. We describe an extinct sheep, Protovis himalayensis, gen. et sp. nov., from the Pliocene of the Zanda Basin in western Himalaya. Smaller than the living argali, this new form shares with Ovis posterolaterally arched horncores and partially developed sinuses and possesses several transitional characters leading to Ovis. Protovis likely subsisted on C 3 plants, which are the dominant vegetation in the Zanda area during the Pliocene. With the discovery of this new genus and species, we extend the fossil record for the sheep clade into the Pliocene of the Tibetan Plateau, consistent with our previous out-of-Tibet hypothesis. Ancestral sheep in the Pliocene were presumed adapted to high altitude and cold environments, and during the Ice Age, sheep became anatomically modern and dispersed outside of the Tibetan Plateau. Both this new fossil datum and the existing molecular phylogeny suggest that the Tibetan Plateau, possibly including Tianshan-Altai, represents the ancestral home range(s) of mountain sheep and that these basal stocks were the ultimate source of all extant species. Most sheep species survived along their Pleistocene route of dispersal, offering a highly consistent pattern of zoogeography. http://zoobank.org/urn:lsid:zoobank.org:pub:F2AEE746-0A5B-4F40-89B7-8EF0C04F21FD SUPPLEMENTAL DATA-Supplemental materials are available for this article for free at www.tandfonline.com/UJVP Citation for this article: Wang, X., Q. Li, and G. T. Takeuchi. 2016. Out of Tibet: an early sheep from the Pliocene of Tibet, Protovis himalayensis, gen. et sp. nov. (Bovidae, Caprini), and origin of Ice Age mountain sheep. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2016.1169190.

Fossil ceratioid anglerfishes are described from the Upper Miocene (upper Mohnian) deposits of the Puente Formation, Los Angeles Basin, California. The specimens were collected from the laminated turbiditic deposits of the Yorba Member in the eastern sector of the Los Angeles Basin during the construction of a new metro rail line. Five taxa (Borophryne cf. apogon; Chaenophryne aff. melanorhabdus; Leptacanthichthys cf. gracilispinis; Linophryne cf. indica; Oneirodes sp.) belonging to two families, Linophrynidae and Oneirodidae, are described based on nine metamorphosed females. A detailed osteological analysis of the fossils has revealed that they can be tentatively assigned to extant species, suggesting that little or no relevant morphological change has characterized these taxa at least since the Late Miocene. Biogeographic considerations suggest that the Late Miocene ceratioid assemblages of the Los Angeles Basin are strikingly similar to those that currently inhabit the tropical and subtropical eastern Pacific region. From a paleoenvironmental point of view, the excellent preservation of the specimens suggests a reduced turbulence and velocity of the turbidity fluxes. Finally, the comparative study of the bathymetric ranges of the ceratioid taxa recognized in the fossil assemblage described in this paper suggests that the minimum depth of the depositional environment might be estimated at approximately 1,000 m.

We describe an associated skull and lower jaws of a new species of primitive skunk, Martinogale (subfamily Mephitinae), from the late Miocene Dove Spring Formation (late Clarendonian), Kern County, California. It is also the first occurrence of this genus in the Tertiary of the West Coast of North America. The new species is among the best-preserved primitive fossil skunks and represents one of the earliest members of the New World mephitines. As the smallest skunk so far known, the new Martinogale features some of the most primitive cranial and dental morphologies in mephitines. The new information permits a phylogenetic analysis of basal taxa from North America, which suggests a New World clade for all known fossil and living taxa from North and South America. We place this New World clade in a tribe (Mephitini) of its own. New World mephitines share the following derived characters: presence of a P4 parastyle, m1 hypoconid dominant in talonid, and presence of a lingual cingulum on lower canines. We further postulate that New World skunks are the result of a single immigration event, and the new California skunk is close to the origin of the New World skunks.

Modern wild sheep, Ovis, is widespread in the mountain ranges of the Caucasus through Himalaya, Tibetan Plateau, Tianshan-Altai, eastern Siberia, and the Rocky Mountains in North America. In Eurasia, fossil sheep are known at a few Pleistocene sites in North China, eastern Siberia, and western Europe, but are so far absent from the Tibetan Plateau. We describe an extinct sheep, Protovis himalayensis, gen. et sp. nov., from the Pliocene of the Zanda Basin in western Himalaya. Smaller than the living argali, this new form shares with Ovis posterolaterally arched horncores and partially developed sinuses and possesses several transitional characters leading to Ovis. Protovis likely subsisted on C3 plants, which are the dominant vegetation in the Zanda area during the Pliocene. With the discovery of this new genus and species, we extend the fossil record for the sheep clade into the Pliocene of the Tibetan Plateau, consistent with our previous out-of-Tibet hypothesis. Ancestral sheep in the Pliocene were presumed adapted to high altitude and cold environments, and during the Ice Age, sheep became anatomically modern and dispersed outside of the Tibetan Plateau. Both this new fossil datum and the existing molecular phylogeny suggest that the Tibetan Plateau, possibly including Tianshan-Altai, represents the ancestral home range(s) of mountain sheep and that these basal stocks were the ultimate source of all extant species. Most sheep species survived along their Pleistocene route of dispersal, offering a highly consistent pattern of zoogeography.

The 'third pole' of the world is a fitting metaphor for the Himalayan-Tibetan Plateau, in allusion to its vast frozen terrain, rivalling the Arctic and Antarctic, at high altitude but low latitude. Living Tibetan and arctic mammals share adaptations to freezing temperatures such as long and thick winter fur in arctic muskox and Tibetan yak, and for carnivorans, a more predatory niche. Here, we report, to our knowledge, the first evolutionary link between an Early Pliocene (3.60-5.08 Myr ago) fox, Vulpes qiuzhudingi new species, from the Himalaya (Zanda Basin) and Kunlun Mountain (Kunlun Pass Basin) and the modern arctic fox Vulpes lagopus in the polar region. A highly hypercarnivorous dentition of the new fox bears a striking resemblance to that of V. lagopus and substantially predates the previous oldest records of the arctic fox by 3-4 Myr. The low latitude, high-altitude Tibetan Plateau is separated from the nearest modern arctic fox geographical range by at least 2000 km. The apparent connection between an ancestral high-elevation species and its modern polar descendant is consistent with our 'Out-of-Tibet' hypothesis postulating that high-altitude Tibet was a training ground for cold-environment adaptations well before the start of the Ice Age.