The earliest modern mongoose (Carnivora, Herpestidae) from Africa (late Miocene of Chad)

We report on the earliest modern mongooses of Africa, from the late Miocene (ca. 7 Ma) of the hominid locality TM 266, Toros-Menalla, Chad. The material is based on fragmentary dentitions of three individuals. The main diagnostic feature of the Chadian species is the great development of the shear in the carnassials, which distinguishes the Chadian specimens from all extant herpestids except Herpestes and Galerella. In comparison with most extinct and extant Herpestes, the species from Toros-Menalla differs by a markedly smaller size and, depending on the species, relatively more elongated carnassials, more transversely elongated M1 and more reduced p4. On the basis of a great morphological similarity and the absence of significant differences, we assign our material to Galerella sanguinea; the Chadian finding therefore represents the earliest appearance of an extant species of Herpestidae. This record ties the first appearance of the genus to a minimum age of ca. 7 Ma, which is consistent with the estimated divergence date of 11.4 Ma known from the literature for the species of Galerella.     

The earliest record of birch mice from the Early Miocene Nei Mongol,China

The earliest species of birch mouse, Sicista primus sp. nov., was recovered from the 17-Ma-old (Early Miocene) Gashunyinadege locality, central Nei Mongol, China. It is ~9 Ma older than the previous first appearance datum of Sicista in Eurasia. This study indicates that North American Macrognathomys is a synonym of Eurasian Sicista, having 12 shared dental characters. As a result, the biogeography of dipodids indicates that Asian Sicista dispersed to North America as opposed to the hypothesis that Sicista originated from the North American clade. Sicista is one of the few extant rodent genera that originated as early as the Early Miocene.     

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).     

The oldest African fox (Vulpes riffautae n. sp., Canidae, Carnivora) recovered in late Miocene deposits of the Djurab desert, Chad

We report on the oldest fox (Canidae) ever found in Africa. It is dated to 7 Ma based on the degree of evolution of the whole fauna. It belongs to a new species. Its overall size and some morphological characteristics distinguish the Chadian specimen from all the other foxes. The presence of Vulpes and of the genus Eucyon in slightly younger African locality, as well as in southwestern Europe in the late Miocene, may indicate that canids migrated in Europe from Africa through a trans-Mediterranean route.     

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.     

Nectar production dynamics and sugar composition in two <Emphasis Type="Italic">Mucuna</Emphasis> species (Leguminosae,Faboideae) with different specialized pollinators

Nectar is secreted in particular rhythms throughout the lifespan of a flower, which allows determining the nectar production dynamics. This paper compares nectar features in Mucuna japira and Mucuna urens describing: dynamics of nectar production, floral response to nectar removal, resorption, nectar sugar composition, and variation in nectar sugar composition. M. japira inflorescence bears 12–21 yellow flowers, which are in anthesis for 7 days, whereas M. urens inflorescence bears 36–54 greenish flowers, but only 1–3 flowers are in anthesis simultaneously that last one night. Nectar volume and sugar concentration were measured, and the amount of sugar was estimated. Qualitative and quantitative nectar sugar composition was determined. Both species had a constant nectar sugar concentration (ca. 10% for M. japira and ca. 16% for M. urens) and secreted high volumes of nectar (ca. 340 μl per flower for M. japira and 310 μl per flower for M. urens), during 5 days for M. japira and 6 h for M. urens, but after the first removal, i.e., when flower opening mechanism is triggered, nectar production stops immediately. Nectar resorption occurred in both species. Nectar sugar composition showed some similarities between the species. Variation in nectar sugar composition occurred in both species. The Mucuna species are dependent on their pollinators to produce fruits and seeds, and they have different strategies to promote the necessary interaction with birds or bats, especially related to nectar and flower characteristics.     

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     

Paleoenvironmental conditions in the Spanish Miocene–Pliocene boundary: isotopic analyses of Hipparion dental enamel

Expansion of C₄ grasses during Late Miocene and Early Pliocene constitutes one of the most remarkable biotic events of the Cenozoic era. The Teruel–Alfambra region (northeastern Spain) contains one of the most complete Miocene–Pliocene sequences of mammalian fossil sites in the world. In this study, stable isotope (δ ¹³C and δ ¹⁸O) analyses have been performed on the tooth enamel from the equid Hipparion from 19 localities spanning a time interval from approximately 10.9 to 2.7 Ma. This time range starts with the first appearance of this genus in Spain and ends at its extinction. An increase in δ ¹³C at about 4.2 Ma has been observed, indicative of a shift toward a more open habitat. This shift may be related to a large scale vegetation change which occurred across the Miocene–Pliocene boundary when C₄ grasses expanded. This expansion might in turn be linked to global tectonic events such as the uplift of the Himalaya and/or the closure of the Panama Isthmus. However, other more regional factors may have ultimately enhanced the trend toward more open habitats in the Western Mediterranean Basin. The Messinian Salinity Crisis was a major environmental event that may have been responsible for the isotopic changes seen in the equid Hipparion from the Iberian Peninsula along with an increase in the aridity detected ~4.6 Ma ago in the Sahara. Even though the exact factor triggering the isotopic change observed in the Hipparion enamel remains mostly unknown, this study demonstrates that the global environmental changes detected across the Miocene–Pliocene boundary are also recorded in the realm of the Iberian Peninsula. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Harvestmen (Arachnida: Opiliones) from the Middle Jurassic of China

Harvestmen (Arachnida: Opiliones) are familiar animals in most terrestrial habitats but are rare as fossils, with only a handful of species known from each of the Palaeozoic, Mesozoic, and Cenozoic eras. Fossil harvestmen from Middle Jurassic (ca. 165 Ma) strata of Daohugou, Inner Mongolia, China, are described as Mesobunus martensi gen. et sp. nov. and Daohugopilio sheari gen. et sp. nov.; the two genera differ primarily in the relative length of their legs and details of the pedipalps. Jurassic arachnids are extremely rare and these fossils represent the first Jurassic, and only the fourth Mesozoic, record of Opiliones. These remarkably well-preserved and modern-looking fossils are assigned to the Eupnoi, whereby M. martensi demonstrably belongs in Sclerosomatidae. It thus represents the oldest record of a modern harvestman family and implies a high degree of evolutionary stasis among one of the most widespread and abundant groups of long-legged, round-bodied harvestmen.     

Out of Africa: Fossils shed light on the origin of the hoatzin,an iconic Neotropic bird

We describe the earliest fossils of the enigmatic avian taxon Opisthocomiformes (hoatzins) from the Oligo-Miocene (22–24 mya) of Brazil. The bones, a humerus, scapula and coracoid, closely resemble those of the extant hoatzin, Opisthocomus hoazin. The very similar osteology of the pectoral girdle in the new Brazilian fossil compared to the extant O. hoazin, in which it reflects peculiar feeding adaptations, may indicate that hoatzins had already evolved their highly specialized feeding behavior by the mid-Cenozoic. We further show that Namibiavis senutae from the early Miocene of Namibia is another, previously misclassified representative of Opisthocomiformes, which documents that the extant Neotropic distribution of hoatzins is relictual. Because of the weak flight capabilities of hoatzins, their occurrence on both sides of the South Atlantic is of particular biogeographic interest. We detail that this distribution pattern is best explained by dispersal from Africa to South America, and that Opisthocomiformes provide the first example of transatlantic rafting among birds.     

A giant termite from the Late Miocene of Styria, Austria (Isoptera)

A giant termite is described and figured from the Late Miocene of the Styrian Basin in southeastern Austria. Gyatermes styriensis gen. n. et sp. n. is represented by a relatively complete forewing, with basal scale. The fossil approximates in size the largest of all termites today and is the largest fossil termite on record. The presence of this species in the Late Miocene fauna of Europe indicates that climatic conditions were appropriate for the persistence of species and colonies requiring relatively stable, warm conditions. The genus is primitive in overall features but shares some similarity with the dampwood termites.     

Eocene bunoselenodont Artiodactyla from southern Thailand and the early evolution of Ruminantia in South Asia

Although Asia is thought to have played a critical role in the basal radiation of Ruminantia, the fossil record of early selenodont artiodactyls remains poorly documented in this region. Dental remains of a new bunoselenodont artiodactyl are described from the late Eocene of Krabi, southern Thailand. This new form, Krabitherium waileki gen. et sp. nov, is tentatively referred to the Tragulidae (Ruminantia) on the basis of several dental features, including a weak Tragulus fold and the presence of a deep groove on the anterior face of the entoconid. Although this new form is suggestive of the enigmatic ?Gelocus gajensis Pilgrim 1912 from the “base of the Gaj” (lower Chitarwata Formation) of the Bugti Hills (Central Pakistan), K. waileki most likely represents an early representative of a relatively bunodont group of tragulids that includes the genus Dorcabune, known from the Miocene of south Asia. This addition to the Eocene record of early ruminants attests to the antiquity of the group in Southeast Asia and lends support to the hypothesis that the Tragulidae represents one of the first offshoots in the evolutionary history of Ruminantia.     

A gravid lizard from the Cretaceous of China and the early history of squamate viviparity

Although viviparity is most often associated with mammals, roughly one fifth of extant squamate reptiles give birth to live young. Phylogenetic analyses indicate that the trait evolved more than 100 times within Squamata, a frequency greater than that of all other vertebrate clades combined. However, there is debate as to the antiquity of the trait and, until now, the only direct fossil evidence of squamate viviparity was in Late Cretaceous mosasauroids, specialised marine lizards without modern equivalents. Here, we document viviparity in a specimen of a more generalised lizard, Yabeinosaurus, from the Early Cretaceous of China. The gravid female contains more than 15 young at a level of skeletal development corresponding to that of late embryos of living viviparous lizards. This specimen documents the first occurrence of viviparity in a fossil reptile that was largely terrestrial in life, and extends the temporal distribution of the trait in squamates by at least 30 Ma. As Yabeinosaurus occupies a relatively basal position within crown-group squamates, it suggests that the anatomical and physiological preconditions for viviparity arose early within Squamata.     

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.     

Cladistic biogeographic analysis suggests an early Caribbean diversification in Mexico

The Great American Biotic Interchange has been the predominant paradigm for explaining biotic diversification in the Nearctic/Neotropical overlap or Mexican Transition Zone, which is commonly explained by the collision of the North and South American continental plates, which began in the Oligocene and fused both landmasses. In the most far-reaching cladistic biogeographical analysis of the area to date, evidence has been found supporting the existence of a remnant Caribbean region extending from eastern Mexico to southeastern USA, a hypothesis that challenges current views of the Great American Biotic Interchange and the Mexican Transition Zone. We show herein that an older terrane, which has drifted to the present day positions of Yucatan and Cuba, may be biogeographically linked to an early ‘Gondwanan’ biota of the Paleocene (ca. 60 Ma). The evidence indicates an east–west biotic divide in Mexico, existing before the collision and formation of Central America. The south–north division of the country, previously recognized by several authors as associated with the Great American Biotic Interchange and the Mexican Transition Zone, is of a younger age. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Reconstructing the anatomy of the 42-million-year-old fossil †Mengea tertiaria (Insecta, Strepsiptera)

Fossilization in amber is unique in preserving specimens with microscopic fidelity; however, arthropod inclusions are rarely examined beyond the exoskeleton as this requires destructive sampling when traditional techniques are used. We report the first complete, digital 3D, non-destructive reconstruction of the anatomy of an insect fossil, a specimen of †Mengea tertiaria embedded in a 42-Ma Baltic amber. This was made possible using Synchrotron μ-CT. The species belongs to the stem group of the phylogenetically enigmatic and extremely specialized Strepsiptera. Most internal structures of the fossil are preserved, but small parts of the lumen had decayed due to incomplete infiltration of the resin. Data on internal organs provided additional information for resolving phylogenetic relationships. A sister group relationship between †Mengea and all extant lineages of the group was confirmed with characters previously not accessible. The newly gained information also yielded some insights in the biology of †Mengea and the early evolutionary history of Strepsiptera. The technique has a tremendous potential for a more accurate interpretation of diverse fossil arthropods preserved in ambers from 130 Ma to the present.     

Numerical dating of the Eckfeld maar fossil site, Eifel, Germany: a calibration mark for the Eocene time scale

 Sediments of the Eckfeld maar (Eifel, Germany) bear a well-preserved Eocene fauna and flora. Biostratigraphically, Eckfeld corresponds to the Middle Eocene mammal reference level MP (Mammals Paleogene) 13 of the ELMA (European Land Mammal Age) Geiseltalian. In the maar crater, basalt fragments were drilled, representing explosion crater eruption products. By ⁴⁰Ar/³⁹Ar dating of the basalt, for the first time a direct numerical calibration mark for an Eocene European mammal locality has been established. The Eckfeld basalt inverse isochron date of 44.3±0.4 Ma suggests an age for the Geiseltalian/Robiacian boundary at 44 Ma and, together with the 1995 time scale of Berggren et al., a time span ranging from 49 to 44 Ma for the Geiseltalian and from 44 to 37 Ma for the Robiacian, respectively. Additional ⁴⁰Ar/³⁹Ar dating on a genetically related basalt occurrence close to the maar confirms a period of volcanism of ca. 0.6 m.y. in the Eckfeld area, matching the oldest Eocene volcanic activity of the Hocheifel volcanic field. Received: 27 December 1999 / Accepted in revised form: 6 April 2000     

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.