A new giraffid (Mammalia, Ruminantia, Pecora) from the late Miocene of Spain, and the evolution of the sivathere-samothere lineage.
ABSTRACT: Giraffids include the only living giraffomorph ruminants and are diagnosed by the presence of bi-lobed canines and a special type of epiphyseal cranial appendages called ossicones. The family Giraffidae ranges from the latest early Miocene until today. However they are currently extant relics with only two living representatives, the African genera Okapia and Giraffa. Giraffids were much more diverse and widespread in the past, with more than 30 fossil species described. For the past decades a number of studies intended to resolve the phylogenetic relationships of the family, but due to the lack of really good cranial material no clear consensus was reached regarding the phylogenetic relationships amongst the different members of the group. The exceptionally complete remains of a new large giraffid from the late Miocene of Spain, Decennatherium rex sp. nov., allows us to improve and reassess giraffid systematics, offering a lot of new data, both anatomic and phylogenetic, on the large late Miocene giraffids of Eurasia. The results of our cladistic analysis show Decennatherium as a basal offshoot of a clade containing the gigantic samotheres and sivatheres, characterized by the presence of a Sivatherium-like ossicone-plan among other features. Decennatherium thus offers the most ancient evidence of this Sivatherium-plan and firmly establishes the early morphological patterns of evolution of a sivathere / samothere-clade that is defined as the less inclusive clade that contains Decennatherium and Sivatherium. Finally, this large group of four-ossiconed giraffids evolutionarily links Miocene Europe and Africa indicating vicariance / migration processes among the giraffid genetic pools separated by the Mediterranean Sea.
Project description:Palaeomerycids were strange three-horned Eurasian Miocene ruminants known through fossils from Spain to China. We here study their systematics, offering the first cladistic phylogeny of the best-known species of the group, and also reassess their phylogenetic position among ruminants, which is currently disputed. The beautifully preserved remains of a new palaeomerycid from middle Miocene deposits of Spain, Xenokeryx amidalae gen. et sp. nov., helps us to better understand palaeomerycid anatomy, especially that of the nuchal region in the skull, significantly improving our current knowledge on these enigmatic ruminants. Our results show two main lineages of palaeomerycids, one containing the genus Ampelomeryx diagnosed by a characteristic type of cranium / cranial appendages and some dental derived traits, and another one that clusters those forms more closely related to Triceromeryx than to Ampelomeryx, characterized by a more derived dentition and a set of apomorphic cranial features. Xenokeryx branches as a basal offshoot of this clade. Also, we find that Eurasian palaeomerycids are not closely related to North American dromomerycids, thus rejecting the currently more accepted view of palaeomerycids as the Eurasian part of the dromomerycid lineage. Instead of this, palaeomerycids are nested with the African Miocene pecoran Propalaeoryx and with giraffoids. On the other hand, dromomerycids are closely related to cervids. We define a clade Giraffomorpha that includes palaeomerycids and giraffids, and propose an emended diagnosis of the Palaeomerycidae based on cranial and postcranial characters, including several features of the cranium not described so far. We also define the Palaeomerycidae as the least inclusive clade of pecorans containing Triceromeryx and Ampelomeryx. Finally, we reassess the taxonomy of several palaeomerycid taxa.
Project description:Several evolutionary theories have been proposed to explain the adaptation of the long giraffe neck; however, few studies examine the fossil cervical vertebrae. We incorporate extinct giraffids, and the okapi and giraffe cervical vertebral specimens in a comprehensive analysis of the anatomy and elongation of the neck. We establish and evaluate 20 character states that relate to general, cranial and caudal vertebral lengthening, and calculate a length-to-width ratio to measure the relative slenderness of the vertebrae. Our sample includes cervical vertebrae (n=71) of 11 taxa representing all seven subfamilies. We also perform a computational comparison of the C3 of Samotherium and Giraffa camelopardalis, which demonstrates that cervical elongation occurs disproportionately along the cranial-caudal vertebral axis. Using the morphological characters and calculated ratios, we propose stages in cervical lengthening, which are supported by the mathematical transformations using fossil and extant specimens. We find that cervical elongation is anisometric and unexpectedly precedes Giraffidae. Within the family, cranial vertebral elongation is the first lengthening stage observed followed by caudal vertebral elongation, which accounts for the extremely long neck of the giraffe.
Project description:Giraffidae are represented by many extinct species. The only two extant taxa possess diametrically contrasting cervical morphology, as the okapi is short-necked and the giraffe is exceptionally long-necked. Samotherium major, known from the Late Miocene of Samos in Greece and other Eurasian localities, is a key extinct giraffid; it possesses cervical vertebrae that are intermediate in the evolutionary elongation of the neck. We describe detailed anatomical features of the cervicals of S. major, and compare these characteristics with the vertebrae of the two extant giraffid taxa. Based on qualitative morphological characters and a quantitative analysis of cervical dimensions, we find that the S. major neck is intermediate between that of the okapi and the giraffe. Specifically, the more cranial (C2-C3) vertebrae of S. major represent a mosaic of features shared either with the giraffe or with the okapi. The more caudal (C5-C7) S. major vertebrae, however, appear transitional between the two extant taxa, and hence are more unique. Notably, the C6 of S. major exhibits a partially excavated ventral lamina that is strong cranially but completely absent on the caudal half of the ventral vertebral body, features between those seen in the giraffe and the okapi. Comprehensive anatomical descriptions and measurements of the almost-complete cervical column reveal that S. major is a truly intermediate-necked giraffid. Reconstructions of the neck display our findings.
Project description:The South Asian river dolphin (Platanista gangetica) is the only extant survivor of the large clade Platanistoidea, having a well-diversified fossil record from the Late Oligocene to the Middle Miocene. Based on a partial skeleton collected from the Chilcatay Formation (Chilcatay Fm; southern coast of Peru), we report here a new squalodelphinid genus and species, Macrosqualodelphis ukupachai. A volcanic ash layer, sampled near the fossil, yielded the 40Ar/39Ar age of 18.78?±?0.08?Ma (Burdigalian, Early Miocene). The phylogenetic analysis places Macrosqualodelphis as the earliest branching squalodelphinid. Combined with several cranial and dental features, the large body size (estimated body length of 3.5?m) of this odontocete suggests that it consumed larger prey than the other members of its family. Together with Huaridelphis raimondii and Notocetus vanbenedeni, both also found in the Chilcatay Fm, this new squalodelphinid further demonstrates the peculiar local diversity of the family along the southeastern Pacific coast, possibly related to their partition into different dietary niches. At a wider geographical scale, the morphological and ecological diversity of squalodelphinids confirms the major role played by platanistoids during the Early Miocene radiation of crown odontocetes.
Project description:Balaenidae (right and bowhead whales) are a key group in understanding baleen whale evolution, because they are the oldest surviving lineage of crown Mysticeti, with a fossil record that dates back ?20 million years. However, this record is mostly Pliocene and younger, with most of the Miocene history of the clade remaining practically unknown. The earliest recognized balaenid is the early Miocene Morenocetus parvus Cabrera, 1926 from Argentina. M. parvus was originally briefly described from two incomplete crania, a mandible and some cervical vertebrae collected from the lower Miocene Gaiman Formation of Patagonia. Since then it has not been revised, thus remaining a frequently cited yet enigmatic fossil cetacean with great potential for shedding light on the early history of crown Mysticeti. Here we provide a detailed morphological description of this taxon and revisit its phylogenetic position. The phylogenetic analysis recovered the middle Miocene Peripolocetus as the earliest diverging balaenid, and Morenocetus as the sister taxon of all other balaenids. The analysis of cranial and periotic morphology of Morenocetus suggest that some of the specialized morphological traits of modern balaenids were acquired by the early Miocene and have remained essentially unchanged up to the present. Throughout balaenid evolution, morphological changes in skull arching and ventral displacement of the orbits appear to be coupled and functionally linked to mitigating a reduction of the field of vision. The body length of Morenocetus and other extinct balaenids was estimated and the evolution of body size in Balaenidae was reconstructed. Optimization of body length on our phylogeny of Balaenidae suggests that the primitive condition was a relatively small body length represented by Morenocetus, and that gigantism has been acquired independently at least twice (in Balaena mysticetus and Eubalaena spp.), with the earliest occurrence of this trait in the late Miocene-early Pliocene as represented by Eubalaena shinshuensis.
Project description:We describe cranial and mandibular remains of three undescribed individuals of the giant mustelid Megalictis ferox Matthew, 1907 from the latest Arikareean (Ar4), Early Miocene mammal fauna of Nebraska, and Wyoming (USA) housed at the American Museum of Natural History (New York, USA). Our phylogenetic hypothesis indicates that Ar4 specimens assigned to M. ferox constitute a monophyletic group. We assign three additional species previously referred to Paroligobunis to Megalictis: M. simplicidens, M. frazieri, and "M." petersoni. The node containing these four species of Megalictis and Oligobunis forms the Oligobuninae. We test the hypothesis that Oligobuninae (Megalictis and Oligobunis) is a stem mustelid taxon. Our results indicate that the Oligobuninae form the sister clade to the crown extant mustelids. Based on the cranium, M. ferox is a jaguar-size mustelid and the largest terrestrial mustelid known to have existed. This new material also sheds light on a new ecomorphological interpretation of M. ferox as a bone-crushing durophage (similar to hyenas), rather than a cat-like hypercarnivore, as had been previously described. The relative large size of M. ferox, together with a stout rostrum and mandible made it one of the more powerful predators of the Early Miocene of the Great Plains of North America.
Project description:Environmental changes can lead to evolutionary shifts in phenotypic traits, which in turn facilitate the exploitation of novel adaptive landscapes and lineage diversification. The global cooling, increased aridity and expansion of open grasslands during the past 50 Myr are prime examples of new adaptive landscapes that spurred lineage and ecomorphological diversity of several mammalian lineages such as rodents and large herbivorous megafauna. However, whether these environmental changes facilitated evolutionary shifts in small- to mid-sized predator morphology is unknown. Here, I used a complete cranial and body morphological dataset to examine the timing of evolutionary shifts in cranial shape, body size and body shape within extant mustelids (martens, otters, polecats and weasels) during the climatic and environmental changes of the Cenozoic. I found that evolutionary shifts in all three traits occurred within extant mustelid subclades just after the onset of the Mid-Miocene Climate Transition. These mustelid subclades first shifted towards more elongate body plans followed by concurrent shifts towards smaller body sizes and more robust crania. I hypothesize that these cranial and body morphological shifts enabled mustelids to exploit novel adaptive zones associated with the climatic and environmental changes of the Mid to Late Miocene, which facilitated significant increases in clade carrying capacity.
Project description:The skull of leporids (rabbits and hares) is highly transformed, typified by pronounced arching of the dorsal skull and ventral flexion of the facial region (i.e., facial tilt). Previous studies show that locomotor behavior influences aspects of cranial shape in leporids, and here we use an extensive 3D geometric morphometrics dataset to further explore what influences leporid cranial diversity. Facial tilt angle, a trait that strongly correlates with locomotor mode, significantly predicts the cranial shape variation captured by the primary axis of cranial shape space, and describes a small proportion (13.2%) of overall cranial shape variation in the clade. However, locomotor mode does not correlate with overall cranial shape variation in the clade, because there are two district morphologies of generalist species, and saltators and cursorial species have similar morphologies. Cranial shape changes due to phyletic size change (evolutionary allometry) also describes a small proportion (12.5%) of cranial shape variation in the clade, but this is largely driven by the smallest living leporid, the pygmy rabbit (Brachylagus idahoensis). By integrating phylogenetic history with our geometric morphometric data, we show that the leporid cranium exhibits weak phylogenetic signal and substantial homoplasy. Though these results make it difficult to reconstruct what the 'ancestral' leporid skull looked like, the fossil records suggest that dorsal arching and facial tilt could have occurred before the origin of the crown group. Lastly, our study highlights the diversity of cranial variation in crown leporids, and highlights a need for additional phylogenetic work that includes stem (fossil) leporids and includes morphological data that captures the transformed morphology of rabbits and hares.
Project description:Monitor lizards (genus Varanus) inhabited Europe at least from the early Miocene to the Pleistocene. Their fossil record is limited to about 40 localities that have provided mostly isolated vertebrae. Due to the poor diagnostic value of these fossils, it was recently claimed that all the European species described prior to the 21st century are not taxonomically valid and a new species, Varanus amnhophilis, was erected on the basis of fragmentary material including cranial elements, from the late Miocene of Samos (Greece). We re-examined the type material of Varanus marathonensis Weithofer, 1888, based on material from the late Miocene of Pikermi (Greece), and concluded that it is a valid, diagnosable species. Previously unpublished Iberian material from the Aragonian (middle Miocene) of Abocador de Can Mata (Vallès-Penedès Basin, Barcelona) and the Vallesian (late Miocene) of Batallones (Madrid Basin) is clearly referable to the same species on a morphological basis, further enabling to provide an emended diagnosis for this species. Varanus amnhophilis appears to be a junior subjective synonym of V. marathonensis. On the basis of the most complete fossil Varanus skeleton ever described, it has been possible to further resolve the internal phylogeny of this genus by cladistically analyzing 80 taxa coded for 495 morphological and 5729 molecular characters. Varanus marathonensis was a large-sized species distributed at relatively low latitudes in both southwestern and southeastern Europe from at least MN7+8 to MN12. Our cladistic analysis nests V. marathonensis into an eastern clade of Varanus instead of the African clade comprising Varanus griseus, to which it had been related in the past. At least two different Varanus lineages were present in Europe during the Neogene, represented by Varanus mokrensis (early Miocene) and V. marathonensis (middle to late Miocene), respectively.
Project description:Killikaike blakei is a new genus and species of anthropoid from the late Early Miocene of southeastern Argentina based on the most pristine fossil platyrrhine skull and dentition known so far. It is part of the New World platyrrhine clade (Family Cebidae; Subfamily Cebinae) including modern squirrel (Saimiri) and capuchin monkeys (Cebus) and their fossil relatives known from Early to Middle Miocene and subrecent periods. Living cebines are relatively large-brained, adroit predatory foragers and live within complex social groups, and wild capuchins exhibit a wide range of behaviors associated with enhanced intelligence. We show that K. blakei lacks diagnostic derived characteristics of the lower face and premolar dentition that are shared by modern cebines, but its strongly vaulted frontal bone and capacious anterior cranial fossa indicate the early evolution of an enlarged forebrain.