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Publisher Summary This chapter discusses the phenotypic and geographic diversity of Parailurus. The evolutionary history of the lesser panda's predecessors was very unclear until recently due to the fragmentary nature of fossil specimens. It is accepted that Ailuridae, and Ailurinae sensu stricto, include the recent Ailurus fulgens and extinct lesser pandas represented by a sing legenus Parailurus from the Pliocene of Europe, Asia, and North America. Although Amphictis ambiguus from the late Oligocene of Quercy (France) may be more closely related to the Simocyoninae and perhaps Ailurinae, both subfamilies are currently included in the Ailuridae. It is shown that the poor fossil record prevented the recognition of further Miocene ailurids first hand. Although Magerictis imperialis from the middle Miocene of Spain was described as anailurid, it was later assigned to Ailurinae. Only recent findings allowed revealing the ailurid identity of other taxa such as Protursus and Simocyon, all already known to specialists. Despite numerous other localities, the paleontological record of the Parailurus lesser panda remains restricted mostly to the isolated teeth, occasionally with postcranial elements, and rarely cranial fragments. Therefore, it is not surprising that more complete skeletal remains are needed to get an idea of how Parailurus might look.

This chapter deals with the advanced members of Ailuridae. Besides a few well-represented taxa such as Simocyon batalleri and Pristinailurus bristoli, the fossil record of Ailuridae mostly consists isolated teeth and fragmentary specimens. However, these two taxa are beginning to shed new light on this group of specialized carnivorans. In particular, the selection of various dental characters appears to have been mosaic, with different teeth specializing at different rates and with different focal zones. In addition, some forms specialize in hypercarnivory, whereas others have become adapted to hypocarnivory to varying degrees. It is quite likely that Simocyon's move to the trees may have directly led to the selection of features which allowed later Ailurids to exploit flowers and/or fruit (in addition to eating meat), and therefore hypocarnivory may have been a direct result of this change in lifestyle. Such a shift in feeding strategy was clearly advantageous to the group, with fossils covering much of the Northern Hemisphere. Moreover, it appears that although both living ‘pandas’ use their so-called ‘false thumb’ for feeding, each lineage acquired this unique trait via different pathways—one for feeding and one for climbing—making this one of the most fascinating examples of convergent evolution. Lastly, as demonstrated by P. bristoli, not every member of this group remained arboreal, which may have reduced its ability to compete with other more generalized omnivores and ultimately to the extinction of nearly every member of this group.

We analyse the evolution of carnivoran guilds (body mass, locomotor pattern, and diet preference) across the Middle to Upper Miocene boundary based on a comparison of the carnivoran fauna from Steinheim (Mammal Neogene zone MN 7/8; Servallian to early Tortonian), and from the Eppelsheim Formation (Mammal Neogene zone MN 9/10; mid-Tortonian), Germany. Results reveal a massive faunal turnover between these two communties due to taxonomic differences up to the family level. Guild structures of both carnivoran faunas were similar, implying that Late Miocene taxa replaced Middle Miocene taxa in their respective niches rather than new ecological strategies being added to the guild. This is not surprising, as (1) the majority of Late Miocene taxa were derived from European Middle Miocene ancestors and mostly share their paleoecology, (2) of the few undisputed Late Miocene immigrant taxa (Eomellivora, Simocyon, Sivaonyx), only Eomellivora adds new ecological strategies to the guild, and (3) both faunas occupied the same kind of partly open woodland paleoenvironment. Our paper supports previous studies, suggesting that ecological diversity remains stable across the Middle to Upper Miocene boundary and that similar environments produce similar carnivoran guild structures irrespective of taxonomical composition.

International audience; The upper Miocene sites of Hadjidimovo and Kalimantsi in South-Western Bulgaria have yielded a few fossils of the ailurid Simocyon Wagner, 1858, a skull and associated mandible of Promeles Zittel, 1890 (a relatively well-known member of the Mustelidae), and several skulls of Promephitis Gaudry, 1861, an Eurasian member of the Mephitidae, a family represented today by three American and one South-Asian genera, to which were also ascribed Palaeomephitis Jäger, 1839, Miomephitis Dehm, 1950, and Proputorius Filhol, 1890, from the Middle Miocene of Europe, as well as several American Clarendonian to Hemphillian genera. We show that the mephitid status of Palaeomephitis, Miomephitis, and Proputorius is doubtful, and that the late Miocene Promephitis is at a lower grade of encephalization than modern Mephitidae and could be the sister taxon of Pleistocene and modern forms, which can be called Mephitinae. Within this subfamily, the American Pleistocene and modern taxa form the sister group of the Asian Mydaus Cuvier, 1821, which displays numerous autapomorphies. A major, previously unnoticed characteristic of the Mephitinae and Promephitis is that their P4 lingual cusp is a hypocone, unlike that of the Clarendonian American forms, in which it is a protocone. If the latter taxa really belong to the Mephitidae, they must have split early in its history.

Simocyon primigenius (Roth & Wagner, 1854) REMARKS This species has recently been discussed by Spassov & Geraads (2011) who confirmed the identification by Bakalov & Nikolov (1962) of two incomplete mandibles from Kalimantsi. An isolated, unnumbered M1 from Hadjidimovo (Fig. 5B), badly worn and damaged, could be of the same species. It is broader mesially than distally. The central basin is open mesially, and a distinct crest descends lingually from the paracone, but the metacone and protocone are worn out. Its buccal length (16.5 mm) is close to the upper end of the range of S. primigenius, to which we tentatively refer it, pending revision of the early Turolian forms of Central Europe., Published as part of Geraads, Denis & Spassov, Nikolaï, 2016, Musteloid carnivores from the upper Miocene of South- Western Bulgaria, and the phylogeny of the Mephitidae, pp. 543-558 in Geodiversitas 38 (4) on page 554, DOI: 10.5252/g2016n4a5, http://zenodo.org/record/4535593, {"references":["GERAADS D., SPASSOV N., HRISTOVA L., MARKOV G. & TZAN- KOV T. 2011. - Upper Miocene mammals from Strumyani, South-Western Bulgaria. Geodiversitas 33: 451 - 484. https: // doi. org / 10.5252 / g 2011 n 3 a 3"]}

Domingo, M.S., Sanchez, I.M., Alberdi, M.T., Azanza, B. & Morales, J. 2011: Evidence of predation/scavenging on Moschidae (Mammalia, Ruminantia) from the Late Miocene of Spain. Lethaia, Vol. 45, pp. 386–400. Fossil remains belonging to three musk deer species (Moschidae) from the Late Miocene pseudo-karstic fossil site of Batallones-1 (Madrid, Spain) bear evidence of corrosion produced by digestive acids. Although moschid bones co-occur with remains of several other vertebrate taxa, modifications due to digestion are restricted to the fossils of these ruminants. In this study, we focus on the particular taphonomic history undergone by the moschid assemblage. A description of the bone-surface modifications together with an analysis of the skeletal part representation and bone breakage are provided. The absence in the fossil site of many moschid skeletal elements and fragments of long bones indicates that these ruminants were preyed on or consumed out of the cavity and eventually became part of the bone assemblage when their predator/scavenger became trapped. The features exhibited by the moschid fossiliferous assemblage allow the evaluation of the trophic interactions in this Late Miocene community. The ailurid Simocyon batalleri and the amphicyonid Magericyon anceps are potentially responsible for the moschid assemblage considering the taphonomical features exhibited by the musk deer remains (type and degree of digestive damage and bone breakage, size of the recovered remains) and the dietary adaptations and preferences of these carnivorans. Together, digestive corrosion traces and moschid bone assemblage characteristics provide strong evidence for specific predator-prey interactions in a Miocene ecosystem. Batallones-1, digestion, Miocene, Moschidae, taphonomy.

” from North Ameri-can Hemphilian of Oregon, Utah, and possibly Nevada (Tedrow etal., 1999). Its migration from Eurasia to North America probablyoccurred at about 8 Ma (Tedrow et al., 1999, Qiu, 2003). Thegeologically oldest record of the genus comes from Asia. Twosmall specimens (IVPP V7732, IVPP V11505) from the latemiddle Miocene of the Chinese Junggar Basin were assigned as

Inferences of function and ecology in extinct taxa have long been a subject of interest because it is fundamental to understand the evolutionary history of species. In this study, we use a quantitative approach to investigate the locomotor behaviour of Simocyon batalleri, a key taxon related to the ailurid family. To do so, we use 3D surface geometric morphometric approaches on the three long bones of the forelimb of an extant reference sample. Next, we test the locomotor strategy of S. batalleri using a leave-one-out cross-validated linear discriminant analysis. Our results show that S. batalleri is included in the morphospace of the living species of musteloids. However, each bone of the forelimb appears to show a different functional signal suggesting that inferring the lifestyle or locomotor behaviour of fossils can be difficult and dependent on the bone investigated. This highlights the importance of studying, where possible, a maximum of skeletal elements to be able to make robust inferences on the lifestyle of extinct species. Finally, our results suggest that S. batalleri may be more arboreal than previously suggested., MJS and JM belong to the Research Group UCM-BSCH 910607.

Some relatively rare carnivore remains from Pikermi are studied. The present specimens include Simocyon primigenius (ROTH and WAGNER, 1854) , Promeles palaeatticus (WEITHOFER, 1888) , Martes woodwardi PILGRIM, 1931 , Felis attica WAGNER, 1857 and Machairodus giganteus (WAGNER, 1848) , as well as some postcranial remains that resemble Enhydriodon? latipes PILGRIM, 1931 and are assigned to Mustelidae gen. et spec. indet. Some of these species were not known until now but from their holotypes as M. woodwardi and E.? latipes. Other species such as F. attica and M. giganteus are better known but from relatively scant material. Since the material of the earlier mentioned species comes from Pikermi, their type locality, it is of especial importance and improves our knowledge for the Pikermi carnivores and their relationships to other species. In some cases where the material comes from ossiferous blocks, the accompanying faunal context is discussed.

A well-preserved skull of Simocyon, an extinct hypercarnivorous procyonid, is described from a late Miocene (Baodean; approximately 5.3–9 Ma) locality in north Shaanxi, China. New knowledge about its cranial morphology allows insight into its phylogenetic position among musteloid camivorans. A sister-group relationship between the simocyonines and the ailurines (the East Asian red panda) is proposed on the basis of cranial and dental evidence. Shared derived characters in support of this relationship include: highly arched zygomatic arch; posteriorly extended posterior palatine border; long bony external auditory meatus; a posterolateral process of promontorium; ventrally ridged paroccipital process; anteriorly inclined coronoid crest; and lateral grooves on canines. The simocyonine-ailurine clade is in turn postulated to be within the procyonid clade because of its common possession of an enlarged M2 and an elongated talonid on m2. The new fossil evidence contradicts several recent phylogenetic ...

The red panda family, the Ailuridae, is unusual because its only extant representative is a derived form, far removed from the group’s morphotypical condition. Effectively, the extreme adaptations of the red panda skull and dentition for a vegetarian diet imply that the study of the fossil record is essential if we are to gain an understanding of the original biology and ecology of the ailurids. Fossil ailurids have remained frustratingly rare and fragmentary findings for many decades, but recent years have seen a dramatic improvement of their record. This has been especially the case with the recent finds of dramatically complete remains of the simocyonine Simocyon batalleri from Batallones-1 in Madrid, Spain, and of the ailurine Pristinailurus bristoli from Gray Fossil Site in Tennessee, United States. Taking into account the information provided by these and other recent findings, it is now possible to gain a more complete picture than ever before of the true place of ailurids within the Carnivora, and of the evolutionary processes that led to the remarkable convergences between the only living ailurid and the unrelated giant panda. In this chapter, we present the early evolution of the ailurids, its Old-World fossil record, and the implications of the palaeontological evidence for the position of the ailurids among other carnivoran families.

Publisher Summary This chapter deals with the advanced members of Ailuridae. A side from a few well-represented taxa such as Simocyon batalleri and Pristinailurus bristoli, the fossil record of the Ailuridae consists of mostly isolated teeth and fragmentary specimens. However, these two taxa are beginning to shed new light on this group of specialized carnivorans. The selection of the various dental characters appears to have been mosaic, with different teeth specializing at different rates (and with different focal zones). In addition, some forms specialized in hypercarnivory, while others were becoming adapted to hypocarnivory to varying degrees. It is quite likely that Simocyon's move to the trees may have directly led to the selection of features which allowed this group to exploit flowers and/or fruit (in addition to eating meat), and therefore hypocarnivory may have been a direct result of this change in lifestyle. Moreover, it appears that although both living “pandas” use their so-called “falsethumb” for feeding, each lineage acquired this unique trait via different pathways—one for feeding and one for climbing—making this one of the most fascinating examples of convergent evolution. Lastly, as demonstrated by Pristinailurus bristoli, not every member of this group remained arboreal, which may have reduced its ability to compete with other more generalized omnivores, and ultimately to the extinction of nearly every member of this group.

A phylogenetic analysis of the Oligocene and some Neogene European arctoid carnivorans of the order-group taxon Mustelida is performed, based on characters of skull and dentition.

The red panda, Ailurus fulgens, is a peculiar recent carnivoran whose systematic relationships have been disputed since its first description in 1825 (Figure 4.1). While occurring with a single species today, Ailuridae was represented by several genera in the past. Flynn and Nedbal (1998) were the first to place Ailurus in its own family based on molecular evidence. While subsequent analyses of molecular data have confirmed this, paleontological findings of the last 10 years shed much more light on the natural history of Ailuridae and the morphology of its members. Before 1997, Ailuridae consisted of 5 genera with about 16 species. Only two genera belonged to Ailurinae (Ailurus and Parailurus). Today, the family contains 9 genera (5 of which are considered to be ailurines) with about 26 species. It now is agreed that the earliest ailurid is Amphictis, the earliest species of which are known from the Late Oligocene (MP 28, about 25 Mya) of Europe. Even though Ailurus today is restricted to Southeast Asia, it is apparent that ailurids were once present in all Northern continents and were most diverse in Europe throughout most of their history. In this chapter, we examine the recent and fossil evidence supporting the assignment of Ailurus fulgens and its ancestors (including Amphictis) to a distinct family, Ailuridae. We find such an assignment fully justified and corroborated by morphological data. We review all molecular and morphological studies in which Ailurus has been included. While new molecular studies continue to support a family Ailuridae, a review of the morphology of A. fulgens led to the identification of a number of characters which could be traced within the fossil relatives of Ailurus. This survey led to the development of emended diagnoses of all included genera. We also include a list of all ailurid species and resolve some outstanding issues concerning the taxonomy or systematic relationships of several taxa. Finally, we briefly recapitulate the natural history of

We describe the postcranial anatomy of the Miocene puma-sized ailurid Simocyon batalleri, discussing some aspects of its biomechanics and inferring a probable life style. The postcranial anatomy of S. batalleri was previously unknown due the fragmentary nature of its fossil record, and most of the available fossils corresponded to cranial and dental material. With the discovery of a rich sample from the Late Miocene natural trap of Batallones-1, to the south of Madrid (Spain), including the remains of at least two individuals of S. batalleri, it is possible to study for the first time the anatomy of the cervical and lumbar vertebrae, complete forelimb and partial hind limb. The morphology of the forelimb, the lumbar region and the unexpected presence of a false-thumb are strongly suggestive of arboreal capabilities. Such locomotor abilities are consistent with a palaeobiological model of a generalized carnivore that foraged mainly on the ground but could readily climb to trees for safety if faced with the threat of larger competing carnivores. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 593–621.

The “false thumb” of pandas is a carpal bone, the radial sesamoid, which has been enlarged and functions as an opposable thumb. If the giant panda ( Ailuropoda melanoleuca ) and the red panda ( Ailurus fulgens ) are not closely related, their sharing of this adaptation implies a remarkable convergence. The discovery of previously unknown postcranial remains of a Miocene red panda relative, Simocyon batalleri , from the Spanish site of Batallones-1 (Madrid), now shows that this animal had a false thumb. The radial sesamoid of S. batalleri shows similarities with that of the red panda, which supports a sister-group relationship and indicates independent evolution in both pandas. The fossils from Batallones-1 reveal S. batalleri as a puma-sized, semiarboreal carnivore with a moderately hypercarnivore diet. These data suggest that the false thumbs of S. batalleri and Ailurus fulgens were probably inherited from a primitive member of the red panda family (Ailuridae), which lacked the red panda's specializations for herbivory but shared its arboreal adaptations. Thus, it seems that, whereas the false thumb of the giant panda probably evolved for manipulating bamboo, the false thumbs of the red panda and of S. batalleri more likely evolved as an aid for arboreal locomotion, with the red panda secondarily developing its ability for item manipulation and thus producing one of the most dramatic cases of convergence among vertebrates.