Journal of Mammalian Evolution

Functional and systematic implications of the postcranial anatomy of a Late Miocene feline (Carnivora, Felidae) from Batallones-1 (Madrid, Spain) --Manuscript Draft--

Manuscript Number: JOMM-D-17-00026R2 Full Title: Functional and systematic implications of the postcranial anatomy of a Late Miocene feline (Carnivora, Felidae) from Batallones-1 (Madrid, Spain) Article Type: Original Article Keywords: Morphology; locomotion; Vallesian; Leptofelis Corresponding Author: Manuel J Salesa Museo Nacional de Ciencias Naturales-CSIC Madrid, Madrid SPAIN Corresponding Author Secondary Information: Corresponding Author's Institution: Museo Nacional de Ciencias Naturales-CSIC Corresponding Author's Secondary Institution: First Author: Manuel J Salesa First Author Secondary Information: Order of Authors: Manuel J Salesa Gema Siliceo Mauricio Antón Stéphane Peigné Jorge Morales Order of Authors Secondary Information:

Funding Information: Ministerio de Economía y Competitividad Dr Jorge Morales (ES) (CGL2015-68333-P (MINECO/FEDER, UE)) Fundação para a Ciência e a Tecnologia Dr. Manuel J Salesa (IF/00351/2014/CP1216/CT0003)

Abstract: The Spanish late Miocene locality of Batallones-1 yielded a rich sample of large carnivorans, including saber-toothed felids, amphicyonids, and ailurids, but also of smaller species, with the small cats being especially interesting. Two species are known from Batallones-1, one of them the size of a wildcat, Felis silvestris, the other one the size of a caracal, Caracal caracal. The former is represented by skulls, mandibles, and postcranial bones, whereas the latter is known from a collection of long bones. Both species are less abundant than their larger relatives, the saber-toothed felids Promegantereon ogygia and Machairodus aphanistus, but the available sample allows us to assess body proportions and adaptations of the smallest species, and to propose a new genus for this feline, Leptofelis vallesiensis. Its limb bones are remarkably gracile compared to fossils of the earlier genera Pseudaelurus, Miopanthera, and Styriofelis, and comparable in cursorial adaptations to the wildcat, very different from extant arboreal cats. While middle Miocene felids were likely semi- arboreal forest dwellers, L. vallesiensis would be mostly terrestrial, climbing essentially for protection. This indicates an adaptation to a mosaic of habitats, including relatively open terrain, and may be related to the climatic changes detected in Eurasia during the late Miocene.

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Click here to view linked References 1 2 3 4 Manuel J. Salesa, Gema Siliceo, Mauricio Antón, Stéphane Peigné, Jorge Morales 5 6 7 8 9 10 Functional and systematic implications of the postcranial anatomy of a Late Miocene 11 12 feline (Carnivora, Felidae) from Batallones-1 (Madrid, Spain) 13 14 15 16 17 Manuel J. Salesa (corresponding author) 18 19 Departamento de Paleobiología, Museo Nacional de Ciencias Naturales-CSIC, 20 21 22 C/ José Gutiérrez Abascal, 2. 28006 Madrid, Spain; 23 24 Departamento de Geologia, Faculdade de Ciências da Universidade de Lisboa, Campo 25 26 27 Grande, Edifício C6 - 1749-016 Lisboa, Portugal; 28 29 30 31 32 Gema Siliceo 33 34 Departamento de Ciencias de la Vida, Universidad de Alcalá, Edificio de Ciencias, Campus 35 36 Universitario, 28805 Alcalá de Henares, Spain; 37 38 39 40 41 Mauricio Antón 42 43 44 Departamento de Paleobiología, Museo Nacional de Ciencias Naturales-CSIC, 45 46 C/ José Gutiérrez Abascal, 2. 28006 Madrid, Spain; 47 48 49 50 51 Stéphane Peigné 52 53 CR2P–UMR 7207 CNRS, Museum national d’Histoire naturelle, Université Paris 06, 57 54 55 56 rue Cuvier, CP 38, 75005 Paris, France; 57 58 59 60 61 62 63 1 64 65 1 2 3 4 Jorge Morales 5 6 7 Departamento de Paleobiología, Museo Nacional de Ciencias Naturales-CSIC, 8 9 C/ José Gutiérrez Abascal, 2. 28006 Madrid, Spain 10 11 12 13 14 E-mail: [email protected]; telephone number: +34 915668979; fax number: +34 15 16 915668960 17 18 19 20 21 22 Acknowledgements This study is part of the research projects CGL2015-68333-P 23 24 (MINECO/FEDER, UE) and IF/00351/2014/CP1216/CT0003 (Fundação para a Ciência e a 25 26 27 Tecnologia, Portugal). MJS is member of the Research Groups CSIC 641538 and IDL-RG2 28 29 (Coast, Water and Earth surface processes). GS is member of the Research Group 30 31 32 “Bioacústica Evolutiva y Paleoantropología”, and a postdoctoral researcher funded by the 33 34 Postdoctoral Programme from the Universidad de Alcalá (Madrid, Spain). JM is member of 35 36 the Research Groups CSIC 641538 and CAM-UCM 910607. We thank the Comunidad 37 38 39 Autónoma de Madrid (Dirección General de Patrimonio Histórico) for its continuous 40 41 funding support and research permissions. We especially thank Dr. Juan Francisco Pastor 42 43 44 (Universidad de Valladolid, Spain) for the loan of the extant specimens used for 45 46 comparison. We thank Enrique Cantero for his excellent preparation of the fossils of 47 48 49 Leptofelis vallesiensis from Batallones-1. We also thank Dr. Gertrud Rößner (Curator of 50 51 Fossil Mammals) and Manuela Schellenberger (photographer) both from the Bayerische 52 53 Staatssammlung für Paläontologie und Geologie (Munich, Germany), for providing us with 54 55 56 images of the tibia of Styriofelis turnauensis from Wintershof-West (catalogue number 57 58 59 60 61 62 63 2 64 65 1 2 3 4 SNSB-BSPG 1937 II 12806). Finally, we thank Dr. John Wible, and two anonymous 5 6 7 reviewers for their helpful and constructive comments on an earlier version of this paper. 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 3 64 65 1 2 3 4 Abstract The Spanish late Miocene locality of Batallones-1 yielded a rich sample of large 5 6 7 carnivorans, including saber-toothed felids, amphicyonids, and ailurids, but also of smaller 8 9 species, with the small cats being especially interesting. Two species are known from 10 11 Batallones-1, one of them the size of a wildcat, Felis silvestris, the other one the size of a 12 13 14 caracal, Caracal caracal. The former is represented by skulls, mandibles, and postcranial 15 16 bones, whereas the latter is known from a collection of long bones. Both species are less 17 18 19 abundant than their larger relatives, the saber-toothed felids Promegantereon ogygia and 20 21 Machairodus aphanistus, but the available sample allows us to assess body proportions and 22 23 24 adaptations of the smallest species, and to propose a new genus for this feline, Leptofelis 25 26 vallesiensis. Its limb bones are remarkably gracile compared to fossils of the earlier genera 27 28 29 Pseudaelurus, Miopanthera, and Styriofelis, and comparable in cursorial adaptations to the 30 31 wildcat, very different from extant arboreal cats. While middle Miocene felids were likely 32 33 semi-arboreal forest dwellers, L. vallesiensis would be mostly terrestrial, climbing essentially 34 35 36 for protection. This indicates an adaptation to a mosaic of habitats, including relatively open 37 38 terrain, and may be related to the climatic changes detected in Eurasia during the late 39 40 41 Miocene. 42 43 44 45 46 Keywords Morphology; Locomotion; Vallesian; Leptofelis. 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 4 64 65 1 2 3 4 Introduction 5 6 7 8 9 Fossils of primitive felids are recorded in several early and middle Miocene sites of Europe, 10 11 Africa, and North America (Thenius 1949; Viret 1951; Crusafont-Pairó 1952; de Beaumont 12 13 14 1961; Ginsburg 1961a, 1983, 1999, 2002; Crusafont-Pairó and Ginsburg 1973; Rothwell 15 16 2001, 2003; Morales et al. 2003; Werdelin et al. 2010). Concerning those showing feline 17 18 19 affinities, there are up to four species classically included in the genus Pseudaelurus 20 21 Gervais, 1850: Ps. romieviensis Roman and Viret, 1934, from La Romieu (France, MN4), 22 23 24 Ps. turnauensis (Hoernes, 1882) from Göriach (Germany, MN 5), Ps. transitorius Depéret, 25 26 1892 and Ps. lorteti Gaillard, 1899, both from La Grive-Saint-Alban (France, MN 7/8). All 27 28 29 of them are mostly known from cranial and dental fragments, which show a quite similar 30 31 pattern, but unfortunately, very few postcranial bones are known from these early felines, 32 33 and thus it is difficult to make inferences about their locomotor adaptations. A fifth species 34 35 36 of primitive felid from Sansan (France, MN 6) was also included in Pseudaelurus as Ps. 37 38 quadridentatus (Blainville, 1843) (in fact this form is the type species of the genus), but 39 40 41 based on its particular features (moderately elongated and laterally flattened upper canines 42 43 and relatively robust Mc I) it is currently considered as the first known saber-toothed felid, 44 45 46 hence the genus Pseudaelurus should be included within the subfamily Machairodontinae 47 48 (de Beaumont 1978; Werdelin et al. 2010; Salesa et al. 2012a; Peigné 2012). Considering 49 50 this, and the feline dental characters (basically, relatively short upper canines with rounded 51 52 53 section) that they displayed, most of the other species of Pseudaelurus (i.e., Ps. lorteti, Ps. 54 55 turnauensis, and Ps. transitorius, the last being a junior synonym of Ps. turnauensis) 56 57 58 mentioned above were assigned in different genera, Styriofelis Kretzoi, 1929, for Ps. 59 60 turnauensis, and Miopanthera Kretzoi, 1938, for Ps. lorteti (Geraads and Peigné 2016). To 61 62 63 5 64 65 1 2 3 4 summarize, in the middle Miocene at least three different taxa are known in Western 5 6 7 Europe: the primitive machairodontine Ps. quadridentatus, the lynx-sized feline M. lorteti, 8 9 and the wild cat-sized feline S. turnauensis. Nonetheless, we should consider that the 10 11 scarcity of postcranial bones of these taxa is probably concealing a greater generic diversity 12 13 14 and preventing an in-depth taxonomic knowledge of these early Felinae. 15 16 Cranial remains of European late Miocene Felinae are much better known than 17 18 19 those from the lower and middle Miocene, and several skulls and mandibles are known 20 21 from different Turolian European localities (Wagner 1857; Schlosser 1924; de Mecquenem 22 23 24 1924; Riabinin 1927; Kretzoi 1951, 1952; de Beaumont 1961; Schmidt-Kittler 1976; 25 26 Solounias 1981; Forsten and Kaya 1995; Morlo 1997; Ginsburg 1999; Roussiakis 2002; de 27 28 29 Bonis 2005). All of them were classically assigned to the wildcat-sized Felis attica 30 31 Wagner, 1857, but separated by Salesa et al. (2012a) into the new genus Pristifelis as P. 32 33 attica. Nevertheless, the postcranial skeleton of this animal is as poorly known as those 34 35 36 from the earliest Felinae. In fact, few papers including comparative descriptions or 37 38 functional inferences on the postcranial anatomy of these Miocene felines have been 39 40 41 published (de Beaumont 1961, 1986; Salesa et al. 2011). Finally, a new species was 42 43 described from the Vallesian of Batallones-1 and Batallones-3 (Madrid, Spain) by Salesa et 44 45 46 al. (2012a) under the name of Styriofelis vallesiensis. This is the smallest of the two species 47 48 of Felinae known from these Spanish localies. When Salesa et al. (2012a) assigned the 49 50 small feline from Batallones-1 and Batallones-3 to the genus Styriofelis as the new species 51 52 53 S. vallesiensis, they chose the most conservative taxonomic point of view based on a set of 54 55 dental similarities between the Batallones sample and the Middle Miocene feline S. 56 57 58 turnauensis, although in their cladogram these two species are not sister-taxa. 59 60 61 62 63 6 64 65 1 2 3 4 Here, we describe the postcranial anatomy of S. vallesiensis, based on a well- 5 6 7 documented sample from Batallones. The objective of our study is first to address the 8 9 generic affinities of the two species of Styriofelis: do their postcrania provide an additional 10 11 support of their close generic relationships? At a wider scale, we also want to address the 12 13 14 systematic relationships and postcranial functional anatomy of the small- to medium-sized 15 16 felines of the middle and late Miocene of Europe mentioned above: Miopanthera lorteti, 17 18 19 Styriofelis turnauensis, Styriofelis vallesiensis, and Pristifelis attica. We posit that their 20 21 postcranial anatomy presents marked differences that, in addition to their particular cranio- 22 23 24 dental features, supports both their taxonomic distinction and different adaptations and 25 26 ecology. Thus, in the Systematic Palaeontology section we provide with a proper 27 28 29 discussion on this issue, and the proposal of the new genus Leptofelis for the species S. 30 31 vallesiensis. In consequence, we will refer throughout the text to the species from 32 33 Batallones-1 with the new name of Leptofelis vallesiensis. 34 35 36 37 38 Material and methods 39 40 41 42 The studied postcranial fossils of Leptofelis vallesiensis come from the fossil site of 43 44 Batallones-1, which is part of the Cerro de los Batallones paleontological complex, located 45 46 47 in a low hill of around 700 m of elevation, 30 km south of the city of Madrid (Spain). Up to 48 49 nine localities have been found since 1991 thanks to the exploitation of the hill as an 50 51 52 opencast mine of sepiolite. The sites were formed through a geological process of piping, 53 54 which consisted in the erosion of the sepiolite levels by water flowing along fractures, 55 56 57 causing collapses and the development of karst-like (‘pseudokarst’) topography (Pozo et al. 58 59 2004; Calvo et al. 2013). These cavities acted as natural traps for the animals inhabiting the 60 61 62 63 7 64 65 1 2 3 4 area, mostly carnivorous vertebrates that entered the holes attracted by the carcasses of 5 6 7 other previously trapped animals (Antón and Morales 2000; Morales et al. 2000). One of 8 9 them, Batallones-1, excavated during 1991-2008, has provided an amazing assemblage of 10 11 Carnivorans, including the ailurid Simocyon batalleri, the amphicyonid Magericyon 12 13 14 anceps, three species of mustelids, the primitive hyaenid Protictitherium crassum, the 15 16 machairodont felids Machairodus aphanistus and Promegantereon ogygia, and two species 17 18 19 of felines (Morales et al. 2000, 2004; Antón et al. 2004; Peigné et al. 2005, 2008; Salesa et 20 21 al. 2005, 2006a, b, 2008, 2010a, b; Siliceo et al. 2015, 2017). In their revision of the late 22 23 24 Miocene felines, Salesa et al. (2012a) proposed the new species Styriofelis vallesiensis for 25 26 the smaller of these two species of Felinae, mainly based on the relative dental similarities 27 28 29 of this species with the Middle Miocene S. turnauensis. Leptofelis vallesiensis is present in 30 31 two localities of Batallones: Batallones-1 and Batallones-3, separated by no more than one 32 33 hundred meters, and with their saber-toothed felids populations showing enough 34 35 36 morphological differences to support also a temporal separation between them (Monescillo 37 38 et al. 2014; Siliceo et al. 2014). 39 40 41 The fossils of L. vallesiensis described here are housed at the Museo Nacional de 42 43 Ciencias Naturales-CSIC (Madrid, Spain). They were found during the excavation 44 45 46 campaigns of 1991-2008 at the late Vallesian locality of Batallones-1 (Torrejón de Velasco, 47 48 Spain), coordinated by J. Morales. The list of studied material is as follows: BAT-1’06 D4- 49 50 64a, BAT-1’06 D4-64b and BAT-1’06 D4-64c, L7-L4; B-4968, right humerus; BAT-1’02 51 52 53 D6-58, left humerus; B-2074 (5), right radius; B/S-575, left radius; B-2074 (6), BAT-1’08 54 55 C8-24, and BAT-1’06 E5-50, right ulnae; BAT-1’03 D4-239, right Mc I; B-2849, left Mc I; 56 57 58 BAT-1’03 D5-162, right Mc II; BAT-1’04 D4-184, right Mc III; BAT-1’03 E4-164, right 59 60 Mc IV; BAT-1’04 E5-129, right Mc V; BAT-1’04 E5-212, left Mc V; BAT-1’06 D4-66a, 61 62 63 8 64 65 1 2 3 4 left and right coxae; BAT-1’06 D4-66b, sacrum; BAT-1’06 D4-67a, right femur; BAT-1’06 5 6 7 D4-67b, left femur; BAT-1’06 D4-103, right tibia; BAT-1’06 D4-110, left tibia; BAT-1’04 8 9 D6-173, right talus; BAT-1’03 D6-300, left talus; BAT-1’04 D6-173 and BAT-1’06 F5-61, 10 11 right calcanei; BAT-1’04 D6-62 and BAT-1’03 D6-301, left calcanei. 12 13 14 Nine species of small extant Felidae were used for anatomical comparisons: 15 16 Caracal caracal (Schreber, 1776), Felis silvestris Schreber, 1775, Felis lybica Forster, 17 18 19 1780, Leopardus wiedii (Schinz, 1821), Leopardus geoffroyi (d’Orbigny and Gervais, 20 21 1844), Leptailurus serval (Schreber, 1776), Lynx pardinus (Temminck, 1827), Lynx lynx 22 23 24 (Linnaeus, 1758), and Lynx rufus (Schreber, 1777). We also used specimens from other 25 26 families of Carnivora: the viverrids Genetta genetta Linnaeus, 1758, and Civettictis civetta 27 28 29 (Schreber, 1776), and the mustelid Martes martes (Linnaeus, 1758). All this material 30 31 belongs to the collections of the Museo Anatómico, Facultad de Medicina, Universidad de 32 33 Valladolid (Spain), and the Comparative Anatomy Collections of the Museo Nacional de 34 35 36 Ciencias Naturales-CSIC. We also compared the fossils of Leptofelis vallesiensis from 37 38 Batallones-1 with the sample of Miopanthera lorteti (Blainville, 1842) from the classical 39 40 41 fossil site of Sansan (MN 6, France), belonging to the collections of the Muséum national 42 43 d’Histoire naturelle (Paris, France); comparisons were also made with published material of 44 45 46 Styriofelis turnauensis (Gaillard 1899; de Beaumont 1961) and M. lorteti (Ginsburg and 47 48 Antunes 1995; Salesa et al. 2011) and the late Miocene feline Pristifelis attica (Roussiakis 49 50 2002). The anatomical descriptions follow the terminology used by Barone (2010), Evans 51 52 53 (1993), Julik et al. (2012) and the Nomina Anatomica Veterinaria (2012). The 54 55 measurements were taken with digital calipers, and are shown in Fig. 1 and Tables 1–2. 56 57 58 59 60 Abbreviations 61 62 63 9 64 65 1 2 3 4 Institutional abbreviations: AMPG PA, Athens Museum of Palaeontology and Geology, 5 6 7 National and Kapodistrian University of Athens; MAN, Museo Anatómico de la 8 9 Universidad de Valladolid; MNCN, Museo Nacional de Ciencias Naturales-CSIC; L. Gr. 10 11 and LGR, La Grive-Saint-Alban, Muséum d’Histoire Naturelle de Lyon; SNSB BSPG, 12 13 14 Staatliche Naturwissenschaftliche Sammlungen Bayerns - Bayerischen Staatssammlung für 15 16 Paläontologie und Geologie. 17 18 19 20 21 Anatomical and measurement abbreviations: ah, articular proximodistal height; aw, 22 23 24 articular mediolateral width; dw, distal mediolateral width; Mc, metacarpal; Mt, metatarsal; 25 26 oh, olecranon proximodistal height; pl, proximal craniocaudal length; pw, proximal 27 28 29 mediolateral width; tl, total proximodistal length. 30 31 32 33 Use of cladistic terminology 34 35 36 Unfortunately, and due to the scarcity of postcranial fossils of Miocene felines, no cladistic 37 38 analysis could be performed in the present study. Nevertheless, the discussed characters are 39 40 41 polarized by outgroup comparisons to non-felid Feliformia or Caniformia on one end, and 42 43 crown-group felids on the other. Given this, and in absence of a proper cladistic analysis, 44 45 46 we cannot rule out the possibility that apparently plesiomorphic features are actually 47 48 reversals, and that features that appear to be derived are convergences with crown taxa 49 50 (instead of shared derived characters). Only a future cladistic analysis could elucidate these 51 52 53 issues, but we have considered of interest to use the terms “primitive” and “derived” in the 54 55 context of our discussion on functional adaptations. 56 57 58 59 60 61 62 63 10 64 65 1 2 3 4 Data availability Statement 5 6 7 All data generated or analyzed during this study are included in this published article. 8 9 10 11 12 Anatomical descriptions and comparisons 13 14 15 In general, the postcranial bones of L. vallesiensis from Batallones-1 show a good state of 16 17 18 preservation, and practically, all their anatomical structures can be observed. In this section, 19 20 we describe the most remarkable features of this material. 21 22 23 24 25 Lumbar vertebrae 26 27 28 29 30 Four articulated lumbar vertebrae of L. vallesiensis, the cranial one just preserving its 31 32 caudal half, are known from Batallones-1 (Fig. 2). The most caudal of these vertebrae has a 33 34 35 craniocaudally shorter body and wider caudal articulation processes than the others, which 36 37 identifies it as the seventh lumbar vertebra, and so the rest are the sixth, the fifth, and a 38 39 40 caudal fragment of the fourth lumbar vertebrae (formally, L7-L4). Comparing this lumbar 41 42 series with those of the comparative sample of extant felines, the L7 of L. vallesiensis is 43 44 relatively shorter than those of the other species, especially those of F. silvestris, F. lybica 45 46 47 and Lp. geoffroyi, whereas the L7 of Le. serval, C. caracal, Ly. lynx, Ly. pardinus, and Ly. 48 49 rufus are only slightly craniocaudally longer than that of L. vallesiensis. The costal 50 51 52 processes are severely broken in all the specimens, but the preserved portions show that 53 54 they were flat and cranially curved. The spinous processes were developed from the cranial 55 56 57 to the caudal border of the dorsal surface, and at least in the sixth and fifth vertebrae were 58 59 triangular (in the seventh lumbar just the base is preserved), although their tips are broken 60 61 62 63 11 64 65 1 2 3 4 (Fig. 2a). Both the cranial and caudal articular processes are elliptical and dorsocranially or 5 6 7 dorsocaudally oriented, respectively; they show this same morphology and orientation in 8 9 the rest of the compared species. In the ventral surface of the body, a shallow central ridge 10 11 is observed in all the compared felines (Fig. 2b). 12 13 14 15 16 Humerus 17 18 19 20 21 The overall morphology of the humerus of L. vallesiensis is similar to that of other small- 22 23 24 sized felines. The diaphysis is quite straight in cranial view, whereas in lateral view it is 25 26 gently craniocaudally curved (Fig. 3). Among the compared felines, only Le. serval shows 27 28 29 a slightly straighter diaphysis. In proximal view, the proximal epiphysis of L. vallesiensis 30 31 shows an elliptic articular head, slightly laterally flattened, with a smooth greater tubercle 32 33 developed along the craniolateral margin, and a craniomedially projected lesser tubercle on 34 35 36 the medial margin. The greater tubercle of Lp. geoffroyi is less cranially expanded onto the 37 38 articular head than those of L. vallesiensis and the other extant compared felines. Also, the 39 40 41 articular head is rounded (not elliptic) in Lp. wiedii, C. caracal, Ly. lynx, Ly. pardinus, and 42 43 Ly. rufus. The intertubercular groove of L. vallesiensis is well developed and relatively 44 45 46 wide, very similar to that of the compared felines. 47 48 In medial view, the greater tubercle of L. vallesiensis, as well as those of F. 49 50 silvestris, Le. serval, Ly. lynx, Ly. pardinus, and Ly. rufus is more proximally projected than 51 52 53 those of Lp. wiedii and Lp. geoffroyi, clearly surpassing the level of the articular head. A 54 55 right humerus of S. turnauensis from La Grive-Saint-Alban illustrated by Galliard (1899), 56 57 58 with the catalogue number L. Gr. 1216 shows a reduced greater tubercle, similar to those of 59 60 Lp. wiedii and Lp. geoffroyi. In cranial view, the lesser tubercle of L. vallesiensis is 61 62 63 12 64 65 1 2 3 4 medially projected (Fig. 3a), with an almost vertical medial border, and a small, round 5 6 7 proximal tuberosity that is part of the attachment area of the m. subscapularis, located on 8 9 the medial surface of the lesser tubercle. Leopardus wiedii, Lp. geoffroyi, F. silvestris, Le. 10 11 serval, Ly. lynx, Ly. pardinus, and Ly. rufus show this pattern, although the lesser tubercle 12 13 14 is much more medially projected than in L. vallesiensis. Just distally to the lesser tubercle, 15 16 there is a short crest that barely extends distally onto the diaphysis, similarly developed in 17 18 19 all the compared species. The attachment area for the m. subscapularis, which occupies the 20 21 cranial margin of the lesser tubercle, is much more distally elongated in L. vallesiensis, F. 22 23 24 silvestris, Le. serval, Ly. lynx, Ly. pardinus, and Ly. rufus than in Lp. wiedii, whereas Lp. 25 26 geoffroyi shows an intermediate pattern. In cranial view, two different orientations of the 27 28 29 proximal margin of the greater tubercle can be clearly observed: in L. vallesiensis, F. 30 31 silvestris, Le. serval, Ly. lynx, Ly. pardinus, and Ly. rufus, this margin is more cranially 32 33 inclined than those of Lp. wiedii and Lp. geoffroyi, probably due to the lesser development 34 35 36 of the greater tubercle. From the illustration by Gaillard (1899), the primitive S. turnauensis 37 38 shows a close morphology to that of these latter South American felines. The crest of the 39 40 41 greater tubercle of L. vallesiensis is very gentle, and it extends along the cranial border of 42 43 the humerus until the middle part of the diaphysis, where a very smooth deltoid tuberosity 44 45 46 is developed; this morphology is also observed in F. silvestris, Lp. wiedii, Lp. geoffroyi, C. 47 48 caracal, and Le. serval, whereas in Ly. lynx, Ly. pardinus, and Ly. rufus the crest is much 49 50 more marked, even ridged. This crest is the attachment area for the mm. pectorales 51 52 53 (superficialis and profundus) and m. deltoideus, this latter attaching on the deltoid 54 55 tuberosity (Davis 1964; Evans 1993; Barone 2010; Ercoli et al. 2015). 56 57 58 On the middle of the lateral surface of the greater tubercle of L. vallesiensis, there is 59 60 a marked elliptical scar for the attachment of the m. infraspinatus (Fig. 3b); it has ridged 61 62 63 13 64 65 1 2 3 4 margins, and it is oriented with its proximal border slightly inclined caudally, as in F. 5 6 7 silvestris and Le. serval, and also as in S. turnauensis; on the contrary, in Ly. lynx, Ly. 8 9 pardinus, Ly. rufus, Lp. wiedii, and Lp. geoffroyi, the scar is almost rounded and sub- 10 11 parallel to the diaphysis, although in the two Leopardus species, the scar is relatively 12 13 14 smaller than those of the other compared felines. In all the compared taxa, the articular 15 16 head is projected caudally, and the neck is marked, clearly distinguished from the head. The 17 18 19 tricipital line of L. vallesiensis (for the attachment of the lateral branch of the m. triceps 20 21 brachii) is marked, slightly ridged, and it develops from the attachment area for the m. 22 23 24 infraspinatus, to the distal tip of the deltoid tuberosity. In the compared sample, the line is 25 26 only absent in Lp. wiedii, being especially marked in Le. serval and Ly. pardinus. In L. 27 28 29 vallesiensis, just distal to the attachment area for the m. infraspinatus, and on the proximal 30 31 end of the tricipital line, there is a rough and irregular scar for the attachment of the m. teres 32 33 minor, similarly marked in F. silvestris and Lp. wiedii, less evident in Lp. geoffroyi, and 34 35 36 much more developed, even showing a strong ridge, in Le. serval, Ly. lynx, Ly. pardinus, 37 38 and Ly. rufus. Caudal to this attachment area, L. vallesiensis shows a strongly excavated 39 40 41 groove for the accessory branch of the m. triceps brachii; this area is shallower in F. 42 43 silvestris, Lp. wiedii, and Lp. geoffroyi, whereas in Le. serval, Ly. lynx, Ly. pardinus, Ly. 44 45 46 rufus, and S. turnauensis, it is more excavated, with a proximal strongly ridged border. 47 48 In the distal epiphysis, the medial epicondyle of L. vallesiensis, Le. serval, and Ly. 49 50 lynx is moderately medially expanded (Fig. 3a), whereas those of F. silvestris, Lp. wiedii, 51 52 53 Lp. geoffroyi, C. caracal, Ly. pardinus, and Ly. rufus are much more projected. The fossil 54 55 species S. turnauensis and M. lorteti show well-developed medial epicondyles too, much 56 57 58 more projected than that of L. vallesiensis, as shown by the specimen L. Gr. 1216 (Gaillard 59 60 1899) and an almost complete humerus of the larger M. lorteti from the Portuguese site of 61 62 63 14 64 65 1 2 3 4 Quinta das Flamengas (Ginsburg and Antunes 1995); finally, a distal fragment of a left 5 6 7 humerus of the Turolian species Pristifelis attica (AMPG PA 2803/91) published by 8 9 Roussiakis (2002) shows a medial epicondyle very similarly developed as that of L. 10 11 vallesiensis. On the other hand, the lateral epicondyle, which shows less projection than the 12 13 14 medial one, is clearly less expanded in L. vallesiensis than in any of the compared species, 15 16 which are quite similar in the development of this structure (including S. turnauensis, M. 17 18 19 lorteti, and P. attica). There is a well-developed supracondylar foramen in all the compared 20 21 species, including L. vallesiensis and P. attica. In cranial view, the trochlea is strongly 22 23 24 distally projected in all the species, although less in Lp. wiedii and Lp. geoffroyi. In medial 25 26 view, the cranial margin of the trochlea is craniodistally oriented in all the species, but 27 28 29 more projected in Ly. lynx and Ly. rufus than in L. vallesiensis, Lp. wiedii, Lp. geoffroyi, Le. 30 31 serval, Ly. pardinus, and C. caracal. In L. vallesiensis, Lp. geoffroyi, and Le. serval, the 32 33 scar for the attachment of the m. pronator teres, located on the distal end of the 34 35 36 supracondylar bar, is relatively smaller than those of F. silvestris, Lp. wiedii, C. caracal, 37 38 Ly. pardinus, Ly. rufus, and Ly. lynx, mainly because in all these latter species it is 39 40 41 proximally extended, significantly increasing the attachment surface of this muscle. Just 42 43 distal to the attachment area for the m. pronator teres there is a similarly sized surface for 44 45 46 the attachment of the m. flexor carpi radialis; this surface is relatively larger in Lp. wiedii 47 48 than in the rest of the compared species, including L. vallesiensis. In the caudal face of the 49 50 distal epiphysis, the olecranon fossa of L. vallesiensis is deep, with a ridged lateral border, 51 52 53 as observed in the rest of the compared taxa. This olecranon fossa is relatively wider in F. 54 55 silvestris, Ly. rufus, Lp. wiedii, and Lp. geoffroyi than in L. vallesiensis, Ly. lynx, Ly. 56 57 58 pardinus, C. caracal, and Le. serval, mostly because the medial margin of the olecranon 59 60 fossa is much more vertical in the latter group, whereas in the former, it opens distally. 61 62 63 15 64 65 1 2 3 4 Also in caudal view, L. vallesiensis shows a relatively mediolaterally narrower distal 5 6 7 epiphysis than those of the compared felines. The caudal surface of the medial epicondyle 8 9 shows a small round attachment surface for the medial branch of the m. triceps brachii, 10 11 very similarly developed in all the compared species except in Ly. lynx, in which it is very 12 13 14 smooth, almost unnoticeable. In the distal tip of the medial epicondyle there are a couple of 15 16 elliptical and rough attachment surfaces for the m. flexor carpi radialis and m. palmaris 17 18 19 longus, which do not show remarkable differences among the studied sample. In the caudal 20 21 surface of the lateral epicondyle of L. vallesiensis there is an excavated groove for the 22 23 24 articular capsule, especially marked in Le. serval, Ly. lynx, Ly. pardinus, and Ly. rufus. The 25 26 lateral supracondylar crest of L. vallesiensis is slightly ridged, mostly in its distal portion, 27 28 29 although it continues proximally to the middle of the diaphysis as a smooth but marked 30 31 crest; this pattern is observed in F. silvestris, Lp. wiedii, and Lp. geoffroyi, and in the fossil 32 33 species S. turnauensis and P. attica, whereas in Ly. pardinus, Ly. rufus, C. caracal, and, 34 35 36 especially in Le. serval, the crest is expanded forming a bony wall; on the other hand, in Ly. 37 38 lynx the crest, although present does not project from the bone surface, and it is formed 39 40 41 more as a cord rather than as a real crest. On the caudal surface of the crest there is a flat 42 43 and smooth surface for the attachment of the m. anconeus; the size of this surface is related 44 45 46 to the different development of the lateral supracondylar crest, and thus it is relatively 47 48 larger in Ly. pardinus, Ly. rufus, C. caracal, and, especially in Le. serval. 49 50 51 52 53 Radius 54 55 56 57 58 The radius of L. vallesiensis is slender, with a straight and strongly craniocaudally flattened 59 60 diaphysis (Fig. 3c, d). The proximal epiphysis is slightly medially inclined; its proximal 61 62 63 16 64 65 1 2 3 4 articulation facet is concave and elliptical, with a marked notch in the middle of its cranial 5 6 7 margin forming a clear capitular eminence; the lateral margin of this facet is ridged, 8 9 whereas the medial one is inclined distally, developing a smooth and slightly concave 10 11 surface. This pattern is similar in all the compared species, although in Le. serval and F. 12 13 14 lybica both the cranial notch and capitular eminence are absent, they being less developed 15 16 in C. caracal, Ly. rufus, and Ly. lynx. The proximal epiphysis is inclined medially in all the 17 18 19 compared species, with the medial border markedly medially projected, whereas the lateral 20 21 one does not stand out from the diaphysis border. The radial tuberosity of L. vallesiensis is 22 23 24 proximodistally elongated (Fig. 3d), located on the lateral border of the caudal face, and it 25 26 shows a slightly ridged lateral margin; its development is similar in all the compared 27 28 29 species. The diaphysis of L. vallesiensis, as that of the other compared species, is 30 31 craniocaudally flattened, elongated and quite straight, showing few muscular scars on the 32 33 cranial and caudal faces. 34 35 36 The distal epiphysis of L. vallesiensis shows a slightly convex cranial face, 37 38 developed as a continuation of the diaphysis, and a concave caudal face, separated from the 39 40 41 diaphysis by means of a marked ridge; this pattern is also observed in the compared sample 42 43 of felines. The medial border of the distal epiphysis of L. vallesiensis is distally projected in 44 45 46 a moderate styloid process, very similarly developed in the rest of the studied species. In L. 47 48 vallesiensis, the lateral facet for articulation with the ulna is elliptical and located in the 49 50 craniodistal border of this lateral face; the compared felines show this morphology, 51 52 53 although in Ly. lynx, Ly. pardinus, Ly. rufus, C. caracal, and Le. serval, the facet is laterally 54 55 projected; also, in the last species, the facet is not elliptical, but round. In L. vallesiensis, the 56 57 58 medial border of this distal epiphysis has a ridge, as a continuation of the medial margin of 59 60 the radius, for the attachment of the m. brachioradialis; this ridge is slightly more medially 61 62 63 17 64 65 1 2 3 4 projected in F. silvestris, F. lybica, and Lp. geoffroyi, and much more projected in Ly. lynx, 5 6 7 Ly. pardinus, Ly. rufus, C. caracal, and Le. serval. The cranial face of the distal epiphysis 8 9 of L. vallesiensis shows a couple of bony ridges, proximodistally oriented, developed for 10 11 separating the passages of the tendons of several extensor muscles: a central, 12 13 14 proximodistally shorter but thicker ridge separates a medial groove for the tendon of the m. 15 16 extensor carpi radialis from a lateral one for the tendon of the m. extensor digitorum 17 18 19 communis, whereas a lateral ridge separates this latter tendon from that of the m. extensor 20 21 digitorum lateralis, placed into a groove located on the lateral border of the distal epiphysis 22 23 24 (Evans 1993; Vollmerhaus and Roos 2001; Barone 2010). The development of these bony 25 26 ridges shows differences within the studied feline sample: in F. lybica and Le. serval, the 27 28 29 lateral ridge is markedly displaced medially in relation to its position in L. vallesiensis, F. 30 31 silvestris, Ly. lynx, Ly. rufus, Ly. pardinus, C. caracal, and Lp. geoffroyi. The distal 32 33 articular facet of L. vallesiensis is well developed, but it does not occupy the whole distal 34 35 36 surface of the epiphysis; it is strongly concave, sub-elliptical, with a marked ridge on its 37 38 craniolateral margin. This pattern is common to the other felines, although it is remarkable 39 40 41 that in Le. serval and F. lybica, the lateral border of the facet is expanded in both lateral and 42 43 caudal directions, thus producing an overall slightly larger articular surface for the 44 45 46 scapholunar than in the rest of the species. 47 48 49 50 Ulna 51 52 53 54 55 The ulna of L. vallesiensis is slender, very similar to that of other felines. The diaphysis 56 57 58 shows a gentle caudal convex curvature in its distal half, and it is markedly mediolaterally 59 60 flattened (Fig. 3e, f). The olecranon of L. vallesiensis is well developed, similarly elongated 61 62 63 18 64 65 1 2 3 4 as those of the compared felines; its proximal border, in both lateral and medial views, is 5 6 7 clearly caudally inclined, as in F. silvestris, F. lybica, Lp. wiedii, and Lp. geoffroyi, whereas 8 9 in Ly. lynx, Ly. rufus, Ly. pardinus, C. caracal, and Le. serval, the proximal border of the 10 11 olecranon is almost horizontal due to the greater proximal projection of its caudal border. 12 13 14 The length of the olecranon is similar in all the compared species, although in Ly. lynx, Le. 15 16 serval, and C. caracal, it is slightly shorter. The olecranon has a pair of proximal tubercles, 17 18 19 lateral and medial, that correspond to the attachment areas for two muscles involved in the 20 21 extension of the forearm: the m. anconeus, which attaches on the lateral tubercle, expanding 22 23 24 its attachment surface onto the caudal border of the lateral face of the proximal epiphysis, 25 26 and the medial branch (caput mediale) of the m. triceps brachii, which attaches on the 27 28 29 medial tubercle (Gonyea 1978; Barone 2010; Julik et al. 2012). In L. vallesiensis, F. 30 31 silvestris, F. lybica, and Le. serval, the medial tubercle is well developed, markedly 32 33 projected proximally, and surpassing the level of the lateral tubercle. Lynx pardinus shows 34 35 36 the opposite morphology, with the lateral tubercle much more proximally projected than the 37 38 medial one, whereas in Ly. lynx, Ly. rufus, C. caracal, Lp. wiedii, and Lp. geoffroyi both 39 40 41 tubercles show a similar proximal projection. In cranial view, the tubercles of L. 42 43 vallesiensis, F. lybica, Lp. wiedii, Lp. geoffroyi, Le. serval, Ly. rufus, and Ly. pardinus are 44 45 46 separated by means of a shallow groove, whereas in F. silvestris, Ly. lynx, and C. caracal 47 48 this groove has a similar width, but it is much deeper. 49 50 On the lateral surface of the proximal epiphysis of L. vallesiensis, originated from 51 52 53 the level of the middle point of the trochlear notch, there is a proximodistally elongated 54 55 triangular groove for the attachment of the m. extensor digiti I et II (Fig. 3e); the groove 56 57 58 continues distally onto the diaphysis by means of a caudal ridge until the distal epiphysis, 59 60 thus forming an attachment surface for the m. abductor digiti I longus (=m. extensor carpi 61 62 63 19 64 65 1 2 3 4 obliquus) (Barone 2010; Julik et al. 2012); the most proximal portion of this groove is also 5 6 7 well marked in F. lybica, F. silvestris, and Le. serval, and much less excavated in Ly. lynx, 8 9 Ly. pardinus, Ly. rufus, C. caracal, Lp. wiedii, and Lp. geoffroyi, probably indicating a 10 11 more powerful m. extensor digiti I et II in the former group. On the medial surface of the 12 13 14 proximal epiphysis, close to the beginning of the diaphysis there is a proximodistally 15 16 elongated scar for the common attachment of the m. brachialis and m. cleidobrachialis, very 17 18 19 similarly developed in all the compared species. On the medial surface of the diaphysis 20 21 there is a caudal proximodistally elongated groove for the attachment of the ulnar branch of 22 23 24 the m. flexor digitorum profundus (Fig. 3f); also, in the distal half of the diaphysis, a 25 26 marked ridge is developed, forming a cranially oriented surface for the attachment of the m. 27 28 29 pronator quadratus; this structure does not show differences among the compared felines. 30 31 The distal epiphysis of L. vallesiensis shows the basic feline morphology: that is 32 33 mediolaterally flattened with a distocaudally oriented styloid process. 34 35 36 37 38 Metacarpal I 39 40 41 42 43 The Mc I of L. vallesiensis shows similar proportions as in F. silvestris and F. lybica, 44 45 46 which have slenderer Mc I than Le. serval, Lp. wiedii, Lp. geoffroyi, Ly. lynx, Ly. rufus, and 47 48 Ly. pardinus. A marked, dorsopalmarly developed ridge (Fig. 4a) divides the base into a 49 50 smooth and dorsally expanded articular surface for the trapezium, and a medial tuberosity 51 52 53 for the attachment of the m. abductor digiti I longus and one of the dorsal carpometacarpal 54 55 ligaments (Fig. 4b). In proximal view, both parts show a similar width in all the compared 56 57 58 species. The body is straight and more or less cylindrical in L. vallesiensis, F. silvestris, and 59 60 F. lybica, and relatively shorter and slightly dorsopalmarly flattened in Le. serval, Lp. 61 62 63 20 64 65 1 2 3 4 wiedii, Lp. geoffroyi, Ly. lynx, Ly. rufus, and Ly. pardinus. The head is markedly 5 6 7 asymmetrical, as the lateral border is much more distally projected than the medial one; on 8 9 its palmar surface, there is a sharp keel, palmarly projected and similarly developed in all 10 11 the compared species. 12 13 14 15 16 Metacarpal II 17 18 19 20 21 The Mc II of L. vallesiensis shows similar proportions than those of Lp. wiedii, Lp. 22 23 24 geoffroyi, C. caracal, Ly. lynx, Ly. rufus, and Ly. pardinus, whereas F. silvestris, F. lybica, 25 26 and Le. serval have slenderer Mc II. The overall morphology of this bone is very similar in 27 28 29 L. vallesiensis and the compared felines. The proximal articular surface has a trapezoid 30 31 shape, with its dorsal border mediolaterally wider than the palmar one. In dorsal view, the 32 33 base is clearly laterally inclined, and it shows a groove for the passage of the radial artery. 34 35 36 In lateral view (Fig. 4c), the palmar tubercle is large and projected; the groove for 37 38 articulation with the medial face of the Mc III is proximodorsally located and well 39 40 41 excavated. In medial view (Fig. 4d), the round facet for articulation with the trapezium 42 43 barely surpasses the level of the proximal border in L. vallesiensis, F. lybica, and F. 44 45 46 silvestris, whereas in Lp. wiedii, Lp. geoffroyi, C. caracal, Le. serval, Ly. lynx, Ly. rufus, 47 48 and Ly. pardinus the facet is more proximally projected, widely surpassing the level of the 49 50 proximal border. Distal to this facet, there is a rough tubercle for the attachment of the short 51 52 53 interosseous ligament that connects Mc I and Mc II, similarly developed in L. vallesiensis 54 55 and all the compared species. In the palmar border of the base there is a small but distinct 56 57 58 scar for the attachment of the m. flexor carpi radialis, slightly more distally extended in F. 59 60 silvestris, F. lybica, Le. serval, Ly. lynx, Ly. rufus, Lp. wiedii, and Lp. geoffroyi than in L. 61 62 63 21 64 65 1 2 3 4 vallesiensis, Ly. pardinus, and C. caracal. The body is slightly curved in lateral and medial 5 6 7 view in all the species, with Ly. lynx and Ly. pardinus showing the straightest ones. The 8 9 distal head of L. vallesiensis, Lp. geoffroyi, C. caracal, and Ly. pardinus has a well- 10 11 developed proximopalmarly projected central keel, whereas in F. silvestris, F. lybica, Le. 12 13 14 serval, Ly. lynx, and Ly. rufus, this structure is clearly reduced. Also, the head is relatively 15 16 smaller in F. lybica, F. silvestris, and Le. serval than in L. vallesiensis and the rest of 17 18 19 compared species. 20 21 22 23 24 Metacarpal III 25 26 27 28 29 The Mc III of L. vallesiensis shows similar proportions as in F. silvestris, F. lybica, C. 30 31 caracal, Ly. rufus, and Ly. pardinus; in this group, the Mc III shows indeed an intermediate 32 33 robustness between that of the slenderer Ly. lynx and Le. serval (which show the most 34 35 36 gracile Mc III) and that of the more robust Lp. wiedii and Lp. geoffroyi. In all the compared 37 38 species, the proximal articular surface is triangular, with a marked notch on its medial 39 40 41 border; in dorsal view, the base is clearly laterally inclined. The lateral face of the base 42 43 shows the articulation surfaces for the Mc IV (Fig. 4e): a large dorsally located groove and 44 45 46 a dorsopalmarly elongated facet, located proximally to the former. This facet shows several 47 48 differences within the compared sample of felines: although it occupies the whole 49 50 dorsopalmar margin in all the species, in L. vallesiensis, Ly. lynx, and Le. serval it clearly 51 52 53 lacks the distal expansion of its palmar end observed in F. silvestris, F. lybica, C. caracal, 54 55 Ly. rufus, Ly. pardinus, Lp. wiedii, and Lp. geoffroyi; all these taxa show a central 56 57 58 constriction in the facet, more marked in F. silvestris, and absent in L. vallesiensis, Ly. lynx, 59 60 and Le. serval. The medial face of the base (Fig. 4f) shows two articular facets for the Mc 61 62 63 22 64 65 1 2 3 4 II, one dorsally and another one palmarly located, widely separated by a rough fossa. These 5 6 7 facets show subtle differences within the sample: in Lp. wiedii, Lp. geoffroyi, Ly. pardinus, 8 9 and Ly. rufus the dorsal facet is relatively larger than those of L. vallesiensis, F. silvestris, 10 11 F. lybica, C. caracal, Le. serval, and Ly. lynx, whereas the palmar facet shows less size 12 13 14 differences. Finally, the head is relatively larger in Lp. wiedii and Lp. geoffroyi than in L. 15 16 vallesiensis and the rest of the compared species. 17 18 19 20 21 Metacarpal IV 22 23 24 25 26 As in the case of the Mc III, the Mc IV of L. vallesiensis shows similar proportions as those 27 28 29 of F. silvestris, F. lybica, C. caracal, Ly. rufus, and Ly. pardinus, which are more robust 30 31 than those of Ly. lynx and Le. serval. Leopardus wiedii and Lp. geoffroyi show a much 32 33 more robust Mc IV than the rest of the compared sample. All the compared species show a 34 35 36 rectangular proximal articular surface, with a medially projected facet for the Mc III; in 37 38 dorsal view, the proximal border of the base is laterally inclined. The lateral face of the 39 40 41 base (Fig. 4g) is occupied by the articulation surfaces for the Mc V: a large and deep central 42 43 groove and a curved and elongated facet developed along the dorsal, proximal, and palmar 44 45 46 margins. Except for its shorter dorsopalmar length in Le. serval and Ly. lynx, the 47 48 morphology of this face is very similar in all the compared species. The medial face of the 49 50 base (Fig. 4h) shows two articular facets for the Mc III, both located near the proximal 51 52 53 border, one dorsally and the other one palmarly located, and widely separated by a smooth 54 55 fossa. Whereas the dorsal facet has a similar development in all the taxa, the palmar facet of 56 57 58 L. vallesiensis is much less proximodistally expanded, having the shape of a narrow cord, 59 60 very different from the round to triangle facet seen in the rest of the compared felines. 61 62 63 23 64 65 1 2 3 4 Finally, the head is relatively larger again in Lp. wiedii and Lp. geoffroyi than in L. 5 6 7 vallesiensis and the rest of compared species. 8 9 10 11 Metacarpal V 12 13 14 15 16 The Mc V of L. vallesiensis shows similar proportions to those of F. silvestris, F. lybica, C. 17 18 19 caracal, Ly. lynx, Ly. rufus, and Ly. pardinus. Once again, the Mc V of Le. serval is the 20 21 most gracile of the sample, whereas those of Lp. wiedii and Lp. geoffroyi are relatively 22 23 24 much shorter and stouter, with the latter being remarkably robust. In proximal view, the Mc 25 26 V of L. vallesiensis shows a semicircular proximal articulation surface that is divided in 27 28 29 two parts by means of a marked dorsopalmarly developed groove; this morphology is more 30 31 or less similar in all the compared species. On the lateral face of the base (Fig. 4i) there is 32 33 an elliptical attachment surface for the m. extensor carpi ulnaris, dorsopalmarly elongated 34 35 36 in L. vallesiensis and Le. serval, and round and more distally expanded in F. silvestris, F. 37 38 lybica, Ly. lynx, Ly. rufus, Ly. pardinus, C. caracal, Lp. wiedii, and Lp. geoffroyi. The 39 40 41 medial face of the base (Fig. 4j) shows a central medially expanded and proximodistally 42 43 elongated bony sheet, surrounded by a smooth facet occupying the proximal and dorsal 44 45 46 margins. The head is relatively larger again in Lp. wiedii and Lp. geoffroyi than in L. 47 48 vallesiensis and the rest of compared species. 49 50 51 52 53 Os Coxae 54 55 56 57 58 The Batallones-1 sample includes the complete right and left os coxae of a single individual 59 60 of L. vallesiensis (Fig. 5). Their general shape and morphology are very similar to those of 61 62 63 24 64 65 1 2 3 4 the compared felines. The acetabular fossa is rounded and located close to the cranial 5 6 7 margin of the obturator foramen. The wing of the ilium of L. vallesiensis is slender, 8 9 craniocaudally elongated, with more or less parallel dorsal and ventral margins and a gently 10 11 curved cranial border (crest of the ilium). The gluteal fossa is shallow but evident, with a 12 13 14 thickened dorsal margin. In Ly. lynx, Ly. pardinus, Ly. rufus, F. lybica, F. silvestris, Lp. 15 16 wiedii, and Lp. geoffroyi, the body of ilium shows a central constriction due to the ventral 17 18 19 projection of the tuber coxae and the marked expansion of the tuber sacrale, whereas in L. 20 21 vallesiensis, C. caracal, and Le. serval, the corpus of the ilium does not show any 22 23 24 constriction. In the cranioventral margin of the wing of Ly. lynx, Ly. pardinus, Ly. rufus, F. 25 26 lybica, Le. serval, and C. caracal there is a triangular surface, with ridged margins, for the 27 28 29 attachment of the m. sartorius; this surface is very reduced in L. vallesiensis, F. silvestris, 30 31 Lp. wiedii, and Lp. geoffroyi. The ischiatic spine of L. vallesiensis, Le. serval, and C. 32 33 caracal is located slightly more caudally than in Ly. lynx, Ly. pardinus, Ly. rufus, F. lybica, 34 35 36 F. silvestris, Lp. wiedii, and Lp. geoffroyi. In dorsal view, the orientation of the wings of L. 37 38 vallesiensis is very similar to that of the compared species, slightly laterally curved but 39 40 41 keeping the dorsoventral plane. In this dorsal view, the acetabular fossa is slightly more 42 43 laterally projected than in Le. serval, C. caracal, F. lybica, F. silvestris, Lp. wiedii, and Lp. 44 45 46 geoffroyi, and similar to that of Ly. lynx, Ly. rufus, and Ly. pardinus. The caudal half of 47 48 BAT-1’06 D4-66a is damaged, and the actual shape of the obturator foramen cannot be 49 50 assessed, but it probably was similar to that of the compared species. The ischiatic 51 52 53 tuberosity of L. vallesiensis and the compared species is rounded and rough, but the 54 55 ischiatic arch shows some differences within the studied sample: in L. vallesiensis, F. 56 57 58 lybica, Lp. wiedii, and Lp. geoffroyi, the caudal margin of the ischium is straight, and so the 59 60 61 62 63 25 64 65 1 2 3 4 ischiatic arch is small, whereas in F. silvestris, Ly. lynx, Ly. pardinus, Ly. rufus, Le. serval 5 6 7 and C. caracal this margin describes a gentle curvature, producing a wide and larger arch. 8 9 10 11 Sacrum 12 13 14 15 16 A complete sacrum of L. vallesiensis, which was articulated to the left and right os coxae 17 18 19 described above, is known from Batallones-1. It is composed, as that of other felids, of 20 21 three fused sacral vertebrae. It is triangular in overall shape, with its maximum mediolateral 22 23 24 width at its cranial margin. This morphology is very similar to that of the compared species, 25 26 although Ly. lynx, Ly. pardinus, Ly. rufus, and C. caracal show a relatively more elongated 27 28 29 sacrum. The available sacrum of L. vallesiensis is almost complete, but it lacks the dorsal 30 31 spines, which only preserve their lower parts, making it impossible to assess their actual 32 33 development. An intermediate sacral crest, mediolaterally flattened, is preserved in the right 34 35 36 side: its cranial portion is cranially oriented, whereas the caudal one is smaller, lower, and 37 38 vertical (Fig. 6a). This pattern is also observed in F. silvestris, F. lybica, and Lp. geoffroyi, 39 40 41 whereas in Ly. lynx, Ly. pardinus, Ly. rufus, Le. serval, and C. caracal, the caudal process 42 43 is absent. In L. vallesiensis, the caudal portion of the lateral sacral crest (=transverse 44 45 46 process of the last sacral vertebra) is developed as a long and caudally pointed sheet of 47 48 bone, as in F. silvestris and C. caracal. In F. lybica, Lp. geoffroyi, Ly. lynx, Ly. pardinus, 49 50 and Ly. rufus, these processes are shorter and less pointed, whereas in Le. serval they are 51 52 53 extremely thin and slender. This caudal portion of the lateral crest in the Batallones feline 54 55 shows a central keel, also present, in more or less extent, in the rest of the compared felines. 56 57 58 The lateral sacral crest is also laterally projected at the level of the second sacral vertebra in 59 60 L. vallesiensis, C. caracal, and Le. serval, it being thus much more developed than in F. 61 62 63 26 64 65 1 2 3 4 silvestris, F. lybica, Ly. lynx, Ly. pardinus, Ly. rufus, and Lp. geoffroyi. At least four dorsal 5 6 7 sacral formamina are observed in all the compared extant felines, but they are absent in S. 8 9 vallesiensis. On ventral view, the sacrum of L. vallesiensis shows four ventral sacral 10 11 foramina (Fig. 6b), as the rest of the studied felines. 12 13 14 The wings of the sacrum of L. vallesiensis are very similar to those of the compared 15 16 felines, that is, clearly ventrally projected, with a rough lateral surface. Dorsally, the wings 17 18 19 contact with the cranial articular surfaces of the sacrum, the articulation area for the last 20 21 lumbar vertebra. These two cranial articular surfaces are separated by a caudally expanded 22 23 24 notch. In L. vallesiensis, C. caracal, and Le. serval, this notch is relatively wider and less 25 26 caudally expanded than those of F. silvestris, F. lybica, Ly. lynx, Ly. pardinus, Ly. rufus, 27 28 29 and Lp. geofrroyi. 30 31 32 33 Femur 34 35 36 37 38 The Batallones-1 sample includes two femora of L. vallesiensis belonging to the same 39 40 41 individual. The femoral head is projected proximomedially by means of a well-developed 42 43 neck and it is located at the same level of the greater trochanter (Fig. 7); only in C. caracal 44 45 46 and Lp. geoffroyi does the head surpass the level of the greater trochanter. In lateral view, 47 48 the femur of L. vallesiensis has a rough gluteal tuberosity with a strongly ridged cranial 49 50 margin that contacts proximally with the greater trochanter (Fig. 7a); distal to the gluteal 51 52 53 tuberosity, a less marked ridge continues distally on the lateral border, until the middle of 54 55 the diaphysis. This pattern is seen in the rest of the studied species, and the only remarkable 56 57 58 difference is the position of the gluteal tuberosity in L. vallesiensis and Lp. wiedii, located 59 60 distal to the lesser trochanter, whereas in the other species it is clearly proximal to this 61 62 63 27 64 65 1 2 3 4 structure. On the proximal tip of the greater trochanter, the attachment areas for the m. 5 6 7 gluteus profundus and m. gluteus medius show a similar pattern in all the taxa, with the 8 9 attachment area for the m. gluteus medius located proximal to that of the m. gluteus 10 11 profundus, both being restricted to the proximal surface of the greater trochanter. In caudal 12 13 14 view (Fig. 7b), the proximal epiphysis of L. vallesiensis shows a deep trochanteric fossa, a 15 16 ridged and relatively short intertrochanteric crest, and a caudally projected lesser 17 18 19 trochanter; in the Batallones feline, as well as in F. lybica, Ly. lynx, Ly. pardinus, Ly. rufus 20 21 and Lp. geoffroyi, the lesser trochanter is located close to the medial margin, but laterally 22 23 24 displaced in relation to its position in F. silvestris, Le. serval, C. caracal, and Lp. wiedii. 25 26 This different position of the lesser trochanter is also related to the orientation of the 27 28 29 attachment area for the m. iliopsoas, which is more or less caudally oriented in the former 30 31 group of felines, and medially faced in the latter. 32 33 In the distal epiphysis of L. vallesiensis, the lateral condyle is mediolaterally wider 34 35 36 than the medial one. In caudal view (Fig. 7b), the medial condyle is vertical, parallel to the 37 38 proximodistal axis of the femur, whereas the lateral one is slightly laterally inclined, with 39 40 41 its proximolateral vertex showing a proximolaterally oriented expansion. Although F. 42 43 silvestris, Lp. wiedii, and Lp. geoffroyi show this pattern, in Le. serval, F. lybica, Ly. lynx, 44 45 46 Ly. pardinus, Ly. rufus, and C. caracal, the medial condyle is slightly but clearly laterally 47 48 inclined. Also, in L. vallesiensis, F. silvestris, F. lybica, Lp. wiedii, and Lp. geoffroyi, both 49 50 condyles reach the same level distally, whereas in Le. serval, Ly. lynx, Ly. rufus, Ly. 51 52 53 pardinus, and C. caracal, the medial condyle distally surpasses the level of the lateral one. 54 55 In caudal view, the intercondyloid fossa of L. vallesiensis is well developed and shows a 56 57 58 similar mediolateral width than those of the other compared felines. In distal view, all the 59 60 compared taxa show a medial condyle slightly more caudally projected than the lateral one. 61 62 63 28 64 65 1 2 3 4 In lateral and medial views, both the craniocaudal length of the distal epiphysis, and the 5 6 7 curvature of the femoral trochlea of L. vallesiensis are very similar to those of the other 8 9 compared felines. The trochlea of L. vallesiensis is more proximally extended than those of 10 11 any of the compared feline species (Fig. 7a), which indicates a greater range of patellar 12 13 14 extension, as has been suggested for other carnivorans (Siliceo 2015). 15 16 17 18 19 Tibia 20 21 22 23 24 The two tibiae of L. vallesiensis from Batallones-1 are complete but strongly laterally 25 26 compressed, thus preventing a clear assessment of some of their characters. It is slender in 27 28 29 overall view, with a more or less straight diaphysis and a caudally curved proximal 30 31 epiphysis (Fig. 7c–e). The proximal epiphysis of L. vallesiensis shows a triangular-shaped 32 33 proximal surface, with the two articular condyles (medial and lateral) separated by a rough 34 35 36 and narrow intercondyloid area. In cranial or caudal views, the lateral condyle in L. 37 38 vallesiensis is located slightly more proximal than the medial one (Fig. 7c), mainly due to 39 40 41 the ridged medial margin of the lateral condyle (the intercondyloid eminence); this pattern 42 43 is observed in all the compared felines. In proximal view, the cranial border is strongly 44 45 46 cranially projected, with a marked notch on its lateral margin for the passage of the tendon 47 48 of the m. extensor digitorum longus (Fig. 7e). The medial condyle of L. vallesiensis is 49 50 clearly less caudally projected than the lateral one, as in the rest of the compared felines. 51 52 53 The tibial tuberosity has a smooth proximal surface separated by a ridge from the cranial 54 55 border, which runs distally until the middle of the cranial margin of the diaphysis. 56 57 58 The diaphysis of L. vallesisensis and the compared felines is strongly mediolaterally 59 60 flattened on its proximal half, whereas the distal half has a square section. Also, the 61 62 63 29 64 65 1 2 3 4 proximal half is laterally inclined in cranial view and caudally in both lateral and medial 5 6 7 views; the distal half is quite straight. The morphology of the diaphysis is similar in all the 8 9 compared species, except for the development of a strong ridge, proximodistally developed, 10 11 on the caudal face of the diaphysis of the tibia of L. vallesiensis (Fig. 7c) and Lp. geoffroyi. 12 13 14 Interestingly, this ridge has been also observed by de Beaumont (1986) in the Asian golden 15 16 cat (Catopuma temminckii) and the fossil feline Pristifelis attica, whereas it is absent in the 17 18 19 tibia of S. turnauensis figured by de Beaumomt (1961) and that of M. lorteti figured by 20 21 Salesa et al. (2011). Among the studied sample of carnivorans, it is absent in Genetta 22 23 24 genetta, Martes foina, Gulo, gulo, Meles meles, Ailurus fulgens, and Canis lupus, and 25 26 present in Ursus americanus. This ridge seems to have separated in life the m. flexor 27 28 29 digitorum lateralis and m. flexor digitorum medialis + m. tibialis caudalis, whose tendons 30 31 run together along the same groove (Barone 2010). These muscles originate on the 32 33 proximal epiphysis of the tibia, and are very closely disposed on the caudal surface of the 34 35 36 diaphysis (Fisher et al. 2008; Ercoli et al. 2012). Both muscles have strong and long distal 37 38 tendons, but the relation between the fleshy part and the tendon varies among the 39 40 41 Carnivora, with felids and canids having in general longer tendons (and then shorter fleshy 42 43 parts) than ailurids, ursids, procyonids, or mustelids (Davis 1964; Feeny 1999; 44 45 46 Vollmerhaus and Roos 2001; Fisher et al. 2008; Ercoli et al. 2012). Thus, the described 47 48 ridge in the caudal face of the tibia of L. vallesiensis and Lp. geoffroyi, which in the former 49 50 reaches the proximal border of the distal epiphysis, would be actually forming a groove for 51 52 53 the accommodation of the tendons of both muscles, m. flexor digitorum medialis + m. 54 55 tibialis caudalis on the medial side of the ridge, and m. flexor digitorum lateralis on its 56 57 58 lateral side. 59 60 61 62 63 30 64 65 1 2 3 4 The distal epiphysis of L. vallesiensis strongly resembles that of the compared 5 6 7 felines, with no feature showing significant differences. 8 9 10 11 Talus 12 13 14 15 16 The overall morphology and proportions of the talus of L. vallesiensis are quite similar to 17 18 19 those of the compared felines. The trochlea is wide, with a shallow central groove that 20 21 divides it into two asymmetrical lips (Fig. 8a), as the medial one is narrower than the lateral 22 23 24 one; the distal border of the lateral lip projects distally surpassing the level of the medial 25 26 one, which does not expand onto the neck. In proximal view, the lateral lip is more dorsally 27 28 29 projected than the medial one in L. vallesiensis and all the compared felines. The proximal 30 31 surface of the trochlea of L. vallesiensis shows a transversal shallow groove and a small, 32 33 medially located talar foramen for the posterior tibial artery, as in F. silvestris, F. lybica, 34 35 36 Le. serval, Lp. wiedii, Ly. pardinus, and Ly. rufus, whereas in C. caracal, Lp. geoffroyi, and 37 38 Ly. lynx it is absent. In plantar view (Fig. 8b), a relatively narrow sulcus tali separates two 39 40 41 large articular facets for the calcaneus: the lateral facet is rectangular and strongly concave, 42 43 whereas the medial one is elliptical, convex, and occupies most of the plantar surface of the 44 45 46 neck, joining the distal articular surface of the talus head by means of a narrow projection; 47 48 this morphology is observed in most of the compared species, although in F. silvestris, F. 49 50 lybica, C. caracal, and Le. serval the medial facet is not connected to the distal facet for the 51 52 53 navicular. The neck of L. vallesiensis is as long as those of F. silvestris, F. lybica, C. 54 55 caracal, Le. serval, Lp. wiedii, and Lp. geoffroyi, and longer than those of Ly. lynx, Ly. 56 57 58 pardinus, and Ly. rufus. The head of the talus in all the species is well distinguished from 59 60 the neck as it has a gently ridged dorsal margin; in distal view, the head is elliptical in L. 61 62 63 31 64 65 1 2 3 4 vallesiensis, with the lateral border being slightly orientated dorsally relative to the medial 5 6 7 one; this orientation is more or less the same in the compared felines, although the head is 8 9 round rather than elliptical in all the species. 10 11 12 13 14 Calcaneus 15 16 17 18 19 The calcaneus of L. vallesiensis and the compared feline species shows a very similar 20 21 morphology and relative proportions. In dorsal view (Fig. 8c), the fibular tubercle 22 23 24 (distolateral expansion for the attachment of the ligament calcaneofibular or collaterale tarsi 25 26 laterale longum) shows little lateral projection, and is located close to the distolateral 27 28 29 border; its morphology is very similar in all the compared species. The sustentaculum tali is 30 31 well medially projected in all the compared species, showing a rounded medial talar facet 32 33 that continues distally through a narrow and long talar facet until the distomedial border of 34 35 36 the talar surface; this facet is almost dorsally oriented in L. vallesiensis, Lp. wiedii, Lp. 37 38 geoffroyii, F. silvestris, and F. lybica, whereas in C. caracal, Le. serval, Ly. lynx, Ly. rufus, 39 40 41 and Ly. pardinus, it is clearly dorsomedially oriented. The tuber calcanei is relatively 42 43 slender and mediolaterally flattened in all the compared species. Also, the medial tubercle 44 45 46 of the tuber is more proximally projected than the lateral one, although in Le. serval and Ly. 47 48 rufus the difference is not so marked. In plantar view (Fig. 8e) the sustentaculum tali shows 49 50 a similar medial projection in all the compared felines, although in C. caracal and Le. 51 52 53 serval, this structure is slightly more proximally located; the groove for the tendon of the 54 55 m. flexor digitorum lateralis is similarly marked in all the species. On the distal end of this 56 57 58 plantar face, a scar for the ligament plantare longum is observed. In lateral view, the fibular 59 60 tubercle of L. vallesiensis is located very close to the distal border of the lateral margin, it 61 62 63 32 64 65 1 2 3 4 showing a marked oblique groove, also seen in F. silvestris, F. lybica, C. caracal, Le. 5 6 7 serval, and Ly. lynx. Plantar to this tubercle, there is a shallow depression with fairly ridged 8 9 plantar and dorsal margins for the attachment of the m. quadratus plantae (Fig. 8e); in L. 10 11 vallesiensis and Lp. geoffroyii, this attachment area is proximally developed until the level 12 13 14 of the coracoid process, whereas in Lp. wiedii and F. silvestris, it is even more proximally 15 16 elongated, reaching the middle point of the tuber calcanei; on the contrary, the attachment 17 18 19 surface is reduced in F. lybica, C. caracal, Le. serval, Ly. lynx, Ly. rufus, and Ly. pardinus, 20 21 as it barely surpasses the level of the fibular tubercle. In proximal view, the tuber calcanei 22 23 24 shows an elliptical shape, dorsoplantarly elongated, with the medial tubercle larger than the 25 26 lateral one. In distal view, the articular facet for the cuboid is almost round in all the 27 28 29 compared feline species. 30 31 32 33 Comparative anatomy and biomechanical implications 34 35 36 37 38 Lumbar vertebrae 39 40 41 42 43 Both the overall morphology and relative proportions of the L7-L5 of L. vallesiensis are 44 45 46 very similar to those of the compared felines: in all of them the L7 is shorter than the rest of 47 48 the lumbar vertebrae, with F. silvestris, F. lybica, and Lp. geoffroyi showing the relatively 49 50 longest L7 (Fig. 9). In felids and other carnivorans, the extension and flexion movements of 51 52 53 the lumbar region (and the vertebral column in general) are very important in galloping 54 55 (Hildebrand 1959; Goslow et al. 1973; Gambaryan 1974; Taylor 1978; English 1980; 56 57 58 Alexander and Jayes 1981). These movements of the column basically contribute to stride 59 60 length and limb speed through the action of the epaxial and abdominal muscles (Hildebrand 61 62 63 33 64 65 1 2 3 4 1959; Goslow et al. 1973; English 1980). Thus, a great flexibility of the lumbar series in 5 6 7 felines is necessary for increasing both stride length and limb speed, something critical in 8 9 cursorial species such as Acinonyx jubatus (Hildebrand 1959; Gambaryan 1974). A 10 11 relatively short L7 is typical of arboreal viverrids and mustelids, such as G. genetta or 12 13 14 Martes foina, animals that do not move primarily on the ground, but the relative length of 15 16 this vertebra in A. jubatus is very similar to that of these carnivorans, and slightly shorter 17 18 19 than those of F. silvestris, F. lybica, and Lp. geoffroyi. In consequence, there are only subtle 20 21 differences in the relative length of L7 among the compared sample of felines, and the 22 23 24 significance of a relative short L7 in L. vallesiensis (Fig. 9), similar to those of viverrids 25 26 (but also mustelids) should be considered as a primitive feature with no clear functional 27 28 29 implications. 30 31 32 33 Humerus 34 35 36 37 38 The differences in the lateral width of the humeral head observed among our sample of 39 40 41 felines have been previously described in other carnivorans; for example, Taylor (1974) 42 43 already noticed it in his study on herpestids and viverrids, with the more terrestrial species 44 45 46 having mediolaterally flattened heads and a relatively limited range of movement in the 47 48 shoulder, both in the craniocaudal and mediolateral planes. Argot (2001), and Argot and 49 50 Babot (2011) proposed that a similar lateral compression of the humeral head observed in 51 52 53 Paleocene and Eocene marsupials suggested that movements of the glenohumeral 54 55 articulation were restricted to flexion/extension, indicative of terrestrial habits. This would 56 57 58 be the case of L. vallesiensis, which has a less rounded humeral head than other compared 59 60 felines, a probable indication of a mostly terrestrial way of life. Other humeral structures 61 62 63 34 64 65 1 2 3 4 have been revealed as good indicators of terrestrial versus arboreal habits in several groups 5 6 7 of mammals, including the development of the greater tubercle (Jolly 1967; Taylor 1974; 8 9 Fleagle and Simons 1982; Ciochon 1986; Larson and Stern 1989; Gebo and Rose 1993; 10 11 Siliceo et al. 2015). All these authors concluded that the more arboreal species had less 12 13 14 projected tubercles than those moving mainly on the ground. Focusing on the sample of 15 16 Felinae studied here, Lp. wiedii and Lp. geoffroyi show greater tubercles that are less 17 18 19 proximally projected than that of L. vallesiensis and those extant species with typically 20 21 terrestrial lifestyles, such as the lynxes and the wildcat (Fig. 10); this is interesting, as Lp. 22 23 24 wiedii is one of the most arboreal felines, and although Lp. geoffroyi occupies a great 25 26 diversity of habitats, it is often found in woodlands (Sunquist and Sunquist 2009). The 27 28 29 greater tubercle is the attachment area for the m. supraspinatus, which originates on the 30 31 supraspinous fossa of the scapula (Barone 2010; Julik et al. 2012), extending the humerus 32 33 and rotating it laterally (Taylor 1974; Barone 2010; Ercoli et al. 2015). The proximal 34 35 36 expansion of the greater tubercle produces the lengthening of the moment arm of the m. 37 38 supraspinatus at the glenohumeral articulation (Jolly 1967; Larson and Stern 1989) and it 39 40 41 has been interpreted in baboons as a way of achieving a more economical locomotion on 42 43 the ground (Jolly 1967); on the contrary, a less projected greater tubercle produces a shorter 44 45 46 lever arm for the m. supraspinatus, which favours fast (although weak) flexion of the 47 48 humerus during arboreal locomotion (Jolly 1967; Larson and Stern 1989). It is remarkable 49 50 that the middle Miocene feline S. turnauensis resembles Lp. wiedii and Lp. geoffroyii in this 51 52 53 feature, suggesting an important arboreal component in its locomotor behavior. On the 54 55 contrary, the development of the greater tubercle of the late Miocene L. vallesiensis 56 57 58 indicates a primarily terrestrial locomotion, in a similar way to other extant felines. These 59 60 different locomotor adaptations would reflect the climate changes at the beginning of the 61 62 63 35 64 65 1 2 3 4 late Miocene in Europe, which produced the predominance of drier, less structured 5 6 7 landscapes than those from the more humid middle Miocene (Agustí and Moyà-Solà 1990; 8 9 Agustí et al. 1997; Fortelius and Hokkanen 2001; Hernández-Fernández et al. 2006). 10 11 Among the Carnivora, the medial epicondyle of the humerus is the attachment area 12 13 14 for several pronator and flexor muscles of the forearm and carpus (Davis 1964; Barone 15 16 1967, 2010; Taylor 1974; Salesa et al. 2008; Fisher et al. 2009; Julik et al. 2012; Ercoli et 17 18 19 al. 2015). The degree of medial projection of this humeral structure is an indicator of the 20 21 development of these muscles in fossil carnivorans, and thus it has been used to infer 22 23 24 several aspects of their paleoecology: those species with reduced medial epicondyles show 25 26 smaller flexor muscles, and have slender limbs, are cursorial hunters, and use to live in 27 28 29 open, low-structured habitats, whereas those with well-developed medial epicondyles have 30 31 more robust proportions, hunt by ambush, and inhabit highly-structured environments, such 32 33 as woodlands (Argot 2001, 2004; Salesa et al. 2008; Argot and Babot 2011). Thus, the 34 35 36 smaller development of the medial epicondyle of L. vallesiensis (Fig. 11a) (similar to those 37 38 of Le. serval and Ly. lynx) suggests that this feline was a capable runner, occupying the less 39 40 41 structured, more open patches of the mixed landscape inferred for the Batallones sites 42 43 (Antón and Morales 2000; Salesa et al. 2006a, 2008; Morales et al. 2008). The extant feline 44 45 46 F. silvestris, a species of similar size and proportions as L. vallesiensis, occupies a great 47 48 diversity of habitats, including grasslands and more or less dense woodlands (Nowak 2005; 49 50 Sunquist and Sunquist 2009), but it has a more developed medial epicondyle than the 51 52 53 Batallones feline and the Turolian P. attica. This implies that these two latter species had a 54 55 more limited range of preferred habitats than F. silvestris, favoring more open habitats than 56 57 58 their extant relative. On the contrary, the much more projected medial epicondyles of the 59 60 middle Miocene species S. turnauensis and M. lorteti suggest their preference for the 61 62 63 36 64 65 1 2 3 4 densely-wooded habitats that predominated during that time. Besides this, the presence of a 5 6 7 strongly projected medial epicondyle of the humerus has been also associated in Felidae to 8 9 the capture of relatively large prey (Meachen-Samuels and Van Valkenburgh 2009); 10 11 following this interpretation, L. vallesiensis would take small prey, whereas the middle 12 13 14 Miocene S. turnauensis and M. lorteti would prey upon relatively larger ones. 15 16 Other features observed in the humerus of L. vallesiensis also seem to be related to 17 18 19 its running abilities, such as the mediolaterally narrower distal epiphysis (Fig. 12), and 20 21 especially, the narrower olecranon fossa than those of F. silvestris, Ly. rufus, Lp. wiedii, 22 23 24 and Lp. geoffroyi. This feature groups L. vallesiensis with taxa showing more cursorial 25 26 postcranial skeletons, such as Ly. lynx, Ly. pardinus, C. caracal, and Le. serval, but also 27 28 29 with the Turolian P. attica, whose narrow olecranon fossa was described by Roussiakis 30 31 (2002). A narrower olecranon fossa prevents the lateral and medial torsion of the ulna in 32 33 mainly terrestrial primates and carnivorans when moving on the ground (Gonyea 1978; 34 35 36 Brundrett 2002; Argot and Babot 2011), hence the presence of this feature in the humerus 37 38 of S. vallesiensis is an additional evidence supporting an efficient terrestrial locomotion in 39 40 41 this species. 42 43 44 45 46 Radius 47 48 49 50 The radii of the compared sample of felines, included L. vallesiensis, are similar in 51 52 53 proportions and morphology. In fact, the most remarkable difference, the absence of notch 54 55 and capitular eminence on the proximal facet, is only shown by Le. serval and F. lybica, 56 57 58 without a clear functional explanation. Davis (1964) suggested that the capitular eminence 59 60 limits the rotatory mobility of the radius, but Gebo and Rose (1993) noticed its presence in 61 62 63 37 64 65 1 2 3 4 arboreal carnivorans (such as the procyonid Potos flavus), suggesting that this structure 5 6 7 could not be an impediment for the pronation-supination movements of the forearm when 8 9 climbing. Nevertheless, Heinrich and Rose (1997) suggested that the capitular eminence 10 11 might stabilize the radius when the elbow is flexed, and then its absence in terrestrial forms 12 13 14 such as Le. serval and F. lybica, which tend to maintain extended limb articulations, would 15 16 be more understandable. 17 18 19 The other difference observed within the studied sample of radii is the larger lateral 20 21 and medial development of the distal facet for the scapholunar, again seen only in Le. 22 23 24 serval and F. lybica. This mediolateral elongation of the facet produces a less rounded 25 26 facet, and thus a restriction in the rotational movements of the scapholunar on the radius, 27 28 29 and a predominance of the wrist flexion and extension actions. It is not surprising that the 30 31 two species that show this morphology are mostly terrestrial animals that move fast on the 32 33 ground, and it is not strange either that the primitive morphology, a more rounded facet, is 34 35 36 observed in L. vallesiensis, most of the compared felines, and G. genetta. 37 38 39 40 41 Ulna 42 43 44 45 46 The caudal border of the olecranon (attachment area for the long branch of the m. triceps 47 48 brachii) is clearly more inflated (proximally projected) in Ly. lynx, Ly. rufus, Ly. pardinus, 49 50 C. caracal, and Le. serval than in L. vallesiensis, F. silvestris, F. lybica, Lp. wiedii, and Lp. 51 52 53 geoffroyi (Fig. 13). In Viverridae, a greater development of this caudal vertex is indicative 54 55 of a relatively larger long branch of the m. triceps brachii (Taylor 1974). Thus, in the 56 57 58 lynxes and lynx-like felines, this muscle should be relatively larger than those of L. 59 60 vallesiensis, F. silvestris, F. lybica, Lp. wiedii, and Lp. geoffroyi. Among extant felids, the 61 62 63 38 64 65 1 2 3 4 cursorial species have relatively greater long branches of the m. triceps brachii than those 5 6 7 species with more robust proportions, as shown by Gambaryan (1974). Nevertheless, 8 9 although our observations seem to indicate a difference in the cursorial capacities of the 10 11 analyzed felines, the group showing more inflated caudal part of the olecranon is composed 12 13 14 by the largest taxa, and so the observed differences could be strongly influenced by 15 16 allometry. Anyway, the larger size of the long branch of the m. triceps brachii in Ly. lynx, 17 18 19 Ly. rufus, Ly. pardinus, C. caracal, and Le. serval is also suggested by the presence, in the 20 21 cranial part of the olecranon, of a marked groove for the passage of the tendon of this 22 23 24 muscle; this feature is typical of terrestrial viverrids, and is lacking in the most arboreal 25 26 forms (Taylor 1974; Feeney 1999). 27 28 29 The degree of proximal projection of the olecranon tuberosities seems to be closely 30 31 related to the development of the lateral and medial branches of the m. triceps brachii in 32 33 fossil and extant Felidae (Gonyea 1978; Salesa et al. 2010a, 2011). In L. vallesiensis, F. 34 35 36 silvestris, F. lybica, and Le. serval (Fig. 14a, d), the lateral tuberosity is much less 37 38 proximally projected than the medial one, a morphology also observed in canids and the 39 40 41 cursorial felid Acinonyx jubatus (Gonyea 1978); these two latter carnivorans have larger 42 43 medial branches of the m. triceps brachii than those species with larger lateral tuberosities 44 45 46 in the ulna (Gambaryan 1974; Gonyea 1978). Interestingly, the lateral tuberosity is more 47 48 proximally projected than the medial one in those felids inhabiting woodland habitats, and 49 50 they have significantly larger lateral branches of the m. triceps brachii than those felids that 51 52 53 occupy low-structured, open landscapes (Gambaryan 1974; Gonyea 1978). Nevertheless, 54 55 Gonyea (1978) also found that several species occupying both open and close environments 56 57 58 (such as C. caracal, Panthera leo, Ly. lynx, or Ly. rufus) had olecranon tuberosities of 59 60 similar size. All these observations made Gonyea (1978) suggest that the development of 61 62 63 39 64 65 1 2 3 4 the tuberosities was actually an indicator of the degree of deviation of the anterior limb 5 6 7 from the parasagittal plane during locomotion, with those felids having larger lateral 8 9 tuberosities exhibiting the greatest parasagittal deviation of the elbow during locomotion, 10 11 and those with larger medial tuberosities having little deviation of the forelimb from a 12 13 14 'pendulum-like' motion, and thus exhibiting high cursorial abilities, such as A. jubatus and 15 16 large canids. Felis silvestris, F. lybica and Le. serval also belong to this latter group, and all 17 18 19 have a mostly terrestrial way of life, climbing primarily as an escape mechanism (Sunquist 20 21 and Sunquist 2009; Julik et al. 2012). Considering its olecranon morphology, L. vallesiensis 22 23 24 would fit well within the group of mostly terrestrial and gracile felines, and probably its 25 26 lifestyle was not very different from that of the small species of the genus Felis. The same 27 28 29 morphology is observed in the middle Miocene, more primitive felines such as M. lorteti 30 31 (Salesa et al. 2011). Nevertheless, this pattern might be the plesiomorphic for Felidae, as it 32 33 is also present in the most primitive felid, Proailurus lemanensis, and the earliest members 34 35 36 of the saber-toothed subfamily, Pseudaelurus quadridentatus and Promegantereon ogygia 37 38 (Salesa et al. 2010a, 2011). 39 40 41 The attachment groove for the m. extensor digiti I et II is more excavated in L. 42 43 vallesiensis, F. lybica, F. silvestris, and Le. serval than in the other species. This muscle 44 45 46 extends the thumb and assists the m. extensor digitorum communis in extending the carpus 47 48 and the digit II (Davis 1964; Barone 2010; Julik et al. 2012). The attachment surface is 49 50 much shallower in Ly. lynx, Ly. pardinus, Ly. rufus, C. caracal, Lp. wiedii, and Lp. 51 52 53 geoffroyi, and also in large felids such as Puma concolor, Panthera pardus, or the cursorial 54 55 A. jubatus This similar morphology in felids with differing locomotor adaptations prevents 56 57 58 the proposal of any functional interpretation explaining the development of the muscle. In 59 60 fact, an ulna of M. lorteti from Sansan illustrated by Salesa et al. (2011) shows a similar 61 62 63 40 64 65 1 2 3 4 pattern to that of L. vallesiensis, and thus a marked groove for the m. extensor digiti I et II 5 6 7 might be the primitive condition for the Felinae. 8 9 10 11 Metacarpal I 12 13 14 15 16 The Mc I of L. vallesiensis, F. silvestris, and F. lybica are slenderer than those of the 17 18 19 smaller species Lp. wiedii and Lp. geoffroyi, the larger felines Le. serval, Ly. lynx, Ly. 20 21 rufus, and Ly. pardinus, and the middle Miocene species S. turnauensis and M. lorteti 22 23 24 (Peigné 2012; Salesa et al. 2012). The main function of the Mc I in Felidae is grasping 25 26 when climbing and hunting, thanks to the abduction of the thumb from the second finger 27 28 29 (Barone 1967; Gonyea and Ashworth 1975; Gonyea 1978; Akersten 1985; Taylor 1989; 30 31 Anyonge 1996; Turner and Antón 1997; Julik et al. 2012). As a result of its slenderness, the 32 33 thumb of L. vallesiensis, F. silvestris, and F. lybica would be able to withstand smaller 34 35 36 forces than those of the rest of the compared felines, which has interesting implications for 37 38 their climbing abilities and their ability to subdue prey. For example, if L. vallesiensis 39 40 41 preyed upon smaller animals that those killed by its middle Miocene counterparts (as the 42 43 morphology of its humerus suggests), it would not be necessary to have the robust Mc I 44 45 46 observed in the latter species. The extant felines F. silvestris and F. lybica hunt mostly 47 48 small prey (Smithers 1971; Kingdon 1977; Roberts 1977; Sharma 1979; Palmer and Fairall 49 50 1988; Sunquist and Sunquist 2009) using head shaking movements that kill them very fast 51 52 53 (Leyhausen 1973, 1979; Pellis and Officer 1987). Thus, both species exemplify the 54 55 ecological role that L. vallesiensis probably played in the late Miocene faunas of Europe. 56 57 58 On the contrary, the middle Miocene primitive saber-toothed felid Pseudaelurus 59 60 quadridentatus had a robust Mc I (Ginsburg 1961a; Salesa et al. 2010a; Peigné 2012), even 61 62 63 41 64 65 1 2 3 4 more than the extant large pantherin felines (Salesa et al. 2010a), in accordance with the 5 6 7 huge dewclaw and robust forelimbs present in Machairodontinae, traits developed for 8 9 achieving the rapid immobilization of prey (Gonyea 1976a; Emerson and Radinsky 1980; 10 11 Akersten 1985; Rawn-Schatzinger 1992; Turner and Antón 1997; Salesa et al. 2005, 2006a, 12 13 14 2010a, 2014). The small Mc I of L. vallesiensis would also help in reducing the weight of 15 16 the forelimb, and thus the energy employed when moving on the ground. 17 18 19 20 21 Metacarpals II-V 22 23 24 25 26 The Mc II-V of L. vallesiensis are very similar to those of the compared felines, with minor 27 28 29 differences mainly related to their robustness. In general, the metacarpals II-V of L. 30 31 vallesiensis are more robust than those of the most cursorial felines, such Ly. lynx and Le. 32 33 serval, but slenderer than those of the most arboreal and forest-dwelling felines Lp. wiedii 34 35 36 and Lp. geoffroyi. But, while this observation provides an overall view of the locomotor 37 38 abilities of the hand, there are some features that deserve a more detailed discussion, as 39 40 41 they seem primitive traits rather than a reflection of functional adaptations. For example, in 42 43 the Mc II, the dorsomedial facet for the trapezium is not proximally projected in L. 44 45 46 vallesiensis, F. lybica, and F. silvestris, whereas in the rest of the compared species it is 47 48 strongly projected. The viverrids C. civetta and G. genetta, with different locomotor 49 50 adaptations, have non-projected facets, thus probably illustrating the primitive morphology. 51 52 53 Another probable primitive feature observed in the Mc IV of L. vallesiensi is the palmar 54 55 facet for the Mc III, which is much less proximodistally expanded than in the compared 56 57 58 felines, showing the same morphology seen in the Mc IV of G. genetta. Interestingly, in the 59 60 terrestrial viverrid C. civetta, this palmar facet is larger, resembling those of the majority of 61 62 63 42 64 65 1 2 3 4 the compared felines. The presence of a large facet between Mc III and IV produces a more 5 6 7 stable and controlled articulation, an adaptation seen in several cursorial mammals 8 9 (Rasmussen and Simons 2000; Stevens et al. 2009), so its presence in most of the felines as 10 11 well as in terrestrial viverrids fits well with this idea, and would suggest a very primitive 12 13 14 state in L. vallesiensis, similar to arboreal viverrids. Finally, the facet for the attachment of 15 16 the muscle extensor carpi ulnaris, on the lateral face of the head of the McV, is less distally 17 18 19 expanded in L. vallesiensis and Le. serval than in F. silvestris, F. lybica, Ly. lynx, Ly. rufus, 20 21 Ly. pardinus, C. caracal, Lp. wiedii, and Lp. geoffroyi. This muscle, despite its name, is a 22 23 24 flexor of the carpus and an external rotator of the forearm, being developed from the lateral 25 26 epicondyle of the humerus to the base of the Mc V (Evans 1993; Barone 2010). Thus, the 27 28 29 distal expansion of the attachment surface on the Mc V observed in some species slightly 30 31 increases the length of the tendon, which given the actual size of the muscle, would suggest 32 33 a larger range of carpus flexion rather than a larger muscle. 34 35 36 37 38 Os Coxae 39 40 41 42 43 The absence of constriction in the ilium of L. vallesiensis, C. caracal, and Le. serval is a 44 45 46 consequence of the lesser dorsoventral expansion of the wing in relation to those species 47 48 with a more or less developed greater ischiatic notch and an expanded spina alaris (Ly. lynx, 49 50 Ly. pardinus, Ly. rufus, F. lybica, F. silvestris, Lp. wiedii, and Lp. geoffroyi) (Fig. 15). This 51 52 53 difference affects the attachment area of the m. sartorius, which in felids originates as a thin 54 55 strip on the ventral border of the iliac crest, inserting onto the medial surface of the patella, 56 57 58 the tibial crest, and the medial condyle of the tibia (Reighard and Jennings 1901; Barone 59 60 2010; Carlon and Hubbard 2012). Its main function is to adduct and rotate the femur, and to 61 62 63 43 64 65 1 2 3 4 extend the tibia (Reighard and Jennings 1901; Evans 1993; Barone 2010; Carlon and 5 6 7 Hubbard 2012). Thus, in L. vallesiensis, C. caracal, and Le. serval, this muscle would have 8 9 a more dorsal origin than in Ly. lynx, Ly. pardinus, Ly. rufus, F. lybica, F. silvestris, Lp. 10 11 wiedii, and Lp. geoffroyi. This configuration could imply a longer muscle, as its origin is 12 13 14 more dorsally located, but the difference, when comparing in detail the coxae of L. 15 16 vallesiensis and F. silvestris (very similar in size) seems really small. In any case, the more 17 18 19 dorsally located origin of the m. sartorius could improve the flexion of the femur during the 20 21 recovery phase of the locomotion, when the limb is preparing for foot-strike, and this bone 22 23 24 is carried cranially by the contraction of the m. iliopsoas and m. sartorius (Argot 2002). 25 26 Besides this, in L. vallesiensis, the attachment area of the m. sartorius is not a rough and 27 28 29 dorsally ridged area, such as those of Ly. lynx, Ly. pardinus, Ly. rufus, Le. serval, and C. 30 31 caracal, but a thin bony sheet along the ventrocranial border of the ilium wing. This 32 33 probably implies a relatively weaker m. sartorius, at least regarding its insertion on the 34 35 36 wing. Nevertheless, this seems to be related to size, given the differences between small 37 38 and large species within the compared sample, and the largest species would need a 39 40 41 relatively stronger m. sartorius. 42 43 Another interesting difference in the coxal morphology of L. vallesiensis and the 44 45 46 compared felines is the position of the ischiatic spine. In the Batallones feline, as well as in 47 48 Le. serval and C. caracal, this structure is more caudally located than in Ly. lynx, Ly. 49 50 pardinus, Ly. rufus, F. lybica, F. silvestris, Lp. wiedii, and Lp. geoffroyi (Fig. 15). In felids, 51 52 53 the ischiatic spine is the origin surface for the m. gemellus cranialis, whereas the m. 54 55 gemellus caudalis originates on the dorsal border of the ischium between the ischiatic spine 56 57 58 and the cranial border of the ischiatic tuberosity; both muscles, developed as two small 59 60 flattened fleshy masses, attach into the trochanteric fossa of the femur (Barone 2010; 61 62 63 44 64 65 1 2 3 4 Carlon and Hubbard 2012). Their function is to help the m. obturatorius internus in 5 6 7 abducting and rotating the femur (Reighard and Jennings 1901; Evans 1993; Barone 2010). 8 9 The more caudal position of the ischiatic spine in L. vallesiensis, Le. serval, and C. caracal 10 11 produces a shorter area for the attachment of the m. gemellus caudalis, and thus a smaller 12 13 14 muscle. But also, it implies relatively longer fibers in the m. gemellus cranialis in relation 15 16 to those of Ly. lynx, Ly. pardinus, Ly. rufus, F. lybica, F. silvestris, Lp. wiedii, and Lp. 17 18 19 geoffroyi. In fact, in these latter species, the origin and insertion areas of this muscle are 20 21 located in such a way that the fibers of the muscle are more or less perpendicular to the 22 23 24 craniocaudal axis of the os coxae. The relative caudal migration of the origin area in L. 25 26 vallesiensis, Le. serval, and C. caracal would make the fibers longer, as well as increasing 27 28 29 the range of rotation of the femur. Although both mm. gemelli are relatively small in felids 30 31 (Reighard and Jennings 1901), the described difference in their disposition might have an 32 33 important impact on their function as assistant to the m. obturatorius internus. In primitive 34 35 36 viverrids such as Genetta genetta, the ischiatic spine is as caudally located as in L. 37 38 vallesiensis, Le. serval, and C. caracal, and although this could point towards well- 39 40 41 developed arboreal capacities (because of the greater rotation range of the femur), it could 42 43 be also reflecting a retained, primitive state for this character. 44 45 46 The shape of the ischiatic arch clearly distinguishes two groups, one with more or 47 48 less straight caudal margin of the ischium (L. vallesiensis, F. lybica, Lp. wiedii, and Lp. 49 50 geoffroyi) and another one with a curved margin, developing a marked ischiatic arch (F. 51 52 53 silvestris, Ly. lynx, Ly. pardinus, Ly. rufus, Le. serval, and C. caracal). Once again, Genetta 54 55 genetta shares the pattern observed in the former group. Feeny (1999) related the presence 56 57 58 of ischiatic arch, with a reduction in the area for the origin of the m. biceps femoris, m. 59 60 semitendinosus, and m. semimembranosus, and an increase in the area for the origin of the 61 62 63 45 64 65 1 2 3 4 m. adductor femoris (=m. adductor magnus et brevis). This muscle adducts and extends the 5 6 7 coxofemoral articulation (Reighard and Jennings 1901; Evans 1993; Barone 2010). Argot 8 9 (2002) has noticed that the adductor muscles of the thigh are essential in stabilizing the foot 10 11 against the medially lying surface in arboreal forms using a wide range of femoral 12 13 14 abduction. This means that a strong development of these muscles would point towards 15 16 arboreal capacities, and that fits with the presence of a reduced ischiatic arch in G. genetta 17 18 19 as well as L. vallesiensis, F. lybica, Lp. wiedii, and Lp. geoffroyi. It is remarkable that Lp. 20 21 wiedii is one of the most arboreal felids (Alderton 1998), but the presence of a similar 22 23 24 morphology in L. vallesiensis could be indicating the retention of a primitive morphology 25 26 in a species that is not an ecological equivalent of the extant Lp. wiedii. 27 28 29 30 31 Sacrum 32 33 34 35 36 The sacrum of L. vallesiensis is quite similar to that of other felines. It shows some 37 38 differences, but they do not seem to have a relevant importance. An exception is the wider 39 40 41 and less caudally expanded notch separating the cranial articular surfaces and by the great 42 43 lateral projection of the lateral sacral crest, both features observed also in C. caracal and 44 45 46 Le. serval, although to a lesser degree (Fig. 16). The presence of this wider cranial notch in 47 48 the sacrum of these species suggests an also wider range of lateral movements between the 49 50 sacrum and the lumbar series, something that is evident when these bones are articulated 51 52 53 experimentally (personal observation). The sacrum of G. genetta shows a relatively 54 55 narrower notch than those of L. vallesiensis, C. caracal, and Le. serval (Fig. 16g), whereas 56 57 58 other arboreal carnivorans, such as Martes foina, exhibit a relatively wide notch (Fig. 16i), 59 60 with its cranial articular surfaces separated in a similar way to those of these former felines. 61 62 63 46 64 65 1 2 3 4 A cranial fragment of sacrum of M. lorteti from Sansan, described by Salesa et al. (2011) 5 6 7 strongly resembles L. vallesiensis in the development of the notch. Unfortunately, no 8 9 additional sacra of early and middle Miocene felines are known, so we lack data on the 10 11 primitive morphology of this group. It is interesting that both L. vallesiensis and M. lorteti 12 13 14 show the morphology of the sacral cranial articulation seen in an arboreal mustelid, instead 15 16 of resembling that of semiarboreal viverrids such as G. genetta. In the phylogeny of the 17 18 19 Carnivora, the families Viverridae and Felidae are closely related, whereas the Mustelidae 20 21 are included in another clade (Gaubert and Veron 2003; Gaubert and Cordeiro-Estrela 22 23 24 2006; Wolsan and Sato 2010). Thus, the similar sacral morphology observed in M. martes, 25 26 L. vallesiensis, and M. lorteti probably indicates a convergent adaptation for increasing the 27 28 29 lumbosacral mobility rather than a phylogenetic relationship. 30 31 Besides this, the sacrum of L. vallesiensis, C. caracal, and Le. serval shows a strong 32 33 lateral projection of the lateral sacral crest (Fig. 16b, d), a bony sheet serving as attachment 34 35 36 surface for the dorsal sacroiliac ligaments and the m. piriformis (Evans 1993; Ercoli et al. 37 38 2013). This might be related with the great lateral mobility of the sacrum of these three 39 40 41 species, inferred from the morphology of the cranial articular surface, which would need 42 43 relatively stronger ligaments stabilizing this region. 44 45 46 Some features of the sacrum of mammals can be associated to their tail length; for 47 48 example, Russo and Shapiro (2011) and Russo (2016) found that long-tailed species of 49 50 cercopithecoid primates and felids had significantly more circularly shaped caudal articular 51 52 53 surfaces, more acute sacrocaudal articulation angles, and more laterally expanded 54 55 transverse processes of the last sacral vertebra. Interestingly, the relative length of the 56 57 58 sacrum has not been found to be a good indicator of the tail length (Ankel 1965, 1972; 59 60 Ward et al. 1991; Nakatsukasa et al. 2004), although Ly. lynx, Ly. pardinus, Ly. rufus, and 61 62 63 47 64 65 1 2 3 4 C. caracal show relatively elongated and narrower sacra when comparing to those of L. 5 6 7 vallesiensis, Le. serval, F. silvestris, F. lybica, and Lp. geoffroyi. 8 9 The development of the intermediate sacral crest (bearing two mammillo-articular 10 11 processes) of L. vallesiensis is also observed in F. silvestris, F. lybica, and Lp. geoffroyi, 12 13 14 species with relatively long tails, whereas in Ly. lynx, Ly. pardinus, Ly. rufus, Le. serval, 15 16 and C. caracal, species with relatively short tails, the caudal process is absent. These 17 18 19 processes are part of the attachment areas for the mm. intertransversarii dorsales caudae, 20 21 one of the main caudal muscular groups (Evans 1993; Ercoli et al. 2013), and thus a 22 23 24 reduction of these structures would be expected when the tail is short. Given this, the 25 26 presence of well-developed mammillo-articular processes in extant species with long tails 27 28 29 would suggest the inclusion of L. vallesiensis in this group. Russo and Shapiro (2011) on 30 31 their study on primates, found that the development of the caudal transverse process of the 32 33 sacrum separated “long” and “tailless” groups from the “short”/“very short” groups, 34 35 36 although the study did not distinguish “short” and “very short” groups (Tojima 2013). In 37 38 the case of our compared sample of felines, there is no clear relationship between the 39 40 41 morphology of the caudal transverse process and the length of the tail, as for example Le. 42 43 serval shows an extremely thin and slender process, more developed than those of F. lybica 44 45 46 and Lp. geoffroyi, which have longer tails. Nevertheless, given the morphology of the other 47 48 discussed features of the sacrum, a relatively long tail can be inferred for L. vallesiensis. 49 50 51 52 53 Femur 54 55 56 57 58 The femur of L. vallesiensis fits very well within the feline pattern, although it shows some 59 60 differences when compared to those of other felines. The gluteal tuberosity, for instance, is 61 62 63 48 64 65 1 2 3 4 located, like in Lp. wiedii, clearly distal to the lesser trochanter, whereas in the rest of the 5 6 7 compared species of Felinae it is clearly proximal to this structure (Fig. 17). This 8 9 configuration relates to the relative length of the m. gluteus superficialis, which originates 10 11 on the iliac crest and sacrocaudal aponeurosis, and attaches, after partial fusion with the m. 12 13 14 gluteofemoralis, onto the gluteal tuberosity (Fisher et al. 2008; Barone 2010; Ercoli et al. 15 16 2013). Given these origin and insertion areas, the distal displacement of the gluteal 17 18 19 tuberosity in L. vallesiensis and Lp. wiedii would increase the relative length of an 20 21 important part of the fibers of the m. gluteus superficialis. The cranial fibers of the m. 22 23 24 gluteus superficialis flex and weakly abduct the hip joint, while the caudal ones weakly 25 26 extend and abduct the hip joint (Fisher et al. 2008). Basically, the pattern observed in L. 27 28 29 vallesiensis and Lp. wiedii could be reflecting a relatively longer m. gluteus superficialis in 30 31 relation to other felines, and its importance in the flexion and extension of the hind limb 32 33 during locomotion. The m. gluteus superficialis is relatively larger in those mammals that 34 35 36 need a powerful flexion of the hip joint (Endo et al. 2006), and thus, it is especially 37 38 developed in otters and other aquatic species (Gambaryan 1974; Endo et al. 2006). 39 40 41 Nevertheless, among the sample of large extant felids dissected by Gambaryan (1974), 42 43 there were no significant differences in the relative weight of the m. gluteus superficialis 44 45 46 with the exception of the jaguar (Panthera onca), which showed a relatively smaller muscle 47 48 (2.7% vs. more than 4.0% in the rest). Thus, the relatively longer m. gluteus superficialis of 49 50 L. vallesiensis and Lp. wiedii, rather than suggesting the presence of a heavier muscle 51 52 53 would indicate a capacity for rapid movements, as relatively longer muscles produces faster 54 55 contractions (Kardong 2002). 56 57 58 The position of the lesser trochanter also affects the function of other important 59 60 muscle, the m. iliopsoas, which basically flexes the coxofemoral articulation, drawing the 61 62 63 49 64 65 1 2 3 4 pelvic limb forward (Evans, 1993; Barone, 2010). Leptofelis vallesiensis, F. lybica, Ly. 5 6 7 lynx, Ly. pardinus, Ly. rufus, and Lp. geoffroyi show a lesser trochanter located close to the 8 9 medial margin of the femur, but laterally displaced in relation to its position in F. silvestris, 10 11 Le. serval, C. caracal, and Lp. wiedii (Fig. 17). As has been noted, a laterally-located lesser 12 13 14 trochanter produces lateral rotation as well as flexion at the coxofemoral articulation during 15 16 the contraction of the m. iliopsoas (Howell 1944; Munthe 1989). On the contrary, in those 17 18 19 carnivorans with more medially placed lesser trochanters, such as canids and hyaenids, the 20 21 pull of the m. iliopsoas is produced primarily in the sagittal plane (Munthe 1989). 22 23 24 According to this, F. silvestris, Le. serval, C. caracal, and Lp. wiedii would be favouring 25 26 those movements of the femur in the sagittal plane, whereas the others, including L. 27 28 29 vallesiensis, would be able of a higher rank of femur movements. Within the first group, F. 30 31 silvestris, Le. serval, and C. caracal are slender animals, with relatively long limbs and 32 33 terrestrial habits (Nowak 2005; Sunquist and Sunquist 2009), whereas Lp. wiedii has, on 34 35 36 the contrary, a more robust proportions and strong climbing abilities (Alderton 1998; 37 38 Sunquist and Sunquist 2009). Nevertheless, if the medially located lesser trochanter were 39 40 41 indicative of a m. iliopsoas acting close to the sagittal plane and producing strong jumping 42 43 impulse, the observed morphology would fit well with the ecology of F. silvestris, Le. 44 45 46 serval, C. caracal, and Lp. wiedii, all of them proficient jumpers (Alderton 1998; Sunquist 47 48 and Sunquist 2009). At this respect, it is remarkable that the semiarboreal viverrid Genetta 49 50 genetta, also a good jumper (Jennings and Veron 2009) shows a medially located lesser 51 52 53 trochanter, as F. silvestris, Le. serval, C. caracal, and Lp. wiedii. The feline from 54 55 Batallones would exhibit a morphology also observed in other extant felines, but derived 56 57 58 from that of the more primitive viverrids. The same pattern observed in L. vallesiensis is 59 60 present in the femora of S. turnauensis from La Grive-Saint-Alban (LGR 4042) and M. 61 62 63 50 64 65 1 2 3 4 lorteti from Sansan (SAN 879), thus indicating that this character derived as early in the 5 6 7 evolution of Felinae as the middle Miocene. 8 9 One of the few differences observed in the distal epiphysis of the femur is the 10 11 orientation of the medial condyle in caudal view, rather vertical in L. vallesiensis, F. 12 13 14 silvestris, Lp. wiedii, and Lp. geoffroyi, and slightly laterally inclined in Le. serval, F. 15 16 lybica, Ly. lynx, Ly. pardinus, Ly. rufus, and C. caracal. Also, in all these latter species, 17 18 19 except F. lybica, the medial condyle distally surpasses the level of the lateral one. When the 20 21 tibia flexes on the femur, both the inclination and the distal projection of the medial 22 23 24 condyle makes the tibia rotate laterally when it flexes on the femur, which all at once 25 26 makes the talus be placed in a more parasagittal position, and so the foot; on the contrary, 27 28 29 when both the lateral and medial femoral condyles reach the same level distally, the talus is 30 31 located in a more lateral position when the tibia flexes on the femur, which makes the foot 32 33 be rotated laterally. The position of the foot has an enormous importance when running, so 34 35 36 the observed differences would be separating more cursorial felids from those inhabiting 37 38 more closed environments, and thus retaining full climbing abilities. The morphology of 39 40 41 the femoral condyles in G. genetta, similar to those of L. vallesiensis strongly supports this 42 43 interpretation. 44 45 46 But maybe the most striking feature in the femur of L. vallesiensis is the 47 48 proximodistal development of the trochlea, greater than that of any other of the compared 49 50 felines (Fig. 18). Argot (2003) noticed that the femoral trochlea of canids was 51 52 53 proximodistally higher than those of other carnivorans including ursids, mustelids, and 54 55 felids, a feature that, following this author, reflected the great range of excursion of the 56 57 58 patella in cursorial forms. Previously, Howell (1944) had stated that jumping and cursorial 59 60 mammals required stronger extensor muscles of the tibia and have a well-developed 61 62 63 51 64 65 1 2 3 4 trochlea, with high marginal ridges. Arboreal viverrids such as G. genetta also show a 5 6 7 proximodistally high femoral trochlea, very similar to that of L. vallesiensis, which 8 9 probably indicates a good ability for jumping in both species. Nevertheless, the femoral 10 11 trochlea of the first known felid, the early Miocene Proailurus lemanensis from Saint 12 13 14 Gérand-le-Puy (MN 2), although being proximodistally higher than those of extant felines, 15 16 it is mediolaterally much wider than that of L. vallesiensis. If this is the primitive 17 18 19 morphology for Felidae, the pattern seen in extant felines had been achieved by reducing 20 21 the proximodistal height of the trochlea. The middle Miocene felines M. lorteti from 22 23 24 Sansan (MN 6) and S. turnauensis from La Grive-Saint-Alban (MN 7/8) (de Beaumont 25 26 1961; Salesa et al. 2011), both show a proximodistally shorter trochlea than that of L. 27 28 29 vallesiensis. Thus, these former species, older than the Batallones feline, already show a 30 31 modern femoral trochlea, very similar to that of extant felines. 32 33 34 35 36 Tibia 37 38 39 40 41 The tibia of L. vallesiensis shows a similar morphology and proportions to those of other 42 43 compared felines, except for the remarkable development of a strong bony ridge on the 44 45 46 caudal face of the diaphysis (Fig. 7c, 19a). This structure seems to separate the attachment 47 48 surfaces for the belly masses of the m. flexor digitorum lateralis and m. flexor digitorum 49 50 medialis + m. tibialis caudalis, and it could be helpful in increasing their area (as the 51 52 53 sagittal crest in the skull increases the attachment area for the m. temporalis), as well as 54 55 stabilizing the muscles by forming a groove to accommodate them. Although the main 56 57 58 function of both the m. flexor digitorum lateralis and m. flexor digitorum medialis is to flex 59 60 the foot phalanges (Evans 1993; Barone 2010) and, although the latter is an accessory of 61 62 63 52 64 65 1 2 3 4 the former, both have an important participation in the propulsion of the body (Barone 5 6 7 2010). If these two muscles were relatively larger in L. vallesiensis than in other felines, 8 9 they would have produced stronger propulsive forces in the hind limb. Among our studied 10 11 sample of felines, this ridge is also present in Lp. geoffroyi (Fig. 19b), but de Beaumont 12 13 14 (1986) described it also in Pristifelis attica and in the Asian golden cat (Catopuma 15 16 temminckii), which means that at least two species of extant felines that do not have either a 17 18 19 close relationship nor marked ecological differences (Ximenez 1975; Johnson and Franklin 20 21 1991; Mattern and McLennan 2000; Sunquist and Sunquist 2009) do retain this feature 22 23 24 exhibited by these two late Miocene felines. The sharing of this character by L. vallesiensis 25 26 and P. attica, and its absence in the middle Miocene Styriofelis turnauensis (Fig. 19c) and 27 28 29 Miopanthera lorteti poses an interesting question of the phylogenetic relationships of these 30 31 taxa (see Taxonomic Discussion section). The caudal crest is also present, although much 32 33 less marked, in ursids (Fig. 19e), which agrees with the great development in this group of 34 35 36 the attachment area for the m. flexor digitorum medialis + m. tibialis caudalis, something 37 38 that has been associated to plantigrady (Ginsburg 1961b). Nevertheless, when considering 39 40 41 its presence in some extant species of Felinae whose locomotion probably does not differ 42 43 from that of other species, the developed caudal crest of the tibia of L. vallesiensis cannot 44 45 46 be considered as an indication of plantigrady in this feline, moreover when the rest of the 47 48 hind limb elements are relatively slender. 49 50 51 52 53 Calcaneus 54 55 56 57 58 The dorsal orientation of the talar facet of the sustentaculum tali in L. vallesiensis, Lp. 59 60 wiedii, Lp. geoffroyii, F. silvestris, and F. lybica is an important feature (Fig. 20), as it 61 62 63 53 64 65 1 2 3 4 would theoretically imply a greater restriction in the lateral movements of the talus when 5 6 7 articulating with the calcaneus, an expected feature of the most cursorial species. 8 9 Nevertheless, when calcaneus and talus are experimentally articulated in our sample of 10 11 felines, there are not marked differences in the relative orientation of the talus, as in those 12 13 14 species having a dorsally oriented talar facet (L. vallesiensis, Lp. wiedii, Lp. geoffroyii, F. 15 16 silvestris, and F. lybica), the matching facet on the talus is less excavated than in felines 17 18 19 having a dorsomedially oriented talar facet in the sustentaculum tali (C. caracal, Le. serval, 20 21 Ly. lynx, Ly. rufus, and Ly. pardinus). The dorsal orientation of the facet is observed in both 22 23 24 terrestrial and arboreal small felines, as well as in L. vallesiensis, whereas the most 25 26 cursorial taxa, C. caracal, Le. serval, Ly. lynx, Ly. rufus, and Ly. pardinus, show a more 27 28 29 dorsomedial orientation. Interestingly, G. genetta shows a similar pattern to that observed 30 31 in the former group, as well as the primitive felids Proailurus lemanensis from Saint 32 33 Gerand-Le-Puy (SG 3532), Promegantereon ogygia from Batallones-1 (B/S-341) (Siliceo 34 35 36 et al. 2014), and the taxa from Sansan Pseudaelurus quadridentatus (Sa 2427) (Peigné 37 38 2012), S. turnauensis (Sa744) (Peigné 2012), and M. lorteti (Sa 671) (Salesa et al. 2011) 39 40 41 thus suggesting that this is the primitive condition for Felidae, which later derived in larger 42 43 felines. 44 45 46 The attachment area for the m. quadratus plantae shows its greatest size in L. 47 48 vallesiensis, Lp. geoffroyii, Lp. wiedii, and F. silvestris, suggesting the presence of a large 49 50 and fully operative muscle (Fig. 20e, h), whereas it would be reduced (given the 51 52 53 development of its attachment area) in F. lybica, C. caracal, Le. serval, Ly. lynx, Ly. rufus, 54 55 and Ly. pardinus (Fig. 20f, g). In several mammals, the m. quadratus plantae is a wide band 56 57 58 arising from the lateral surface of the calcaneus, and extending obliquely across the sole to 59 60 its insertion on the tendinous sheet of the mm. flexores digitorum profundi, formed by the 61 62 63 54 64 65 1 2 3 4 m. flexor digitorum medialis and m. flexor digitorum lateralis (Davis 1964; Barone 1967; 5 6 7 Ercoli et al. 2013), thus constituting an accessory of it. The m. quadratus plantae has two 8 9 main functions: adjusting the oblique pull of the mm. flexores digitorum profundi, and 10 11 flexing the foot without the involvement of this latter muscle (Reighard and Jennings 1901; 12 13 14 Barone 1967; Turner and Antón 1997; Vollmerhaus and Roos 2001; Benjamin et al. 2008). 15 16 Ginsburg (1961b) linked the great development of this muscle with plantigrady, but in the 17 18 19 case of felids, which are digitigrade carnivorans, the presence of a well-developed m. 20 21 quadratus plantare is clearly a primitive feature most likely related to the retention of the 22 23 24 grasping function of the hind limb (Turner and Antón 1997), and a reduced capacity for 25 26 pedal inversion but powerful flexion capability of the distal phalanges of the foot (Cifelli 27 28 29 1983). A well-developed depression for the attachment of the m. quadratus planate is found 30 31 in many fossil felids, such as P. lemanensis, Ps. quadridentatus, M. lorteti, Pr. ogygia, and 32 33 Metailurus major (Ginsburg 1961a; Salesa et al. 2011, 2012b; Siliceo et al. 2014) 34 35 36 indicative of a relative large muscle. Thus, these fossil species did probably exhibit a good 37 38 climbing ability, with the m. quadratus plantae muscle providing extra force and precision 39 40 41 to the flexion of the foot during this activity (Turner and Antón 1997). In more cursorial 42 43 felids, such as F. lybica, C. caracal, Le. serval, Ly. lynx, Ly. rufus, and Ly. pardinus, both 44 45 46 the necessity for decreasing the weight of the distal elements of the limbs and the restriction 47 48 of the lateral movements of the foot (Sooriakumaran and Sivananthan 2005) probably 49 50 explain the reduction in the size of the m. quadratus plantae (Ginsburg 1961b). Then, in 51 52 53 these cursorial felids, the mm. flexores digitorum profundi (m. flexor digitorum medialis 54 55 and m. flexor digitorum lateralis) became primarily flexors of the toes in relation to the 56 57 58 tibia, their function then changing from grasping to propulsion (Turner and Anton 1997). 59 60 61 62 63 55 64 65 1 2 3 4 Taxonomic discussion on “Styriofelis” vallesiensis 5 6 7 8 When Salesa et al. (2012a) assigned the small feline from Batallones-1 and Batallones-3 to 9 10 the genus Styriofelis as the new species S. vallesiensis, they chose the most conservative 11 12 13 taxonomic alternative based on a set of dental similarities between the Batallones sample 14 15 and the middle Miocene feline S. turnauensis, although their cladogram did not imply a 16 17 18 generic identity between both taxa. Nevertheless, the present new study reveals important 19 20 morphological differences in the postcranial skeleton of S. vallesiensis and S. turnauensis, 21 22 strongly suggesting a generic separation between these two Miocene felines. These features 23 24 25 are: 1) the humerus of S. turnauensis and M. lorteti has a less proximally projected greater 26 27 tubercle, and a much more medially projected medial epicondyle than those of S. 28 29 30 vallesiensis, both features indicating a strong difference in the locomotion of these felines, 31 32 with the two former species being much less cursorial than the feline from Batallones; 2) 33 34 35 the Mc I of S. vallesisensis is slenderer than those of S. turnauensis and M. lorteti; 3) both 36 37 S. turnauensis and M. lorteti have proximodistally shorter femoral trochleae than that of S. 38 39 40 vallesiensis, a feature that links the former species with the extant felines, whereas the 41 42 Batallones feline shows the same morphology observed in some extant viverrids; 4) in S. 43 44 turnauensis and M. lorteti, the attachment area of the m. quadratus plantae is much more 45 46 47 proximally extended than in S. vallesiensis, suggesting a much larger muscle, typical of 48 49 arboreal felids inhabiting vegetated ecosystems; and 5) the caudal face of the tibiae of P. 50 51 52 attica and S. vallesiensis shows a marked proximodistally developed ridge, a feature 53 54 completely absent in S. turnauensis and M. lorteti. 55 56 57 All these differences, besides suggesting a different lifestyle (with S. vallesiensis 58 59 being a more cursorial animal) support the separation of these felines in different genera. 60 61 62 63 56 64 65 1 2 3 4 Recently, Geraads and Peigné (2016) have discussed the status of the genus Styriofelis, 5 6 7 proposing the inclusion of the species lorteti in the genus Miopanthera, which would also 8 9 include the late Miocene Felis pamiri as M. pamiri. This increases the previously assumed 10 11 diversity of the felids from the middle Miocene, as at least three different taxa were present 12 13 14 at this time: the primitive machairodontine Pseudaelurus quadridentatus, the lynx-sized 15 16 feline Miopanthera lorteti, and the wild cat-sized feline Styriofelis turnauensis. Of these 17 18 19 three lineages, at least the first two had a continuity in the late Miocene, with 20 21 Promegantereon ogygia (a primitive saber-toothed felid very similar to Ps. quadridentatus) 22 23 24 and M. pamiri, which would be very close to the pantherin felid clade (Geraads and Peigné 25 26 2016). In both cases, several dental and skeletal similarities support a phylogenetic 27 28 29 relationship between those forms included in the same lineage (de Beaumont 1975; 30 31 Ginsburg 1983; Salesa et al. 2010b, 2012; Peigné 2012; Geraads and Peigné 2016), but the 32 33 assumption of S. vallesiensis as being part of the S. turnauensis lineage is quite doubtful, as 34 35 36 now it is clear that both species represent two different felid models characterized by a 37 38 combination of unique dental and skeletal features. Besides this, the other late Miocene 39 40 41 feline known in the European faunas, the larger P. attica shows a more derived dentition 42 43 than that of S. vallesiensis, which points towards a different generic status (Salesa et al. 44 45 46 2012). In the present study, we have shown the existence of shared features in the 47 48 postcranial skeleton of P. attica and S. vallesiensis (such as the proximodistally developed 49 50 ridge in the caudal face of the tibia), but there are also some others that distinguish both late 51 52 53 Miocene felines (such as the smaller projection of the lateral epicondyle of the humerus in 54 55 S. vallesiensis). In absence of a proper cladistic analysis (unapproachable due to the 56 57 58 scarcity of fossils), these similarities and differences are difficult to assess, but combining 59 60 the results of the phylogenetic tree proposed by Salesa et al. (2012) with the observations of 61 62 63 57 64 65 1 2 3 4 the present study, it seems quite probable that S. vallesiensis and P. attica belonged to the 5 6 7 same lineage of terrestrial felines, although representing two different genera. For example, 8 9 the presence of a P2 in P. attica, whereas S. vallesiensis retains two small upper deciduous 10 11 premolars, D1 and D2 (see discussion in Salesa et al. 2012) gives an idea of the marked 12 13 14 phylogenetic separation existing between both taxa. In consequence, a different generic 15 16 name should be proposed for S. vallesiensis in order to separate it from the primitive and 17 18 19 much more robust S. turnauensis, and from the contemporary and more derived P. attica. In 20 21 absence of any available name, we propose the new genus Leptofelis, with the following 22 23 24 diagnosis: 25 26 27 28 29 Order Carnivora Bowdich, 1821 30 31 Suborder Feliformia Kretzoi, 1945 32 33 Family Felidae Fischer, 1817 34 35 36 Subfamily Felinae Fischer, 1817 37 38 Genus Leptofelis, new genus 39 40 41 Leptofelis vallesiensis (Salesa, et al., 2012) 42 43 44 45 46 Holotype: BAT-3’09 1576, an almost complete skull from Batallones-3. 47 48 49 50 Type locality: Batallones-3 (late Miocene, Vallesian, MN 10) (Madrid, Spain). 51 52 53 54 55 Other localities: Maragheh (MN 11–12, Iran), and Batallones-1 (MN 10, Madrid). 56 57 58 59 60 61 62 63 58 64 65 1 2 3 4 Etymology: from the Greek λεπτός (leptós), meaning “slender”, and from the Latin felis, 5 6 7 cat. 8 9 10 11 Diagnosis: That provided by Salesa et al. (2012) and also, postcranial skeleton with a 12 13 14 combination of primitive and derived features: humerus with a well proximally projected 15 16 greater tubercle and an almost non-projected medial epicondyle; medial tubercle of the ulna 17 18 19 well developed, markedly proximally projected, and surpassing the level of the lateral 20 21 tubercle; slender Mc I, much less robust than those of middle Miocene felines such as S. 22 23 24 turnauensis and M. lorteti; mediopalmar facet of the Mc IV base relatively less 25 26 proximodistally expanded than in most felines; relatively short L7; dorsal sacral foramina 27 28 29 absent; relatively proximodistally elongated femoral trochlea; relatively reduced attachment 30 31 area for the m. quadratus plantae on the lateral face of the calcaneus; presence of a marked 32 33 proximodistally developed ridge on the caudal face of the tibia. 34 35 36 37 38 Differential diagnosis: Only postcranial features are considered. Compared to S. 39 40 41 turnauensis and M. lorteti, humerus with a more proximally projected greater tubercle, and 42 43 a much less medially projected medial epicondyle; Mc I slenderer; proximodistally longer 44 45 46 femoral trochlea; attachment area of the m. quadratus plantae much less proximally 47 48 extended; caudal face of the tibiae with a marked proximodistally developed ridge. 49 50 Compared to P. attica, lesser projection of the lateral epicondyle of the humerus. 51 52 53 54 55 Palaeobiology of Leptofelis vallesiensis 56 57 58 59 60 61 62 63 59 64 65 1 2 3 4 The postcranial anatomy of the late Miocene small feline Leptofelis vallesiensis is 5 6 7 characterized by a combination of derived and primitive characters, a good example of the 8 9 mosaicism typically exhibited by the intermediate forms of several groups of Carnivora. In 10 11 this case, whereas the forelimb of L. vallesiensis is relatively modern, resembling that of 12 13 14 some extant felines, its hind limb shows a set of primitive features in the pelvis, femur, and 15 16 calcaneus, some of them shared with arboreal extant viverrids, and absent in the living 17 18 19 felines. This unique combination of traits is different from that seen in the extant members 20 21 of Felinae, but also from that present in the middle Miocene earliest felines. 22 23 24 Leptofelis vallesiensis had a body mass of around 7–9 kg (see Appendix 1), that is, a 25 26 little larger than the extant F. silvestris and F. lybica, and particular proportions, as its 27 28 29 brachial index (BIn) was within the range of C. caracal and Lp. geoffroyi, its crural index 30 31 (CIn) is close to those of F. silvestris and Ly. lynx, and its intermembral index (IIn) is 32 33 similar to that of C. caracal and G. genetta (Table 3). These indexes provide information 34 35 36 on the locomotor adaptations (see Appendix 2), with the most cursorial species of 37 38 Carnivora having relatively longer radii and tibiae than the corresponding humerii and 39 40 41 femora (Taylor 1974, 1989; Gonyea 1978; Van Valkenburgh 1985, 1987; Anyonge 1996), 42 43 that is, they show BIn and CIn higher or close to 100. In the case of felids, the BIn has also 44 45 46 been related to habitat complexity, with those species inhabiting highly structured habitats, 47 48 with dense vegetational cover, having relatively shorter forearms (and thus BIn between 49 50 80–90) than those occupying low structured habitats (and showing BIn close to 100) 51 52 53 (Gonyea 1976a, b, 1978). Considering this, L. vallesiensis would have inhabited highly 54 55 structured habitats, which does not always mean dense-vegetated areas, but a landscape 56 57 58 composed of relatively open areas mixed with densely-vegetated patches (Fig. 21). This 59 60 kind of environments allows greater species diversity than less structured habitats, by 61 62 63 60 64 65 1 2 3 4 reducing predation efficiency and enabling predator/prey coexistence, and by increasing the 5 6 7 number of available niches (Menge and Sutherland 1976; Petren and Case 1998). This 8 9 would be an optimal habitat for a small feline such as L. vallesiensis, where it could find 10 11 both refuge and a high diversity of available prey. 12 13 14 On the other hand, the IIn provides information on the relative length of the limbs, 15 16 and in most felids, whose hind limb is longer than the forelimb, this index shows values 17 18 19 below 100. A lower IIn (between 70–80) indicates a particularly long hind limb (Fig. 21), 20 21 which has been related to jumping capacities (Howell 1944; Gonyea 1976b; Taylor 1976). 22 23 24 The IIn of L. vallesiensis is almost 81, which supports a well development of its jumping 25 26 capacities, as inferred from the morphology of its pelvic limb (see above). 27 28 29 In summary, L. vallesiensis was very likely a mostly terrestrial species, as shown by 30 31 the morphology and proportions of its forelimb, but retained both primitive knee and ankle, 32 33 associated with efficient jumping, an ability that was probably essential for eluding the 34 35 36 large carnivorans that inhabited Batallones, such as saber-toothed felids, amphicyonids, 37 38 ursids, ailurids, etc., all of them much larger than L. vallesiensis. 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Mammal Species 54: 1–4 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 84 64 65 1 2 3 4 Figure captions 5 6 7 8 9 Fig. 1. Measurements for each studied postcranial element. a, b, left humerus in (a) cranial 10 11 and (b) lateral views. c, lateral view of a right ulna. d, cranial view of right femur. e, lateral 12 13 14 view of a left coxal. f, dorsal view of a right calcaneus. g, dorsal view of a right talus. h, 15 16 ventral view of a sacrum. Abbreviations: ah, articular proximodistal height; aw, articular 17 18 19 mediolateral width; dw, distal mediolateral width; oh, olecranon proximodistal height; pl, 20 21 proximal craniocaudal length; pw, proximal mediolateral width; tl, total proximodistal 22 23 24 length. 25 26 27 28 29 Fig. 2. Caudal part of a lumbar region of Leptofelis vallesiensis from Batallones-1 (BAT- 30 31 1’06 D4-64a, BAT-1’06 D4-64b and BAT-1’06 D4-64c) in left lateral (a), and ventral (b) 32 33 views. From left to right, L7, L6, L5 and caudal fragment of L4. Scale bar = 20 mm. 34 35 36 37 38 Fig. 3. Long bones of Leptofelis vallesiensis from Batallones-1. a, b, BAT-1’02 D6-58, left 39 40 41 humerus in cranial (a), and lateral (b) views. c, d, B-2074 (5), right radius in cranial (c), 42 43 and caudal (d) views. e, f, Right ulna B-2074 (6) in lateral (e), and medial (f) views. Scale 44 45 46 bar = 20 mm. 47 48 49 50 Fig. 4. Metacarpals of Leptofelis vallesiensis from Batallones-1. a, b, BAT-1’03 D4-239, 51 52 53 right metacarpal I in dorsal (a), and medial (b) views. c, d, BAT-1’03 D5-162, right Mc II 54 55 in lateral (c), and medial (d) views. e, f, BAT-1’04 D4-184, right Mc III in lateral (e), and 56 57 58 medial (f) views. g, h, BAT-1’03 E4-164, right Mc IV in lateral (g), and medial (h) views. 59 60 i, j, BAT-1’04 E5-129, right Mc V in lateral (i), and medial (j) views. Scale bar = 20 mm. 61 62 63 85 64 65 1 2 3 4 5 6 7 Fig. 5. BAT-1’06 D4-66a, pelvis of Leptofelis vallesiensis from Batallones-1 in left lateral 8 9 (a) and ventral (b) views. Scale bar = 20 mm. 10 11 12 13 14 Fig. 6. BAT-1’06 D4-66b, sacrum of Leptofelis vallesiensis from Batallones-1 in dorsal (a) 15 16 and ventral (b) views. Scale bar = 20 mm. 17 18 19 20 21 Fig. 7. Long bones of Leptofelis vallesiensis from Batallones-1. a, b, BAT-1’06 D4-67a, 22 23 24 right femur of in cranial (a), and caudal (b) views. c–e, BAT-1’06 D4-110, left tibia in 25 26 caudal (c), medial (d), and lateral (e) views. Abbreviations: cr, caudal ridge. Scale bar = 20 27 28 29 mm. 30 31 32 33 Fig. 8. Tarsals of Leptofelis vallesiensis from Batallones-1. a, BAT-1’04 D6-173, right 34 35 36 talus in dorsal view (digitally separated from the articulated calcaneum BAT-1’04 D6-174). 37 38 b, BAT-1’03 D6-300, left talus in plantar view. c–e, BAT-1’06 F5-61, right calcaneus in 39 40 41 dorsal (c), medial (d), and plantar (e) views. Scale bar = 10 mm. 42 43 44 45 46 Fig. 9. Ventral view of articulated caudal part of the lumbar region (L7 to L4) of different 47 48 species of Carnivora shown at the same size for a better comparison. a, Genetta genetta. b, 49 50 Martes foina. c, Leptofelis vallesiensis from Batallones-1. d, Felis silvestris. e, Leopardus 51 52 53 geoffroyi. f, Leptailurus serval. 54 55 56 57 58 Fig. 10. Lateral view of the proximal epiphysis of the left humerus of different species of 59 60 Felidae, shown at the same size for a better comparison. a, BAT-1’02 D6-58, Leptofelis 61 62 63 86 64 65 1 2 3 4 vallesiensis from Batallones-1. b, Leopardus geoffroyi. c, Lynx lynx. Abbreviations: gt, 5 6 7 greater tubercle. 8 9 10 11 Fig. 11. Cranial view of the distal epiphysis of the left humerus of different species of 12 13 14 Felidae, shown at the same size for a better comparison. a, BAT-1’02 D6-58, Leptofelis 15 16 vallesiensis from Batallones-1. b, Leopardus geoffroyi. c, Lynx lynx. Abbreviations: me, 17 18 19 medial epicondyle. 20 21 22 23 24 Fig. 12. Caudal view of the distal epiphysis of the left humerus of different species of 25 26 Felidae, shown at the same size for a better comparison. a, BAT-1’02 D6-58, Leptofelis 27 28 29 vallesiensis from Batallones-1. b, Leopardus geoffroyi. c, Lynx lynx. Abbreviations: le, 30 31 lateral epicondyle. 32 33 34 35 36 Fig. 13. Lateral view of the proximal epiphysis of the right ulna of different species of 37 38 Felidae, shown at the same size for a better comparison. a, B-2074 (6), Leptofelis 39 40 41 vallesiensis from Batallones-1. b, Leopardus geoffroyi. c, Lynx lynx. d, Felis lybica. e, 42 43 Leopardus wiedii. f, Leptailurus serval. Abbreviations: ltb, attachment area for the lateral 44 45 46 branch of m. triceps brachii. 47 48 49 50 Fig. 14. Cranial view of the proximal epiphysis of the right ulna of different species of 51 52 53 Felidae, shown at the same size for a better comparison. a, BAT-1’06 E5-50, Leptofelis 54 55 vallesiensis from Batallones-1. b, Leopardus geoffroyi. c, Lynx lynx. d, Leptailurus serval. 56 57 58 Abbreviations: lt, lateral tuberosity; mt, medial tuberosity. 59 60 61 62 63 87 64 65 1 2 3 4 Fig. 15. Lateral view of the left coxae of different species of Felidae, shown at the same 5 6 7 size for a better comparison. a, BAT-1’06 D4-66a, Leptofelis vallesiensis from Batallones- 8 9 1. b, Caracal caracal. c, Lynx pardinus. Abbreviations: gsn, greater ischiatic notch; sa, 10 11 spina alaris; is, ischiatic spine. 12 13 14 15 16 Fig. 16. Sacra of different species of Felidae and Viverridae, shown at the same size for a 17 18 19 better comparison. a, b, BAT-1’06 D4-66b, Leptofelis vallesiensis from Batallones-1 in 20 21 cranial (a) and ventral (b) views. c, d, Caracal caracal in cranial (c) and ventral (d) views. 22 23 24 e, f, Felis silvestris in cranial (e) and ventral (f) views. g, h, Genetta genetta in cranial (g) 25 26 and ventral (h) views. i, j, Martes foina in cranial (i) and ventral (j) views. Abbreviations: 27 28 29 cas, cranial articular surface; lsc, lateral sacral crest. 30 31 32 33 Fig. 17. Caudal view of the right femur of different species of Felidae, shown at the same 34 35 36 size for a better comparison. a, BAT-1’06 D4-67a, Leptofelis vallesiensis from Batallones- 37 38 1. b, Felis silvestris. c, Caracal caracal. d, Leopardus geoffroyi. e, Lynx lynx. 39 40 41 Abbreviations: gt, gluteal tuberosity; lt, lesser trochanter. 42 43 44 45 46 Fig. 18. Cranial view of the right femur of different species of Felidae and Viverridae, 47 48 shown at the same size for a better comparison. a, BAT-1’06 D4-67a, Leptofelis 49 50 vallesiensis from Batallones-1. b, Felis silvestris. c, Caracal caracal. d, Genetta genetta. 51 52 53 Arrows indicate the proximal margin of the throclea. 54 55 56 57 58 Fig. 19. Caudal view of the left tibia of different species of Felidae. a, BAT-1’06 D4-110, 59 60 Leptofelis vallesiensis from Batallones-1. b, Leopardus geoffroyi. c, SNSB-BSPG 1937 II 61 62 63 88 64 65 1 2 3 4 12806, Styriofelis turnauensis from Wintershof-West. d, Genetta genetta. e, Ursus 5 6 7 americanus. Abbreviations: cr, caudal ridge. Scale bar 1 (for A–D) = 20 mm Scale bar 2 8 9 (for E) = 40 mm. 10 11 12 13 14 Fig. 20. Calcanei of different species of Felidae, shown at the same size for a better 15 16 comparison. a, e, BAT-1’06 F5-61, Leptofelis vallesiensis from Batallones-1 in medial (a) 17 18 19 and lateral (e) views. b, f, Lynx lynx in medial (b) and lateral (f) views. c, g, Caracal 20 21 caracal in medial (c) and ventral (f) views. d, h, Leopardus wiedii in medial (d) and lateral 22 23 24 (h) views. Abbreviations: fst, talar facet of the sustentaculum tali; gqp, groove for the m. 25 26 quadratus plantae (arrows indicate the proximal margin of this groove). 27 28 29 30 31 Fig. 21. Skeletal (a) and life reconstruction (b) of Leptofelis vallesiensis, based on the 32 33 specimens from Batallones-1 (scale bar = 10 cm). The shown hypothetical coat colour 34 35 36 pattern is congruent with the phylogenetic inferences about coat pattern evolution in felids 37 38 proposed by Werdelin et al. (2010) (artwork by M. Antón). 39 40 41 42 43 Table captions 44 45 46 47 48 Table 1. Measurements of the postcranial bones of L. vallesiensis from Batallones-1. 49 50 51 52 53 Table 2. Measurements of the postcranial bones of L. vallesiensis from Batallones-1. 54 55 56 57 58 Table 3. Brachial, Crural, and Intermembral indices of L. vallesiensis and several species 59 60 of Felidae and Viverridae. Data taken from Salesa (2002). 61 62 63 89 64 65 1 2 3 4 APPENDIX 1 5 6 7 Body mass estimation in Leptofelis vallesiensis 8 9 10 11 For the estimation of the body mass of L. vallesiensis we used the specific formulae for the 12 13 14 family Felidae of Rodríguez (1997) and Anyonge (1993), both based on the following 15 16 measurements of the appendicular skeleton: proximal width of the tibia (PW) and width of 17 18 19 the femoral head (FHW) (Rodríguez, 1997), and distal articular (condylar) area of femur 20 21 (FDA) (Anyonge, 1993). The results are: 22 23 24 For the proximal width of the tibia (PW), the formula is: Log Body Mass = 2.8581 25 26 (log PW) - 0.0786. The PW of the tibia BAT-1’06 D4-110 is 25.32 mm, which results in a 27 28 29 body mass of 8.5 kg. 30 31 For the width of the femoral head (FHW), the formula is: Log Body Mass = 2.9968 32 33 (log FHW) + 0.5668. The FHW of the femur BAT-1’06 D4-67a is 12.54 mm, which results 34 35 36 in a body mass of 7.21 kg. 37 38 For the femoral distal articular (condylar) area (FDA) the formula is: Log Body 39 40 41 Mass = 1.32 (log FDA) – 2.16. The FDA was calculated following the methodology of 42 43 Anyonge (1993) for the femora BAT-1’06 D4-67a and BAT-1’06 D4-67b, obtaining areas 44 45 2 2 46 of 222.63 mm and 229.168 mm , respectively. This results in a body mass of 8.68 kg from 47 48 the femur BAT-1’06 D4-67a, and 9.02 kg from the specimen BAT-1’06 D4-67b. 49 50 Thus, the estimated body mass of L. vallesiensis from Batallones-1 is between 7.21 51 52 53 and 9.02 kg. The published data on the body weight of extant felines shows wider ranges, 54 55 as they take into account the size variation within each species, mostly due to sexual 56 57 58 dimorphism. Anyway, the estimated body weight of L. vallesiensis is close to the maximum 59 60 61 62 63 90 64 65 1 2 3 4 weights of several species of extant small felines such as Felis silvestris, F. chaus, 5 6 7 Leopardus geoffroyi and Puma yaguarondi (Sunquist and Sunquist, 2009). 8 9 10 11 APPENDIX 2 12 13 14 Body proportions in Leptofelis vallesiensis 15 16 17 18 19 The body proportions of the feline from BAT-1 have been calculated using the brachial, 20 21 crural and intermembral indexes (Gonyea, 1976a, 1976b; Taylor, 1976; Anyonge, 1996; 22 23 24 Iwaniuk et al., 1999). The formulae for these indexes are: 25 26 Brachial index (BIn) = (Length radius/Length humerus) x 100. 27 28 29 Crural index (CIn) = (Length tibia/ Length femur) x 100. 30 31 Intermembral index (IIn) = (Length thoracic limb/ Length pelvic limb) x 100. 32 33 The corresponding measurement were taken on the following bones of L. 34 35 36 vallesiensis from Batallones-1: Femora (BAT-1’06 D4-67a and BAT-1’06 D4-67b), Tibiae 37 38 (BAT-1’06 D4-110 and BAT-1’06 D4-103), Humeri (BAT-1’02 D6–58 and B-4968) and 39 40 41 Radii (B/S-575 and B-2074(5)). 42 43 The results of the different indexes for L. vallesiensis are as follows: BIn: 89.16– 44 45 46 91.13; CIn: 103.01–104.04; IIn: 80.23–80.93. 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 91 64 65 Figure 1 Click here to download Figure Fig1.tif Figure 2 Click here to download Figure Fig2.tif Figure 3 Click here to download Figure Fig3.tif Figure 4 Click here to download Figure Fig4.tif Figure 5 Click here to download Figure Fig5.tif Figure 6 Click here to download Figure Fig6.tif Figure 7 Click here to download Figure Fig7.tif Figure 8 Click here to download Figure Fig8.tif Figure 9 Click here to download Figure Fig9.tif Figure 10 Click here to download Figure Fig10.tif Figure 11 Click here to download Figure Fig11.tif Figure 12 Click here to download Figure Fig12.tif Figure 13 Click here to download Figure Fig13.tif Figure 14 Click here to download Figure Fig14.tif Figure 15 Click here to download Figure Fig15.tif Figure 16 Click here to download Figure Fig16.tif Figure 17 Click here to download Figure Fig17.tif Figure 18 Click here to download Figure Fig18.tif Figure 19 Click here to download Figure Fig19.tif Figure 20 Click here to download Figure Fig20.tif Figure 21 Click here to download Figure Fig21.tif Table 1 Click here to download Table Table 1.docx

Table 1. Measurements of the postcranial bones of L. vallesiensis from Batallones-1.

Element Number tl pw dw pl Right femur BAT-1’06 D4-67a 142.13 26.80 24.73 Left femur BAT-1’06 D4-67b 141.04 24.88 23.33 Right tibia BAT-1’06 D4-103 147.28* 25.25 18.21 25.10 Left tibia BAT-1’06 D4-110 144.86 25.30 18.32 22.84* Left radius B/S-575 111.84 9.62 13.60 Right radius B-2074s 108.32 9.72 13.56 Left humerus BAT-1’02 D6-58 122.69 17.30 19.72 24.12 Right humerus B-4968 123.44* 18.00 19.46 23.84 Left coxal BAT-1’06 D4-66a 99.36 - - Right Mc I BAT-1’03 D4-239 13.58 5.14 4.00 Right Mc II BAT-1’03 D5-162 32.31 4.85 5.57 Right Mc III BAT-1’04 D4-184 37.71 6.35 5.82 Right Mc IV BAT-1’03 E4-164 36.37 5.18 5.41 Right Mc V BAT-1’04 E5-129 29.24 5.18 5.58 Left Mc V BAT-1’04 E5-212 29.01 5.17 5.63

Table 2 Click here to download Table Table 2.docx

Table 2. Measurements of the postcranial bones of L. vallesiensis from Batallones-1.

Element Number tl oh ah aw Right ulna BAT-1’08 C8-24 131.06 15.02 16.67 - Right ulna BAT-1’06 E5-50 123.73 13.06 14.80 - Right ulna B-2074(6) 128.12 13.44* 14.84 - Sacrum BAT-1’06 D4-66b 33.94 - - - Left talus BAT-1’03 D6-300 19.42 - 12.46 13.11 Right calcaneus BAT-1’06 F5-61 33.19 - 16.36 14.18 Left calcaneus BAT-1’04 D6-62 31.34 - 15.68 14.93 Left calcaneus BAT-1’03 D6-301 34.40 - 16.10 14.39

Table 3 Click here to download Table Table 3.docx

Table 3. Brachial, Crural and Intermembral indexes of L. vallesiensis and several

species of Felidae and Viverridae. Data taken from Salesa (2002).

Species BIn CIn IIn

Felis lybica 98.75 95.35 85.08 Caracal caracal 91.34 95.45 80.73 Leopardus pardalis 88.39 95.50 85.14 Puma yaguarondi 81.86 95.58 77.33 Profelis aurata 86.11 95.97 80.31 Prionailurus viverrinus 89.79 96.97 82.92 Lynx pardinus 96.31 99.72 82.11 Leopardus wiedii 85.48 100.55 81.05 Leopardus tigrinus 84.50 100.49 81.07 Leopardus geoffroyi 89.34 100.09 84.73 Prionailurus planiceps 88.87 101.69 81.97 Pardofelis marmorata 83.19 102.69 80.65 Prionailurus bengalensis 86.90 102.77 83.49 Lynx lynx 97.89 103.30 82.75 Felis silvestris 94.59 104.81 86.01 Leptailurus serval 95.78 101.87 87.38 Felis chaus 97.66 111.72 93.36 Catopuma temminckii 87.40 118.42 73.95 Neofelis nebulosa 81.92 99.21 82.47 Puma concolor 83.90 93.72 82.70 Uncia uncia 91.41 103.12 84.93 Panthera pardus 82.24 92.63 84.95 Panthera onca 80.80 86.61 87.71 Acinonyx jubatus 103.30 105.00 90.10 Panthera leo 95.35 88.77 91.91 Genetta genetta 85.77 106.05 80.55 Civettictis civetta 88.64 96.44 81.88 Leptofelis vallesiensis 89.16–91.13 103.01–104.04 80.23–80.93