Journal of Vertebrate Paleontology Promegantereon Ogygia (Felidae, Machairodontinae, Smilodontini) from the Vallesian (Late Mioc

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Journal of Vertebrate Paleontology Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/ujvp20 Promegantereon ogygia (Felidae, Machairodontinae, Smilodontini) from the Vallesian (late Miocene, MN 10) of Spain: morphological and functional differences in two noncontemporary populations Gema Siliceo a , Manuel J. Salesa a , Mauricio Antón a , Marcos F. G. Monescillo a & Jorge Morales a a Departamento de Paleobiología , Museo Nacional de Ciencias Naturales–CSIC C/José Gutiérrez Abascal , 2.28006 , Madrid , Spain Published online: 04 Mar 2014.

To cite this article: Gema Siliceo , Manuel J. Salesa , Mauricio Antón , Marcos F. G. Monescillo & Jorge Morales (2014) Promegantereon ogygia (Felidae, Machairodontinae, Smilodontini) from the Vallesian (late Miocene, MN 10) of Spain: morphological and functional differences in two noncontemporary populations, Journal of Vertebrate Paleontology, 34:2, 407-418 To link to this article: http://dx.doi.org/10.1080/02724634.2013.812099

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PROMEGANTEREON OGYGIA (FELIDAE, MACHAIRODONTINAE, SMILODONTINI) FROM THE VALLESIAN (LATE MIOCENE, MN 10) OF SPAIN: MORPHOLOGICAL AND FUNCTIONAL DIFFERENCES IN TWO NONCONTEMPORARY POPULATIONS

GEMA SILICEO, MANUEL J. SALESA,* MAURICIO ANTON,´ MARCOS F. G. MONESCILLO, and JORGE MORALES Departamento de Paleobiologıa,´ Museo Nacional de Ciencias Naturales–CSIC C/Jose´ Gutierrez´ Abascal, 2. 28006 Madrid, Spain, [email protected]; [email protected]; [email protected]; [email protected]; [email protected]

ABSTRACT—We compare two populations of the primitive saber-toothed felid Promegantereon ogygia from the late Miocene (Vallesian, MN 10) of Spain. These populations come from two fossil sites, Batallones-1 and Batallones-3, located very close to each other, within the Cerro de los Batallones complex. The sites show differences in age and in their faunal assemblages, with Batallones-1 being older than Batallones-3. We ﬁnd that the population from this latter site shows slightly derived characters in both dentition and postcranial elements, which clearly indicate evolution within the Promegantereon lineage, but are not strong enough to support a separation at the species level.

INTRODUCTION of excavation it was clear that, like the Batallones-1 locality, its fauna was also composed mainly of carnivorans. Nevertheless, The Cerro de los Batallones is a low hill (elevation 700 m) both Batallones-1 and Batallones-3, although separated by less located 30 km south of the city of Madrid (Spain), between the than 180 m, constitute two different cavities, showing marked dif- villages of Valdemoro and Torrejon´ de Velasco (Fig. 1). Due to ferences in the faunal assemblages, mainly due to the absence the high quality of its sepiolite levels, this hill has been exploited in Batallones-3 of the ailurid Simocyon batalleri, and the pres- as an opencast mine of this mineral since 1974. In November ence of the bear Indarctos arctoides, the giant mustelid Eomel- 1991, during mining work, the mechanical diggers discovered a livora piveteaui, and the derived amphicyonid Thaumastocyon rich accumulation of carnivoran fossils within a lens of green- sp., all absent in the Batallones-1 sample (Salesa et al., 2010b, ish clay intercalated between the levels of sepiolite. The site, 2012; Abella, 2011; Valenciano et al., 2012). Besides these taxa, named Batallones-1, was formed during the late Miocene as an a rich sample of the saber-toothed felids Machairodus aphanis- irregular cavity in the sepiolite levels, at least 12 m deep, later tus and Promegantereon ogygia has been recovered, although filled with greenish clay. It acted as a natural trap for many ani- there are differences in their relative abundance: P. ogygia is the mals, likely due to the fact that when the sepiolite is wet, its sur- most abundant species in Batallones-1 (Anton´ and Morales, 2000; face becomes slippery, which would make escape from the trap Salesa, 2002), followed by M. aphanistus, whereas in Batallones- very difficult. Of all the macromammal bones recovered from 3 M. aphanistus and I. arctoides are more abundant than P. ogy- the site, 98% correspond to members of the order Carnivora, gia (Abella, 2011; Siliceo et al., 2011). Thus, the relative abun- which were probably trapped while attempting to scavenge. This dance of these two large felids are reversed in Batallones-1 and carnivoran assemblage includes well-preserved remains of the Batallones-3, which could imply an actual difference in their pop- ailurid Simocyon batalleri, the amphicyonid Magericyon anceps, ulation densities in both communities, and thus some kind of pa- three species of mustelids, the primitive hyaenid Protictitherium leoecological difference. Recent studies on the dentition of the crassum, the machairodont felids Machairodus aphanistus and cricetid Hispanomys moralesi from three of the Batallones locali- Promegantereon ogygia, and two species of felines, Styriofelis ties (Lopez-Anto´ nanzas˜ et al., 2010) have shown that the popula- vallesiensis and an undetermined, larger species. This impressive tion of this cricetid from Batallones-3 shows a more derived mor- Downloaded by [CSIC Instituto Quimica Fiscia Rocasolano] at 07:36 04 March 2014 sample has allowed several systematic, paleoecological, and func- phology than that from Batallones-1, establishing that the most tional studies (Anton´ and Morales, 2000; Morales et al., 2000, primitive dentition should correspond to the older locality, and 2004; Anton´ et al., 2004; Peigne´ et al., 2005, 2008; Salesa et al., thus proposing a younger age for Batallones-3. Unfortunately, 2005, 2006a, 2006b, 2008, 2010a, 2010b, 2012). based on morphological data, only a relative dating can be pro- Up to now, nine cavities have been identified in the area of posed, the exact chronological separation between both locali- Cerro de los Batallones, all of them responding to a similar geo- ties being impossible to assess. This difference in age between logical process of piping, which consisted in the erosion of the both localities would explain the marked differences in faunal sepiolite levels by water flowing along fractures, causing col- composition. lapses and the development of a karst-like (‘pseudokarst’) topog- The primitive Smilodontini Promegantereon ogygia was a raphy (Pozo et al., 2004). Although the localities of Batallones- poorly known fossil felid until the discovery of Batallones-1 in 2, Batallones-4, and Batallones-5 have yielded some remains of 1991, which yielded a high number of well-preserved cranial and carnivorans, only Batallones-1 and Batallones-3 can be consid- postcranial elements of this species. With these findings, several ered as carnivore traps, due to the predominance of carnivoran aspects of its anatomy, taxonomy, and palaeobiology were as- fossils. Batallones-3 was found in 2000, and since the beginning sessed (Salesa et al., 2005, 2006b, 2010a, 2010b), although all these studies were based on a single population. With the new fossils from Batallones-3, it is now possible, for the first time, *Corresponding author. to accomplish a comparative functional and morphological study

407 408 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 34, NO. 2, 2014

FIGURE 1. A, geographic location of the paleontological area of Cerro de los Batal- lones within Spain and the Madrid Region; B, schematic geological map of Cerro de los Batal- lones showing the location of the paleontological sites discussed in the text.

between two populations of this saber-toothed felid, which con- de la Universidad de Valladolid (Spain) and Museo Nacional de stitutes the primary goal of the present contribution. Ciencias Naturales–CSIC (Madrid, Spain), which provided complete skeletons of the felines Panthera leo, Panthera tigris, Pan- thera pardus, Panthera onca, Puma concolor, Acinonyx jubatus, and Lynx pardinus. Comparisons with the primitive felids Proail- MATERIALS AND METHODS urus lemanensis Filhol, 1879, from the Upper Oligocene of Saint- Material Gerand-Le-Puy fossil site, and Pseudaelurus quadridentatus from Sansan, were made using the collections of the Museum National The fossils of Promegantereon ogygia from the fossil sites of Downloaded by [CSIC Instituto Quimica Fiscia Rocasolano] at 07:36 04 March 2014 d’Histoire naturelle (Paris). Comparisons with other fossil Feli- Batallones-1 and Batallones-3 studied here are housed at the dae, such as Panthera atrox, Smilodon fatalis (Merriam and Stock, paleontological collections of the Museo Nacional de Ciencias 1932), Smilodon gracilis (Berta, 1987), and Megantereon cultrides Naturales–CSIC, Madrid (Spain). Labeled on each specimen is (Turner, 1987; Christiansen and Adolfssen, 2007), were made us- the acronym of Batallones-3 (indicated as ‘BAT-3’) followed ing published data. by the abbreviation of the year of excavation, and the catalog number of the fossil (that may include a letter); in some cases an ‘s’ may be present before the number of fossil, indicating that the specimen was not found in situ, but in the top sedi- Methods ments of Batallones-3 removed during the mining, e.g., BAT- 3’08 s-45A. At least 10 individuals of Promegantereon ogygia are The anatomical descriptions follow the terminology used by represented in Batallones-3 (based on the number of right cal- Barone (2010) and the Nomina Anatomica Veterinaria (2005). canei). The fossils of P. ogygia from Batallones-3 analyzed in this The measurements were taken using digital calipers and are study have been compared both biometrically and morphologi- shown in Figure 2 and Tables 1–5. Quantitative data were ana- cally with the extensive sample of P. ogygia from Batallones-1, lyzed using the Student’s t-test from the statistics software pack- which has been previously studied by Salesa (2002) and Salesa age IBM SPSS Statistics 19.0. Results for the statistically signiﬁ- et al. (2005, 2006b, 2010a, 2010b). Comparisons with extant car- cant differences are shown in Table 6. Only those elements show- nivores were made using the collections of the Museo Anatomico´ ing these differences are included in the discussion. SILICEO ET AL.—PROMEGANTEREON OGYGIA FROM BATALLONES SITES 409

FIGURE 2. Measurements for each studied cranial and postcranial element. A, ventral view of the skull; B, buccal view of the hemimandible; C, buccal view of the m1; D, occlusal view of the P4; E, dorsal view of the calcaneus; F, distal view of the calcaneus; G, plantar view of the talus; H, dorsal view of the talus; I, dorsal view of an idealized metapodial; J, lateral/medial view of an idealized metapodial (modiﬁed from Merriam and Stock, 1932).

Institutional Abbreviations—BAT-1, B, Batallones-1; BAT-3, 344, BAT-3’11 2392, BAT-3’11 1624, and BAT-3’11 1188, com- Batallones-3; B/S, Batallones-1, fossil not found in situ; MNCN, plete left upper canines; BAT-3’07 835 and BAT-3’10 623, incom- Museo Nacional de Ciencias Naturales–CSIC. plete left upper canines; BAT-3’10 2037, skull and mandible with Anatomical Abbreviations—C, upper canine; c, lower canine; observable right C–P4; BAT-3’06 591, skull with right and left I3, I, upper incisor; i, lower incisor; M, upper molar; m, lower mo- P3–P4; BAT-3’06 519, skull with left P3–P4 and right C–P4; BAT- lar; Mc, metacarpal; Mt, metatarsal; P, upper premolar; p,lower 3’07 207, skull with left and right P3–M1; BAT-3’09 1636, skull premolar. with left P3–M1 and right P3–P4; BAT-3’10 1773, skull with left Measurement Abbreviations—bl, skull basal length; blw, buc- and right P3–M1; BAT-3’07 1066, skull with left C–M1; BAT-3’01 colingual width; ch, crown height; del, distal epiphysis length; 60A, fragment of skull with left and right P3–M1; BAT-3’08 833, dew, distal epiphysis width; dfw, mediolateral width of the dis- right lower canines; BAT-3’11 1964, BAT-3’11 1937, and BAT- tal facet for the cuboid; dpl, dorsopalmar length in the center of 3’10 689, left lower canines; BAT-3’11 2020, complete mandible the metapodial; dsl, dental series length; mcl, mandibular length with right and left ramus with c–m1; BAT-3’10 1773, mandible from the mesial border of the incisors to the caudal border of the with complete left ramus with c–m1 and right ramus, whose coro- coronoid process; mdl, mesiodistal length; mh, maximum height noid process is fragmented, with c–m1; BAT-3’11 132, complete of the talus; ml, maximum mandibular length; mlw, mediolateral hemimandible with p3–m1; BAT-3 s-683A and 683B, fragments width in the center of the metapodial; mrh, mandibular ramus of left hemimandible with p3–m1; BAT-3’07 908 and BAT-3’10 1, height; mw, maximum width of the talus; pad, mesiodistal length incomplete mandibles with left c–m1 and right p3–m1; BAT-3’11 of the paraconid; padh, paraconid height; pel, proximal epiph- 1467, incomplete right hemimandible with m1. Postcranial ele- ysis length; pew, proximal epiphysis width; ppl, length from the ments: BAT-3’06 260, BAT-3’06 229, BAT-3’07 1003, BAT-3’01 caudal border of the palate to the caudal border of the foramen 6e, BAT-3’09 1611, BAT-3’11 590, BAT-3’11 827, BAT-3’11 204, Downloaded by [CSIC Instituto Quimica Fiscia Rocasolano] at 07:36 04 March 2014 magnum; prd, mesiodistal length of the protoconid; prdh,proto- BAT-3’11 1124, and BAT-3’11 1938, right calcanei; BAT-3’08 conid height; tash, talar articular surface height; tasw, talar artic- 151, BAT-3 s-235, BAT-3’11 1391, BAT-3’11 838, and BAT-3’11 ular surface width; tcw, tuber calcanei width; tl, total length; trh, 1866, left calcanei; BAT-3’07 403, BAT-3 s-361, BAT-3’11 687, trochlea height of the talus; ysl, jugal dental series length. and BAT-3’2067, right tali; BAT-3’08 905, BAT-3’464, BAT-3’07 278, BAT-3 s-359, BAT-3’08 410, BAT-3 s-101, BAT-3’11 686, SYSTEMATIC PALEONTOLOGY and BAT-3’11 688, left tali; BAT-3’09 869, BAT-3’11 497, and BAT-3’11 1313, right Mc–II; BAT-3 s-222, BAT-3 s-631a, BAT- Order CARNIVORA Bowdich, 1821 3 s-216, BAT-3’11 40, and BAT-3’11 497, left Mc–II. Suborder FELIFORMIA Kretzoi, 1945 Family FELIDAE Fischer, 1817 Subfamily MACHAIRODONTINAE Gill, 1872 Description Tribe SMILODONTINI Kretzoi, 1929 Genus PROMEGANTEREON Kretzoi, 1938 Skull—The size and overall morphology of the skulls of P. PROMEGANTEREON OGYGIA (Kaup, 1832) Kretzoi, 1938 ogygia from Batallones-3 are essentially similar to those from (Figs. 3–6) Batallones-1 (Fig. 3). Most of skulls show a strong lateral or dorsoventral ﬂattening, which prevents the observation of sev- Referred Material—Cranial, dental, and mandibular elements: eral structures. The nasals are relatively narrow, subrectangular BAT-3’09 1507 and BAT-3’11 493, right upper canines; BAT-3’11 in shape, with rostral and caudal margins of similar width, and 410 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 34, NO. 2, 2014

FIGURE 3. Skulls of Promegantereon ogygia from Batallones-3. A, B, BAT-3’06 519 in A, lateral view; B, ventral view; C, D, BAT-3’06 591 in C, dorsal view; D, ventral view.

slightly curved caudally; the caudal margin lacks the narrowing tal protuberance, extending from the caudal margin of the tym- observed in the nasals of large extant felids. The left and right panic bulla. The mastoid process is robust and relatively well de- temporal lines meet dorsally at the level of the glenoid fossa, giv- veloped; it is located on the lateral wall of the tympanic bulla, ing way to a strong sagittal crest that ends in a robust nuchal crest. rostrally to the jugular process and slightly surpassing its ventral The rostral border of the premaxilla is curved rostrally, produc- level. The palate is triangular, with a nasopharyngeal fossa delim- ing a ‘U’-shaped arcade. There is a short diastema between I3 itated by the walls of the pterygoid bone. The palatine fissurae are and the upper canine. The zygomatic process of the frontal is placed close to the rostral border of the palate. sharp, rough, and slightly laterally projected; it is the origin of Upper Dentition—The fossil sample of P. ogygia from a marked temporal line. The infraorbital foramen is large and Batallones-3 does not include either I1 or I2, although from the placed at the level of P3. The lacrimal foramen is placed on the alveoli it can be observed that they were much smaller than I3 inner face of the rostral border of the orbit. The sphenopalatine (Fig. 3B). The I3 crown is sharp and caniniform; its mesial surface foramen is relatively smaller than that of Panthera; it is placed is convex and smooth, whereas the distal one is concave and bears on the orbital wall of the palatine bone, close to the maxillopala- a slight cingulum along its lingual border; on the buccal surface tine suture. The posterior palatine foramen is smaller than the there is a marked ridge from the base to the tip of the crown. The sphenopalatine foramen, and it is located slightly rostrally to the upper canines show the same morphology as those of P. ogygia latter. The oval foramen opens on the basisphenoid bone, be- from Batallones-1 (Fig. 4A–D), with a laterally flattened crown tween the pterygoid process and the postglenoid process. The zy- that curves backwards, with no crenulations on its mesial or Downloaded by [CSIC Instituto Quimica Fiscia Rocasolano] at 07:36 04 March 2014 gomatic arch is much more robust than that of a similarly sized distal margin; both buccal and lingual surfaces are smooth, al- extant felid, and dorsoventrally higher. The frontal process of the though the mesiolingual border bears a marked crest developed zygomatic is very small, although it is present in all the specimens; from the base to the middle of the crown. One skull (BAT-3’07 it is developed as a small bony bulge, ventrocaudally oriented, 1010) shows a very small tooth in both left and right maxillae, with an irregular ventral end. The tympanic bullae are rounded, mesial to P3, clearly separated from it; its crown is round and very less inflated than those of Panthera, and extend from the rostral low, and it could represent a retained D2, rather than a P2. No margin of the jugular process to the caudal margin of the post- trace of this tooth was found in the 11 studied skulls of P. ogygia glenoid process without contacting with the latter. The caudal en- totympanic penetrates rostrally beside the medial margin of the ectotympanic. The external auditory meatus is a single, rostrolat- TABLE 1. Summary of the ventral measurements (mm) of the skulls of erally oriented opening. The stylomastoid foramen and tympa- P. ogygia from Batallones-1 and Batallones-3. nohyal depression are very close to each other. Both are placed in a more rostral position than in Panthera, that is, at the level bl ppl dsl of the central part of the bulla. The jugular process is reduced in Locality N Mean SD Mean SD Mean SD size when compared with that of extant large felids, being dorsally BAT-1 9 166.07 8.58 85.17 3.62 80.18 4.03 displaced in comparison with its position in the latter, and medio- ...... laterally oriented; it is developed as a bony sheet with a small dis- BAT-3 8 159 41 7 67 81 07 5 15 74 88 3 99 SILICEO ET AL.—PROMEGANTEREON OGYGIA FROM BATALLONES SITES 411

TABLE 2. Summary of the measurements (mm) of the dental remains of P. ogygia from Batallones-1 and Batallones-3.

MDL BLW CH Locality Element N Mean SD Mean SD Mean SD BAT-1 C 31 14.89 0.93 8.86 0.53 35.78 2.81 BAT-3 C 10 14.26 1.13 8.58 0.72 35.60 3.44 BAT-1 P3 25 15.37 1.55 7.81 1.15 7.61 0.49 BAT-3 P3 12 15.18 0.85 7.43 0.48 8.81 0.47 BAT-1 P4 25 25.15 1.19 12.64 0.50 11.99 0.69 BAT-3 P4 14 24.69 1.24 11.69 0.69 12.38 0.78 BAT-1 c 32 10.75 0.68 7.61 0.52 20.93 1.78 BAT-3 c 4 10.17 0.51 7.07 0.39 19.37 0.14 BAT-1 p3 37 11.89 0.75 5.64 0.44 6.21 0.59 BAT-3 p3 12 11.83 0.57 5.36 0.31 6.81 0.73 BAT-1 p4 28 16.46 0.90 7.34 0.39 9.38 0.84 BAT-3 p4 10 16.28 0.64 6.77 0.23 10.29 0.47 BAT-1 m1 23 19.24 0.97 8.24 0.33 10.63/10.49∗ 0.78/0.47∗ BAT-3 m1 11 18.74 1.07 7.99 0.26 10.12/9.42∗ 0.79/0.73∗

∗First measurement refers to paraconid height, second measurement refers to protoconid height. Downloaded by [CSIC Instituto Quimica Fiscia Rocasolano] at 07:36 04 March 2014

FIGURE 4. Canines of Promegantereon ogygia from Batallones-3. A, B, left upper canine BAT- 3’11 1188 in A, buccal view; B, lingual view; C, D, right upper canine BAT-3’09 1507 in C, lingual view; D, buccal view; E, F, right lower canine BAT-3’08 833 in E, lingual view; F, buccal view; G, H, left lower canine BAT-3’11 1937 in G, buccal view; H, lingual view. 412 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 34, NO. 2, 2014

FIGURE 5. Detailed view of the upper and lower dentition of Promegantereon ogygia from Batallones-1 (A, B) and Batallones-3 (C–H). A, occlusal view of the left P3–M1 of the maxilla B-3570; B, buccal view of the left p3–m1 of the hemimandible B-462; C, occlusal view of the left P3–M1 of the maxilla BAT-3’01 60A; D, buccal view of the left p3–m1 of the hemimandible BAT-3 s-683A; E, occlusal view of P3–M1 of the skull BAT-3’06 519; F, buccal view of right p3–m1 of the hemimandible BAT-3’11 132; G, occlusal view of the left P3–P4 of the skull BAT-3’06 591; H, buccal view of left p3–m1 of the mandible BAT-3’10 1773 (E and F are shown inverted).

from Batallones-1. The P3 is placed parallel to the maxilla mar- the mandibular condyle. The angular process is caudoventrally gin, with its mesial half slightly displaced lingually (Fig. 3B, D). oriented, with a rough lateral border; its medial surface has a ros- The crown is mesiodistally lengthened, with a slightly convex buc- trocaudally oriented ridge that produces a marked groove. cal border, and a moderate constriction at the level of the main Lower Dentition—The lower canines are clearly smaller than cusp; there is a small distolingual basal protuberance, clearly re- the upper ones; their crown, curved distally and lacking any duced when compared with that from Batallones-1 (Fig. 5A, C, crenulation, shows a smooth lateral compression (Fig. 4E–H). E, G); there is no cingulum, except on the distal border. Most of The lingual face is concave and has a longitudinal ridge, whereas the available P3 lack the mesial cusp, as those from Batallones-1, the buccal face is smooth and slightly convex; the root has a although in the skull BAT-3 ‘07 1010 both P3 show a small but central bulge on its lingual face, whereas the buccal one is rel- patent mesial cusp; the distal and central cusps are strong, with atively ﬂat. A tiny, single-rooted d2, also present in P. ogygia the latter being much higher. The P4 has a relatively reduced pro- from Batallones-1, is present on the middle of the postcanine di- tocone when compared with those from Batallones-1, placed be- astema in the mandible BAT-3’10 1. The p3 crown is triangular tween the parastyle and paracone and with a more variable orien- on occlusal view, with a much wider distal half than the mesial tation, ranging from mesiolingually to almost lingually oriented one (Fig. 5D, F, H); it has a well-developed main cuspid, and a (Fig. 5A, C, E, G). The parastyle is well developed, and there is very reduced but present distal one, with a moderate distal cin- no trace of an ectostyle. The metacone-metastyle edge is slightly gulum; there is no mesial cuspid. The p4 crown is also triangularly shorter than the parastyle, with a marked notch separating both shaped, with a high central cuspid and small mesial and distal cus- structures. There is no cingulum. The M1 is very reduced, al- pids; both mesial and distal cuspids are separated from the central though it retains two roots; it has a buccolingually lengthened one by marked notches (Fig. 5D, F, H). The crown has a marked crown, with a slight ridge extending from the mesiobuccal to the posterior cingulum around the distal cuspid. As in p3, the crown lingual borders (Fig. 5A, C, E). of p4 is sharpened mesially. The m1 is similar to that of P. ogy- Mandible—The mandibular corpus may have either one sin- gia from Batallones-1, although it shows a reduced talonid, with Downloaded by [CSIC Instituto Quimica Fiscia Rocasolano] at 07:36 04 March 2014 gle, large foramen located at the level of p3 (BAT-3’10 1) or absence of metaconid, a cuspid that is present in the m1 from two mental foramina, the rostral one at the level of the post- Batallones-1(Fig.5B,D,F,H);theprotoconidisslightlyhigher canine diastema, and the caudal one, which is smaller, at the and mesiodistally longer than the paraconid, with a marked notch level of p3 (Fig. 6). Thus, as discussed for the sample of P. ogy- separating them. gia from Batallones-1 (Salesa et al., 2010b), the size of these Postcranial Skeleton—The postcranial anatomy of P. ogygia foramina seems to be highly variable, probably lacking any tax- from Batallones-1 was described by Salesa (2002) and Salesa et al. onomic value. The mandibular symphysis has a vertical anterior (2010a), so in this section we will only mention the differences border, and there is an almost straight angle between the ven- with the population from Batallones-3. tral border of the symphysis and the mandibular corpus. In ros- The Mc II of P. ogygia from Batallones-3 are very similar to tral view, the symphysis is high and rectangular, showing a light those from Batallones-1 in both morphology and proportions; mental ridge, more marked in its ventral half. The masseteric nevertheless, there is an interesting difference regarding the at- fossa is deep and its anterior border reaches the level of the m1 tachment area for the muscle ﬂexor carpi radialis; this scar is protoconid. There is just one preserved coronoid process within located on the medial border of the palmar tubercle, and it is the Batallones-3 sample, and it seems to be slightly larger than clearly more distally elongated than those from Batallones-1 (Fig. those from Batallones-1, although smaller than that of the simi- 7E–G). larly sized Pa. pardus; its posterior margin is markedly caudally The overall morphology of the tali from Batallones-3 is similar curved, and it does not surpass the level of the anterior margin of to those from Batallones-1, although some interesting differences SILICEO ET AL.—PROMEGANTEREON OGYGIA FROM BATALLONES SITES 413

FIGURE 6. Mandibles of Promegantereon ogygia from Batallones-3. A, B, complete mandible BAT-3’10 1773 in A, left buccal view; B, occlusal view; C–E, right hemimandible BAT-3’11 132 in C, occlusal view; D, buccal view; E, lingual view.

have been found. In dorsal view, the trochlea is wide, with a cen- lateral and medial borders, the latter showing a greater proximal tral, proximodistal soft groove that divides the trochlea into two projection (Fig. 7C, D). The plantar border of the tuber is a rough similarly wide lips; the trochlea is relatively lower in Batallones- surface for the attachment of the muscle ﬂexor digitorum super- 3 than in Batallones-1, in relation to both the maximum height ﬁcialis. Proximally, a round and smooth surface gives attachment of the talus and the width of the trochlea (Fig. 7A, B). In plan- for the common calcaneal tendon. The dorsal talar facet for the tar view, a relatively wide sulcus tali separates two large articu- lateral plantar facet of the talus is located on the coracoid pro- lar facets for the calcaneus; the lateral facet is rectangular and cess, markedly projected dorsally; it is convex and proximodis- strongly concave, whereas the medial one is round and convex, tally developed. The dorsal talar facet for the medial plantar facet extending onto the astragalus neck to join the talar head on its of the talus is round, with ridged margins, and located on a medi- medial margin. On the lateroplantar border of the head there is a ally projected sustentaculum tali; this second talar facet continues triangular facet for the calcaneus. The neck is well developed, distally by means of a narrow prolongation that ends in a small, and the talar head is elliptic, with its lateral border being dis- square facet. Between both talar facets, there is a deep sinus tarsi,

Downloaded by [CSIC Instituto Quimica Fiscia Rocasolano] at 07:36 04 March 2014 placed dorsally in relation to its medial one. In proximal view, narrower than that from Batallones-1. The plantar surface of the the trochlea may show a talar foramen. Finally, in distal view, the sustentaculum tali has a deep groove for the passage of the ten- angle between the talar head and the mediolateral axis is close to don of the muscle flexor digitorum lateralis. In Batallones-3, the 45◦. whole talar area is mediolaterally narrower, and its distal mar- The calcanei from Batallones-3 are similar to those from gin is less laterally inclined than those from Batallones-1. The Batallones-1 in overall morphology, although it also shows sev- fibular tubercle (distolateral expansion for the attachment of the eral differences. It is proximodistally elongated, and slightly ligament collaterale tarsi laterale longum) shows a shallow lat- mediolaterally flattened. The tuber calcanei is round, with rough eral groove. On the lateral surface of the calcaneus there is a

TABLE 3. Summary of the measurements (mm) of the Mc II of P. ogygia from Batallones-1 and Batallones-3.

tl pew pel mlw del dew Locality N Mean SD Mean SD Mean SD Mean SD Mean SD Mean SD BAT-1 39 59.17 1.84 12.37 0.77 16.27 0.75 7.92 0.55 12.48 0.73 11.79 0.60 BAT-3 8 61.99 2.31 12.12 0.57 16.41 0.89 8.02 0.42 12.32 1.20 12.80 0.55 414 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 34, NO. 2, 2014

TABLE 4. Summary of the measurements (mm) of the talus of P. ogygia from Batallones-1 and Batallones-3.

th mw mh Locality N Mean SD Mean SD Mean SD BAT-1 41 24.62 1.41 26.08 1.34 35.01 2.43 BAT-3 13 21.62 1.49 25.29 2.29 34.02 1.85

gentle depression, proximodistally elongated for the attachment of the muscle quadratus plantae; in both populations of P. ogygia it is developed from the distal border to the middle of the dorsal border of the calcaneus. Between the proximal border of the attachment surface of the muscle quadratus plantae and the dor- solateral border of the calcaneus, there is a marked round fossa for the ligament short collateral lateral. The distal articular surface for the cuboid is semicircular, gently concave, and its plane is laterally much more inclined in Batallones-1 than in Batallones- 3. Close to the medioplantar border of this facet there is a rough tubercle for the long plantar ligament.

Statistical Results The results of the different Student’s t-tests calculated for the quantitative data are shown in Table 6. The most interesting results were as follows:

1. The P3 shows significant differences in the ratio of the height (H) and the mesiodistal length (MDL) of the crown, as shown by a Student’s t-test for this index. For similar crown lengths, the population of Batallones-3 shows P3 with proportionally higher crowns than those from Batallones-1. Also, significant differences were found in the buccolingual width (BLW) of P3, with those specimens from Batallones-3 being narrower than those from Batallones-1. 2. The P4 shows significant differences in Student’s t-test for an index between the buccolingual width (BLW) and the mesiodistal length (MDL) of the crown. The P4 from Batallones-3 are buccolingually narrower in proportion to their length. 3. The mesiodistal length of the P4 (MDL) shows significant differences in relation to the basal length of the skull (BL), the population from Batallones-3 having a relatively longer P4 than those from Batallones-1. 4. In the p3, a Student’s t-test shows significant differences in the index between the height (H) and mesiodistal length (MDL) of the crown. The p3s from Batallones-3 show higher crowns in relation to their length than those from Downloaded by [CSIC Instituto Quimica Fiscia Rocasolano] at 07:36 04 March 2014 Batallones-1. 5. The height of the p4 shows significant differences between both populations. This is demonstrated by a Student’s t-test for an index between the height (H) and length (MDL) of the crown, with the p4s from Batallones-3 showing relatively higher crowns than those from Batallones-1. 6. Concerning the calcaneus, the articular surface for the talus shows significant differences in its proportions, as shown by a Student’s t-test for an index between the width (TASW) and the height (TASH) of the articular surface for the talus, with Batallones-3 showing a relatively narrower talar articu- FIGURE 7. Postcranial elements of Promegantereon ogygia from lar surface. Batallones-3 that showed statistically significant differences with those 7. Also in the calcaneus, the index between the width of the from Batallones-1. A, B, left talus BAT-3’08 905 in A, dorsal view; B, distal facet for the cuboid (DFW) and the width of the ta- plantar view; C, D, right calcaneus BAT-3’06 260 in C, lateral view; D, lar articular surface (TASW) shows significant differences, dorsal view; E–G, left Mc II BAT-3 s-216 in E, dorsal view, F,medial with the calcaneus from Batallones-3 showing a relatively view; G, lateral view. narrower talar articular surface than those from Batallones-1. SILICEO ET AL.—PROMEGANTEREON OGYGIA FROM BATALLONES SITES 415

TABLE 5. Summary of the measurements (mm) of the calcaneus of P. ogygia from Batallones-1 and Batallones-3.

ti tasw dfw tash tuw Locality N Mean SD Mean SD Mean SD Mean SD Mean SD BAT-1 38 62.96 2.56 26.25 1.73 17.26 1.15 25.25 1.67 17.00 1.09 BAT-3 15 62.81 2.97 24.51 1.70 18.10 1.78 24.58 1.87 16.94 1.19

8. Also the talar articular surface of the calcaneus (TASW) is one of the eight available skulls from Batallones-3. This could significantly narrower in relation to the calcaneus tubercle be considered as a plesiomorphic character, because it is present width (TW) in Batallones-3, as indicated by a Student’s t-test in the more primitive machairodontine Pseudaelurus quadriden- for an index between both measurements (TW/TASW). tatus from the Middle Miocene of Europe. On the contrary, the 9. A Student’s t-test for the talus showed significant differences presence in Batallones-3 of a relatively narrow P3, with a reduced in the index between the height of the trochlea (TH) and basal expansion of the crown, implies a morphological separa- the maximum width of the talus (MW), with those tali from tion of this taxon from Ps. quadridentatus, which shares with P. Batallones-3 showing a relatively lower TH than those from ogygia from Batallones-1 the presence of a marked distolingual Batallones-1. basal expansion in the P3 (Fig. 5A, C, E, G). Other Smilodon- 10. Finally, a Student’s t-tests for an index between the height tini such as the Turolian (MN 11–12) Paramachaerodus orientalis, of the trochlea (TH) and the maximum height of the talus the Pliocene Megantereon, and the Pleistocene Smilodon show a (MH) showed significant differences between both popula- relatively small P3, without distolingual expansion (Salesa et al., tions, with that of Batallones-3 having relatively lower TH 2010b). The difference between the P3 from Batallones-1 and than that of Batallones-1 when compared with MH, that is, a Batallones-3 probably indicates the beginning of the strong re- relatively longer talus neck. duction of the P3 observed in more derived taxa. The relatively reduced protocone in the P4 of P. ogygia from It should be noted that although both populations show size Batallones-3 (which is relatively buccolingually narrower than differences in some of the elements, such as the total length of the that from Batallones-1) in comparison with the morphology of calcaneus, these differences are not significant; thus, we should those from Batallones-1 is a derived feature (Fig. 5A, C, E, G), consider the specimens from Batallones-1 and Batallones-3 as also observable in the more advanced Paramachaerodus orien- having a very similar body size. talis, although this species also shows a marked ectostyle, absent in P. ogygia. More derived machairodontines such as Smilodon Discussion and Megantereon show markedly reduced protocones in P4. The primitive condition for this character can be seen in the The identification of the small teeth mesial to P3/p3 as D2/d2 early Miocene felid Proailurus lemanensis, which shows a well- is based on their extremely reduced size and simple morphology, developed P4 protocone, clearly mesiolingually oriented, similar which precludes their identification as permanent premolars. Al- to the condition observed in members of the family Viverridae. though Salesa et al. (2010b) cite the presence of a very small Once again, we observe in the Batallones-3 material a slight mod- p2 in P. ogygia from Batallones-1, such reduced premolars, ob- ification from the dental morphology seen in the older population served in several fossil felids, have been more recently considered from Batallones-1. The reduced protocone creates a more tren- as retained decidual teeth (Salesa et al., 2012), and that is more chant P4, a trait not only observed in more-derived smilodon- likely the case of the small premolars observed in P. ogygia.It tins, but also in derived non-felid saber-toothed taxa, such as is remarkable that whereas none of the skulls of P. ogygia from barbourofelids and nimravids (Bryant, 1991, 1996; Morales et al., Batallones-1 show any trait of D2, this tooth appears in at least 2001; Morlo et al., 2004). This feature, often associated with a lengthening of the P4 crown, is clearly related to more efficient meat consumption, because it enlarges the cutting blade of the TABLE 6. Summarized results of the Student’s t-tests showing the vari- upper carnassial. Also, P3, p3, P4, and p4 show relatively higher ables that showed significant differences. crowns in Batallones-3 than in Batallones-1, a difference for which we found no clear functional implication. These higher and Element Variable P Comments thus more pointed crowns could also increase the shear capacity P3 MDL/H 0.000 BAT-3 shows relatively higher of both teeth, mostly considering that this difference is found in Downloaded by [CSIC Instituto Quimica Fiscia Rocasolano] at 07:36 04 March 2014 P3. all the premolars. P3 BLW 0.001 BAT-3 shows narrower P3. The sample of m1 from Batallones-3 shows a reduced talonid, P4 MDL/BLW 0.010 BAT-3 shows relatively a derived feature associated with a more trenchant lower carnas- narrower P4. sial (Fig. 5B, D, F, H). The talonid of P. ogygia from Batallones-1 . P4 MDL/BL 0 013 BAT-3 shows relatively longer shows a basal expansion retaining a tiny metaconid (Salesa et al., P4. p3 H/MDL 0.001 BAT-3 shows relatively higher 2010b), whereas in Batallones-3 the m1 shows a smaller talonid p3. composed only of a basal expansion, without any trace of a meta- p4 H/MDL 0.000 BAT-3 shows relatively higher conid. The primitive felids Proailurus lemanensis and Pseudaelu- p4. rus quadridentatus show a well-developed talonid with a marked Calcaneus TASW/TASH 0.002 BAT-3 shows relatively metaconid, whereas in derived Smilodontini such as Megantereon narrower TASW. and Smilodon the talonid is absent. Another primitive smilodon- Calcaneus DFW/TASW 0.000 BAT-3 shows relatively narrower TASW. tin, the Eurasiatic Pa. orientalis, shows a very reduced talonid, Calcaneus TW/TASW 0.000 BAT-3 shows relatively whereas in the closely related Pa. maximiliani it is completely ab- narrower TASW. sent (Salesa et al., 2010b). In this reduction in the m1 talonid, the Talus TH/MW 0.000 BAT-3 shows relatively lower population of P. ogygia from Batallones-3 again shows a derived TH. morphology in relation to that from Batallones-1. Talus TH/MH 0.000 BAT-3 shows relatively lower TH. Concerning the postcranial skeleton, some of the observed differences between the populations of Batallones-1 and 416 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 34, NO. 2, 2014

Batallones-3 can be related to functional differences indicating increased cursorial abilities in the specimens from the latter sample. Thus, the main consequence of having a relatively narrower talar area in the Batallones-3 calcanei (Fig. 8A, B), besides producing a less inclined distal border, is the presence of a more vertical orientation of the calcaneus and cuboid, and thus more parasagittal flexion-extension movements of the ankle, something observed in many cursorial carnivorans (Gambaryan, 1974; Taylor, 1989). The relatively lower trochlea of the talus in Batallones-3 would reduce the range of flexion-extension movements of the tibia, which does not have a direct cursorial implication, but could be a consequence of the more vertical movements of the talocalcaneal joint. Another important difference between both samples is the morphology of the Mc II: in Batallones-3, the facet for one of the tendons of the muscle flexor carpi radialis, located on the medial face of the proximal epiphysis, is clearly distally elongated when compared with the Mc II from Batallones-1 (Fig. 8C, D). Among fossil and extant Felidae, the development of this facet is variable, with Panthera leo, Panthera atrox, Puma concolor, Lynx pardinus,andPanthera onca having markedly elongated facets, and Panthera pardus, Smilodon fatalis,andSmilodon gracilis showing relatively shorter attachment scars; the extreme condition is observed in Acinonyx jubatus, which shows a much more elongated facet than that of other species, reaching the middle point of the diaphysis. Thus, the greatest development of this facet is found among cursorial felids, which show a similarly sized facet as in other cursorial carnivorans such as canids (although in this later group the whole facet is distally displaced). The smallest facets seem to appear in relatively robust felids such as S. fatalis or Pa. pardus, and this would imply the presence of a relatively thinner tendon of the muscle flexor carpi radialis, a muscle that participates in flexion of the carpus (Barone, 2010), and together with other extensor and flexor muscles, it has an important role in both prey immobilization and locomotion. An intermediate size for this facet is seen in extant felids with different proportions and cursorial capacities, such as the relatively slender Pa. leo and the much more robust Pa. onca. This difference could reflect a phylogenetic relationship rather than any functional differences between species, especially if we consider that extant individuals of Pa. onca are relatively much more robust than their Pleistocene counterparts (Seymour, 1993). Given this, it is remarkable that strong differences in the development of this facet are found in both studied populations of P. ogygia. The individuals from Batallones-3 would have a thicker tendon for the muscle flexor carpi radialis, and thus probably a larger or better-developed muscle. The presence of a more powerful muscle in the specimens from Batallones-3 would imply an increase in the force of the hand during flexion, something directly related to prey subduing. Although the studied populations of P. ogygia from Batallones-1 and Batallones-3 show interesting differences Downloaded by [CSIC Instituto Quimica Fiscia Rocasolano] at 07:36 04 March 2014 in both dental and postcranial features, all of them seem insuf- ficient to propose a species-level separation between these two samples. On the contrary, they both should be considered as part of a gradual evolution of the ‘chronopopulation’ of this felid that inhabited the central basin of Spain during the late Miocene. Unfortunately, the last known fossils of P. ogygia are those from Batallones-3; thus, it is more likely that this species disappeared FIGURE 8. Comparison of calcaneus and Mc II of Promegantereon at the end of the Vallesian or beginning of the Turolian, when ogygia from Batallones-1 (A, C) and Batallones-3 (B, D), showing the morphological differences discussed in the text. A, B, dorsal view of A, the enigmatic Pa. orientalis, whose postcranial skeleton is almost left calcaneus B/S-341 and B, right calcaneus BAT-3’06 260 (showed in- unknown, was already present in the faunas of Eurasia. verted); C, D, medial view of C, left Mc II B-1598 (2) and D,leftMcII BAT-3 s-216, showing the differences in the development of the area for the attachment of the muscle flexor carpi radialis. CONCLUSIONS The study of the population of P. ogygia from Batallones-3 reveals the existence of a set of derived features in the dentition and postcranial skeleton compared with the older sample from Batallones-1. The differences in the dentition essentially SILICEO ET AL.—PROMEGANTEREON OGYGIA FROM BATALLONES SITES 417

imply the presence in Batallones-3 of more trenchant premolars Kretzoi, N. 1938. Die Raubtiere von Gombaszog¨ nebst einer Ubersicht¨ and molars, whereas those observed in the appendicular skeleton der Gesamtfauna. Annales Historico-Naturales Musei Nationalis can be related to a reduction in the weight of the hind limb in Hungarici 31:88–157. Batallones-3, which points towards an increase in cursorial abili- Kretzoi, N. 1945. Bemerkungen uber¨ das Raubtiersystem. Annales ties, and with the need for prey immobilization during the hunt. Historico-Naturales Musei Nationalis Hungarici 38:59–83. ´ All these differences, although indicating some degree of mor- Lopez-Anto´ nanzas,˜ R., P. Pelaez-Campomanes,´ M. A. Alvarez-Sierra, phological segregation between both populations, do not seem and I. Garcıa-Paredes.´ 2010. New species of Hispanomys (Rodentia, Cricetodontinae) from the Upper Miocene of Batallones (Madrid, strong enough to support the existence of a different species of Spain). Zoological Journal of the Linnean Society 160:725–747. Promegantereon in Batallones-3. Merriam, J. C., and C. Stock. 1932. The Felidae of Rancho La Brea. Carnegie Institution of Washington, Washington, D.C., 231 pp. Morales, J., M. J. Salesa, M. Pickford, and D. Soria. 2001. A new tribe, ACKNOWLEDGMENTS new genus and two new species of Barbourofelinae (Felidae, Car- This study is part of the research projects CGL2008-00034, nivora, Mammalia) from the Early Miocene of East Africa and CGL2008-05813-C02-01, and CGL2011-25754 (Direccion´ Gen- Spain. Transactions of the Royal Society of Edinburgh: Earth Sci- ´ ´ ences 92:97–102. eral de Investigacion, Ministerios de Ciencia e Innovacion y Morales, J., L. Alcala,´ L. Amezua, M. Anton,´ S. Fraile, E. Gomez,´ P. Economıa´ y Competitividad, Spain). G.S. is a contracted predoc- Montoya, M. Nieto, B. Perez,´ M. J. Salesa, and I. M. Sanchez.´ toral researcher (FPI) of the project CGL2008-00034 (Ministe- 2000. El yacimiento de el Cerro de los Batallones; pp. 179–190 in rio de Ciencia e Innovacion,´ reference BES2009-013437). M.J.S. J.Morales,M.Nieto,L.Amezua,S.Fraile,E.Gomez,´ E. Herraez,´ and J.M. belong to the research group UCM-BSCH-910607. We P. Pelaez-Campomanes,´ M. J. Salesa, I. M. Sanchez,´ and D. So- thank the Government of the Comunidad Autonoma´ de Madrid ria (eds.), Patrimonio Paleontologico´ de la Comunidad de Madrid. (Direccion´ General de Patrimonio Historico)´ for its continuous Comunidad de Madrid, Serie “Arqueologıa,´ Paleontologıa´ y Etno- funding support and research permissions, and J. F. Pastor (Uni- grafıa,”´ Volume 6. Comunidad de Madrid, Madrid. versity of Valladolid, Spain) for specimen loan. Morales, J., L. Alcala,´ M. A. Alvarez-Sierra,´ M. Anton,´ B. Azanza, J. P. Calvo, P. Carrasco, S. Fraile, I. Garcıa-Paredes,´ E. Gomez,´ M. Hern´ andez´ Fernandez,´ L. Merino, A. van der Meulen, C. Martın´ LITERATURE CITED Escorza, P. Montoya, M. Nieto, S. Peigne,´ B. 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