This is the accepted version of the journal article: Sánchez Marco, Antonio. «Two new Gypaetinae (Accipitridae, Aves) from the late Miocene of Spain». Historical biology, Vol. 34, Issue 8 (2022), p. 1534-1543. DOI 10.1080/08912963.2022.2053117 This version is available at https://ddd.uab.cat/record/264387 under the terms of the license Page 1 of 24 1 2 3 4 Two New Gypaetinae (Accipitridae, Aves) from the late Miocene of 5 Spain 6 7 Antonio Sánchez Marco 8 Institut Català de Paleontologia Miquel Crusafont, Barcelona, Spain. 9 10 antonio.sanchez@icp.cat 11 http://orcid.org/0000-0003-0654-1935 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 Page 2 of 24 1 2 3 4 Two New Gypaetinae (Accipitridae, Aves) from the late Miocene of 5 Spain 6 7 8 Abstract 9 Only three monospecific genera of extant accipitrids –Gypaetus, Neophron and 10 11 Gypohierax– have been grouped into the Gypaetinae vultures, which currently presents 12 an Eurasian and African geographic distribution. The fossil record of these genera is 13 meager and not older than the Pleistocene of both landmasses. A surprising Old World 14 vulture radiation occurred in North America from the early Miocene to the late 15 Pleistocene, with taxa more closely linked to Gypaetinae than to Aegypiinae. More 16 recently new fossil gypaetines have been described for the late Miocene of North 17 America and China. Two new species assigned to genera Gypaetus and Neophron are 18 19 described here, and their links with other related taxa are evaluated. The new specimens 20 were found at two sites in the Cerro de los Batallones, a rich late Miocene 21 paleontological complex located in the center of Spain. 22 23 Keywords: Avian systematics; Accipitriformes; diurnal raptors; Neogene birds. 24 25 26 27 Introduction 28 29 Old World vultures were traditionally considered closely related (Livezey and Zusi, 30 2007), but are now classified into two distantly related groups, Aegypiinae and 31 Gypaetinae. The extant Gypaetinae includes three monospecific genera: Gypaetus, 32 Neophron and Gypohierax, distributed across southern Europe, Africa and throughout 33 34 Asia, east to Tibet and India (del Hoyo et al., 1994). They constitute a more basal and 35 separate clade from the Aegypiinae (Lerner and Mindell 2005; Lerner 2007), the other 36 vultures within Accipitridae. Lerner and Mindell (2005) suggest that Eutriorchis astur 37 (Sharpe, 1875), an eagle endemic to Madagascar, could be closely related to the 38 Gypaetinae. The fossil record of the Accipitridae dates back to the late Eocene / early 39 Oligocene (Mayr 2009), but the earliest evidence of Gypaetinae is considerably more 40 modern (Mayr, 2017). Two new vultures referred to Gypaetinae have recently been 41 42 reported from the late Miocene of North America and China. Anchigyps voorhiesi 43 Zhang et al., 2012a, from Nebraska (USA), similar in size to Neophron percnopterus 44 and to the congeneric species described here. Mioneophron longirostris Li et al., 2016, 45 from the Gansu Province (China), has been described from a quite complete skeleton 46 and related to the extant Neophron percnopterus. The finding of Gypaetus barbatus 47 (Linnaeus, 1758,) in the early Pleistocene of the Iberian Peninsula (Montoya et al. 1999, 48 49 2001; Mlíkovský 2002) is so far the oldest record of the genus in the Old World 50 (Europe, Asia and Africa). Gypaetus osseticus Burchak-Abramovich, 1973, from the 51 late Pleistocene of Tsona (also known as Bugas-Klde, Georgia), was considered nomen 52 nudum by Brodkorb (1978). Fossil findings referred to Neophron come from many 53 middle and late Pleistocene sites and have been assigned to the recent species Neophron 54 percnopterus (Linnaeus, 1758) (Lambrecht 1933; Tyrberg 1998). 55 The Neogene of North America has yielded a substantial collection of fossils 56 57 referred to Old World vultures, Aegypiinae and Gypaetinae (Wetmore 1943; Rich 1977; 58 Zhang et al. 2012a; Li et al. 2016, and references therein). Neogyps errans Miller, 1916, 59 from the late Pleistocene of Rancho La Brea (California, USA), was closely related to 60 Gypaetus because of its clear resemblance to that genus by Miller (1916). Six species of Page 3 of 24 1 2 3 Neophrontops have been described in North America, dating back from the early 4 Miocene to the late Pleistocene (Brodkorb 1964; Olson and Rasmussen 2001). Also 5 6 fossil bones from early Miocene and to early Pliocene localities of North America have 7 each been attributed to species of the genus Palaeoborus (Cope 1874; Wetmore 1936; 8 Miller and Compton 1939). The absence of Neophron and related forms prior to the 9 Pleistocene in the Eurasian and African record has led to the suggestion that Neophron 10 may have originated from a dispersal of Neophrontops from North America (Feduccia 11 1974). Due to the presence of specific anatomical features for prey capture, 12 Palaeoborus and Neogyps were viewed of more eagle-like aspect than Neophrontops 13 14 and the extant Neophron (Miller and Compton 1939). 15 The paleontological sites of Cerro de los Batallones, located in the south of 16 Madrid, in the middle of Spain, were formed at the beginning of the late Miocene 17 (Tortonian Age). The faunal context points to the biozone MN 10, with deposition ages 18 of circa 9 Ma (Morales et al. 2008, Calvo et al. 2013). So far, in two of the sites, 19 Batallones 1 and Batallones 3, more than one hundred fossils of birds belonging to 20 21 about twelve species have been found. Most of them correspond to diurnal raptors. In 22 these two paleontological sites, Oonly galliform birds have previously reported 23 (Sánchez Marco 2006, 2021). 24 The avian assemblage recorded does not constitute a representative sample of 25 the birds that populated the surrounding areas. Carrion birds are predominate 26 predominant in these paleontological sites, which also happens with regard to mammals 27 (Morales et al. 2008). Two new Gypaetinae vultures found in Cerro de los Batallones 28 29 are described and analysed here. 30 31 32 Material and Methods 33 34 The osteological terminology follows to Baumel and Witmer (1993) and Howard 35 (1980). The systematic classification follows Lerner and Mindell (2005), Griffiths et al. 36 37 (2007) and recommendation of the IOC World Bird List, version 11.2 (Gill et al. 2021). 38 The fossil bones are housed in the Museo Arqueológico Regional de Madrid, Spain, 39 with the labels shown in this paper. 40 Institutional abbreviations: MNCN, Museo Nacional de Ciencias Naturales, 41 Madrid, Spain. NMNH, Division of Birds, National Museum of Natural History, 42 Smithsonian Institution, Washington, USA. CAM, Arturo Morales personal collection, 43 44 Laboratorio de Arqueozología, Universidad Autónoma de Madrid, Spain. LMUM, 45 Veterinary Faculty of the Ludwig Maximilians University of Munich, Germany. 46 Specimens of extant species used for comparisons: Necrosyrtes monachus 47 (NMNH-18894, 291440, 291441, 291442, 291443), Gypohierax angolensis (NMNH- 48 224820, 226143, 291078, 291316), Neophron percnopterus (CAM 2, CAM 3, MNCN 49 26843, NMNH-17835), Milvus migrans (NMNH-319228), Gypaetus barbatus (CAM 2, 50 CAM 3, MNCN 23330, NMNH-17834, 345684), Circaetus cinereus (NMNH-430776), 51 52 Terathopius ecaudatus (NMNH-17836, 289742, 292912, 319919), Trigonoceps 53 occidentalis (NMNH-320859, 347358), Torgos tracheliotus (NMNH-19990, 320977, 54 321827, 347597), Aegipyus monachus (NMNH-289569, 428040, 614152), Gyps 55 africanus (NMNH-431403, 587405), Gyps fulvus (NMNH-229051), Gyps rueppellii 56 (NMNH-430178), Gyps coprotheres (NMNH-561314) and Gyps himalayanus (NMNH- 57 19534), Haliaeetus albicilla (CAM 2, CAM 3, LMUM 2, LMUM 11), Haliaeetus 58 vocifer (LMUM 3), Haliaeetus leucocephalus (LMUM 2, LMUM 3). No morphological 59 60 or size differences have been observed between Necrosyrtes monachus and Necrosyrtes Page 4 of 24 1 2 3 m. pileatus. In consequence, the race pileatus has no relevance for this study, and we 4 refer it as to Necrosyrtes monachus. 5 6 7 8 Systematic Paleontology 9 10 Accipitriformes Voous 1973 11 Accipitridae Vieillot 1816 12 Gypaetinae Vieillot 1816 13 14 Gypaetus Storr 1784 15 Gypaetus georgii n. sp. 16 17 Holotype: One complete left carpometacarpus (Bat 3.2012.437). 18 19 Referred specimens: Batallones 1 site: cranial end of coracoideum and scapula (B- 20 21 1968), two right humeri (Bat 1.171, Bat 1.175), left radius (Bat 1.915), one proximal 22 end and shaft of left radius (Bat 1.2286), left os carpi radiale (Bat 1.2008.35), two left 23 carpometacarpi (Bat 1.2005.287, Bat 1.1403), two right phalanges proximalis digiti 24 majoris (Bat 1.2008.6, Bat 1.2004.68), right tibiotarsus (Bat 1.2005.6), crushed left 25 tibiotarsus and complete fibula (B-1970, not “in situ”). Batallones 3 site: right scapula 26 (Bat 3.2008.616), proximal end of left radius (Bat 3.2007.130), right phalanx proximalis 27 digiti majoris (Bat 3.2008.158), right tibiotarsus (Bat 3.2012.2591), crushed left 28 29 tibiotarsus (Bat 3.2009.912). 30 31 Type locality: Batallones 3, Cerro de los Batallones, Torrejón de Velasco village, 32 Madrid province, Spain. 33 34 Other locality: Batallones 1, Cerro de los Batallones, Torrejón de Velasco village, 35 Madrid province, Spain. Biozone MN 10, early Tortonian, late Miocene (Morales et al. 36 37 2008, Calvo et al. 2013). 38 39 Stratotype: Upper zone of the “Intermediate Unit” (Morales et al. 1992, sensu 40 Lomoschitz et al. 1985, Calvo et al. 2013), biozone MN 10, early Tortonian, late 41 Miocene (Morales et al. 2008). 42 43 44 Etymology: Dedicated to Jorge Morales, a friend and paleontologist, who has conducted 45 the works in the Batallones sites since their discovery. Georgii is the genitive of 46 “Georgius” –the Latin for “Jorge”-. 47 48 Dimensions and figures: Table 1 and Figure 1. 49 50 Diagnosis: Differs from extant Gypaetus barbatus in: 1) smaller size (see Table 1); 2) 51 52 coracoideum, processus procoracoideus large and curved medially and dorsally; 3) 53 scapula, shorter and less developed acromion; 4) humerus, tuberculum dorsale not 54 protruding from the line going between the caput humeri and the crista deltopectoralis,; 55 5) epicondylus dorsalis scarcely developed, 6) more prominent processus flexorius; 7) 56 radius, tuberculum bicipitale radii is closer to the facies articularis ulnaris in the new 57 species than in the other vultures compared; 8) carpometacarpus, processus extensorius 58 more horizontally oriented 9) tibiotarsus in distal view, more square-shaped distal end 59 60 (vs. rectangular in G. barbatus). Page 5 of 24 1 2 3 4 5 6 Descriptions and comparisons 7 8 Coracoideum: The remains of this bone are in a poor state of preservation. The 9 processus procoracoideus is large and curved medially and cranially, as in Gyps, 10 Aegypius and Torgos. In Gypaetus, Neophron and Terathopius the processus 11 procoracoideus is scarcely developed as well as almost not curved. This trait is not 12 observable in Mioneophron longirostris. In Trigonoceps, it is not curved at all. In 13 14 Gypaetus georgii n. sp., the processus procoracoideus is larger than in G. barbatus. 15 There are not remains of coracoids on which to take measures owing to their poor state 16 of preservation. The size of this bone is considerable smaller than in G. barbatus. 17 18 Scapula: In cranial view, the triosseal surface of the acromion is less elongated (less 19 developed dorso-cranially) than in any compared species. In this respect, Trigonoceps, 20 21 Gypaetus and Terathopius show more elongated acromions than Gyps, Aegypius and 22 Torgos. The triosseal surface of the fossil scapula do not bear pneumatic foramens. This 23 trait seems to be connected with the size of the specimen as the smaller species, 24 Trigonoceps occidentalis (Burchell 1824) and Terathopius ecaudatus (Daudin 1800), 25 likewise lack it. Nevertheless, the latter species has a large pneumatic area just in the 26 dorsal surface of the acromion. In Gypaetus, Torgos and Aegypius there is only a single 27 foramen whereas in Trigonoceps, Gyps and in the fossil there is no pneumatization. 28 29 30 Humerus: Both humeri are crushed and in a bad state of preservation. They show the 31 general aspect of the Old World vultures. The tuberculum dorsale is fairly separated 32 from the head, a shared feature within extant Gypaetinae. The region of the processus 33 flexorius is relatively more developed than in Gyps, Aegypius, Neophron and even G. 34 barbatus. In the extant Gypaetinae and in both new species described in this paper, the 35 ventral condyle is larger and fairly more protruding distally than in Neophrontops 36 37 vetustus (Rich 1977). In addition, the region of the flexor process and the 38 entepicondylar prominence is more robust and rounded in Neophron and Gypaetus. A 39 furrow that starts at the attachment of the latissimus dorsi posterioris and continues on 40 the shaft towards the end of the crista deltopectoralis is very marked in the new species. 41 It is present in Gypaetinae (Rich 1977), although not exclusive. 42 43 44 Radius: To a certain extent, size and general shape of the radius, including curvature 45 and twisting of the diaphysis, are distinctive among accipitrids. The overall shape of the 46 fossils is like that observed in other Old World vultures. The tuberculum bicipitale radii 47 is closer to the facies articularis ulnaris in the new species than in the other vultures 48 compared with. The area of the depressio ligamentosa and the facies articularis ulnaris 49 is smooth in the fossil, as well as in Terathopius and Gypaetus. This region is deeper in 50 the other species. The tuberculum bicipitale radii is closer to the facies articularis ulnaris 51 52 than in G. barbatus. On the other extreme of the bone, the sulcus tendinosus is slightly 53 marked out, as also in Torgos and Aegypius. In Torgos, Gypaetus, Aegypius and in the 54 fossil the overall distal end protrudes distally more than in the rest of compared taxa. 55 56 Carpometacarpus: In the new species, on the ventral side of the proximal end there are 57 two deep fossae, the fossa infratrochlearis (Baumel and Witmer 1993; internal 58 ligamental fossa of Howard 1980) and the fovea carpalis cranialis, which is situated 59 60 between the processus pisiformis and the base of the os metacarpale alulare. This Historical Biology Page 6 of 24 1 2 3 pattern is only seen in Torgos and Aegypius. Likewise, at the base of the os metacarpale 4 alulare, but in dorsal view, the fossil presents a marked depression, deeper than in the 5 6 other taxa compared. One exclusive feature is the short distance between both edges of 7 the trochlea carpalis, in cranial as in ventral views. In Aegypiinae, the os metacarpale 8 minus expands cranially considerably towards its junction with the os metacarpale 9 majus. This expansion is relatively broader in G. georgii n. sp. A character only shared 10 with Torgos is that the os metacarpale minus exceeds the os metacarpale majus distally. 11 The processus extensorius is more curved ventrally in the new species than in G. 12 barbatus. 13 14 15 Tibiotarsus: The two proximal thirds of the fossil are crushed. Length and general 16 robustness are like in Gypaetus barbatus and Trigonoceps occidentalis. The cranial 17 edge of the crista cnemialis cranialis is round, like in Gyps, Aegypius and Trigonoceps, 18 not straight, like in the other taxa used for comparisons. The condylus medialis is round, 19 like in Torgos, Gyps, Aegypius, Necrosyrtes, Terathopius, Trigonoceps, Haliaeetus, 20 21 Gypaetus, Neophron and Gypohierax. Both distal condyles are less rounded in eagles. 22 In cranial aspect, the distal end of the fossil is quite symmetric with regard to its 23 longitudinal axe. This pattern is only observed in Torgos tracheliotus (Forster 1791). 24 Likewise, in distal view, the incisura intercondylaris is narrow, like in Torgos 25 tracheliotus, Trigonoceps, Aegypius and Gyps. It is much wider in Gypaetus barbatus. 26 The distal end, in distal view, offers a more rectangular shape in the accipitrids 27 compared, including Gypaetus barbatus, than in the fossil. The new species shows a 28 29 supratendinal bridge much more vertical than Gypaetus barbatus. In Neophron it is 30 practically horizontal. In Anchigyps voorhiesi, it is almost horizontal, but extraordinarily 31 wide (Zhang et al. 2012a), more than in extant Gypaetinae and Aegypiinae. Arikarornis 32 macdonaldi Howard, 1966, was erected on one dital end of tibiotarsus. Neither the 33 description nor the figure of paper correspond to the morphology of this bone in the Old 34 World vultures. The remarkable inclination of the supratendinal bridge in Arikarornis 35 macdonaldi resembles that of Gyps, but the morphology of the rest of the bone is unlike 36 37 in Gypaetinae or Aegypiinae. 38 39 Remarks: All the bones of Gypaetus georgii n. sp. are smaller in size than in the G. 40 barbatus individuals to which they have been compared (Table 1). Forelimbs bones of 41 the new species are 10 to 20 % smaller, while the tibotarsus is only 10 % shorter. This 42 estimate is conjectural since the size ratios would surely change if fossil and 43 44 comparative samples grew. In addition, we lack femora and tarsometatarsi to know if 45 the ratio between fore and hindlimbs has changed. As for the morphological features, 46 scarce differences have been found. Neogyps errans, from the Pleistocene of California, 47 is supported on the description of the tarsometatarsus. It bears morphological 48 resemblance to Gypaetus barbatus, and their sizes are similar, although Neogyps is 49 much less robust at its proximal end (Miller 1916). 50 51 52 Neophron Savigny, 1809 53 Neophron lolis n. sp. 54 55 Holotype: Complete right tarsometatarsus (Bat 1.3087). 56 57 Referred specimens: Batallones 1 site: fragmentary cranium (Bat 1.2009.69), partially 58 crushed right humerus (Bat 1.59), almost complete left ulna (Bat 1.2004.209), proximal 59 60 end and shaft of right ulna (Bat 1.2002.247), two distal ends of right ulnae (Bat 1.3306, Page 7 of 24 1 2 3 Bat 1.2006.128), right radius (Bat 1.6833), proximal end of left radius (Bat 1.3519), 4 fragmentated right radius (Bat 1.3613), crushed distal end of right carpometacarpus (Bat 5 6 1.6835), part of proximal end of right carpometacarpus (Bat 1.2079), fragmentary right 7 carpometacarpus (Bat 1.2002.5), complete pelvis (Bat 1.2004.92), fragmentary pelvis 8 (Bat 1.2003.154), right femur very deteriorated (Bat 1.2006.297), distal end of left 9 tarsometatarsus (Bat 1.216). Batallones 3 site: three fragmentary crania (Bat 10 3.2007.745, Bat 3.2008.329, Bat 3.2012.1432), medial part of left coracoideum (Bat 11 3.2009.1229), fragmentary right radius (Bat 3.2009.1190), proximal end of left radius 12 (Bat 3.2008.301), distal end of right radius (Bat 3.2012.1216), deteriorated right 13 14 carpometacarpus (Bat 3.2008.673), three right first alar phalanges digiti majoris (Bat 15 3.2007.518, Bat 3.2009.741, Bat 3.2012.5), right tarsometatarsus (Bat 3.2012.173), 16 fragmentary right tarsometatarsus ( Bat 3.2009.1397). Batallones 1 or 3 sites: 17 incomplete left carpometacarpus (B-1969, not “in situ”) 18 19 Type locality: Batallones 1 site Cerro de los Batallones, Torrejón de Velasco village, 20 21 Madrid province, Spain. 22 23 Other locality: Batallones 3 site , Cerro de los Batallones, Torrejón de Velasco village, 24 Madrid province, Spain. Biozone MN 10, early Tortonian, late Miocene (Morales et al. 25 2008, Calvo et al. 2013). 26 27 Stratotype: Upper zone of the “Intermediate Unit” (Morales et al., 1992, sensu 28 29 Lomoschitz et al., 1985, Calvo et al. 2013), biozone MN 10, early Tortonian, late 30 Miocene (Morales et al., 2008). 31 32 Etymology: In honour of M. Dolores (‘Loli’) Soria, excellent person and a 33 paleontologist who, in spite of a short life, became a reference for the study of the 34 Spanish Tertiary. Lolis is the genitive of “Loli” (3rd. declension). 35 36 37 Dimensions and figures: Table 1 and Figures 2 and 3. 38 39 Diagnosis: Different morphologically from the other genera of Gypaetinae and 40 Accipitridae compared. Differences with the extant Neophon percnopterus are more 41 subtle. Differs from the extant Neophron percnopterus in: 1) cranium, slimmer and 42 longer postorbital process; 2) os palatinum, more rounded angulus caudolateralis; 3) 43 44 humerus, more distally protruding epicondylus ventralis; 4) pelvis, pubis completely 45 merged with the ala ischii; 5) ischium wings are short and round; 6) tarsometatarsus, 46 longer, though relatively slender (Table 1) The tarsometatarsus is likewise longer than 47 in Necrosyrtes and Mioneophron longirostris (Table 1). 48 49 Descriptions and comparisons 50 An almost complete skeleton of a gypaetine has been recently named Mioneophron 51 52 longirostris by Li et al. (2016). Although it preserves a remarkable amount of the bones, 53 these are badly broken, many of the anatomical features have been lost or remain hidden 54 in the slab matrix and the magnification of the figures is insufficient. 55 56 Cranium: Four crania, deformed and incomplete, have been unearthed. The rostrum 57 maxillare is elongated and narrow, both dorsoventrally and laterally, and the nostrils are 58 elongated horizontal slits, as in Neophron percnopterus and Mioneophron longirostris, 59 60 and reaches a similar length (Table 1). On the contrary, a robust rostrum maxillare, Page 8 of 24 1 2 3 broad dorsoventrally and hook-shaped, is common in Accipitridae, including Gypaetus 4 and Gypohierax. A rostrum as peculiar as this one also occurs in Necrosyrtes and 5 6 Neophrontops americanus. Two features distinguish the new species from its extant 7 congeneric form. In Mioneophron longirostris, the pre-narial portion of the rostrum 8 maxillare is longer than in Neophron percnopterus (Li et al. 2016, fig. 3a). In the new 9 species, it is proportionally similar. The postorbital process is far more slender and 10 longer in Neophron lolis n. sp. than in Neophron percnopterus. The angulus 11 caudolateralis of the os palatinum is more rounded in the specimens from Batallones 12 than in the latter species. The nares extend towards the end of the beak more in 13 14 Neophron (N. percnopterus and N. lolis n. sp.) than in Mioneophron (Li et al. 2016, fig. 15 3a). 16 17 Humerus: The tuberculum dorsale is fairly developed and detached from the head, like 18 in Gypaetus. It is relatively larger than in Aquila, Gyps, Aegypius, Hieraaetus and 19 Milvus, and smaller than in Pandion. This tubercle is poorly developed in Necrosyrtes 20 and Gypohierax. The distal edge of the crista bicipitalis adopts a lobe-shaped form (Fig. 21 2), as in Neophron, Necrosyrtes and Gypohierax, which is more pronounced than in 22 23 Aquila, Circaetus, Haliaeetus, Hieraaetus and Milvus. This part of the edge of the crista 24 bicipitalis is not lobed in Gypaetus barbatus and G. georgii n. sp. The caput humeri is 25 large and protruding, like in both species of Gypaetus, Neophron, Necrosyrtes and 26 Gypohierax, and more developed than in Mioneophron longirostris. The ventral edge of 27 the crista bicipitalis is almost straight and parallel to the longitudinal axis of the bone. 28 This feature is shared with Neophron, both species of Gypaetus, Aquila, Circaetus, 29 Haliaeetus, Hieraaetus and Milvus, whereas it is not observed in Necrosyrtes and 30 31 Gypohierax. The distal margin of the crista bicipitalis is round and forms a marked step 32 with the shaft, as in Neophron percnopterus. This angle is less marked in Gypaetus 33 barbatus and even less in M. longirostris (Li et al (2016). The epicondylus dorsalis is 34 slightly developed in Gyps, Necrosyrtes and Gypohierax; it is larger in Aegypius, 35 Aquila, Gypaetus, Haliaaetus, Hieraeetus and in the fossil, whereas it is more 36 developed in Neophron percnopterus. In the fossil, in cranial view, the condylus 37 38 ventralis is very prominent with respect to the condylus dorsalis, which likewise is 39 observed in Neophron, Gypohierax, and in Gypaetus in a lesser extent. As in Gypaetus 40 georgii n. sp., one marked furrow that starts in the attachment of the m. latissimus dorsi 41 posterioris and continues on the shaft towards the end of the crista deltopectoralis is also 42 present in Neophron lolis n. sp. This furrow is present in Gypaetinae (Rich 1977), 43 although also in other accipitrids. The distal humerus on which Neophrontops vetustus 44 Wetmore, 1943, from the middle Miocene of Nebraska, was erected is similar in 45 46 morphology to Neophron lolis n. sp., although the epicondylus dorsalis is larger in the 47 latter species. The condylus ventralis is more robust and large in Batallones 1. The 48 flexor process is most acute in N. vetustus. Furthermore, the size of N. vetustus is 49 considerably lower than Neophron lolis n.sp. (Table 1). In turn, N. vetustus is only 50 slightly more than half as large than Neophrontops dakotensis Compton 1935, from the 51 52 early Pliocene of South Dakota (Howard 1966). In Palaeoborus umbrosus, from the 53 early Pliocene of New Mexico, the caput humeri is more developed and the edge of the 54 crista bicipitalis is less round than in N. lolis n. sp. The length of the humerus of 55 Mioneophron longirostris (see Table 1) is very small, both in absolute value and in its 56 proportion with ulna and radius. It may not be correct as the distal end of this bone was 57 manipulated by those who previously possessed the specimen. One of the specific 58 diagnostic traits of Mioneophron longirostris noted by Li et al. (2016) is a well- 59 60 Page 9 of 24 1 2 3 projected and globose caput humeri, although this is also seen in N. percnopterus and N. 4 lolis n. sp. 5 6 In summary, Neophron lolis n. sp, is overall similar in shape and size to the 7 extant Neophron percnopterus, with the following exceptions: (1) epicondylus dorsalis 8 is more distinctive in the extant species, and (2) the epicondylus ventralis protrudes 9 distally more in the fossil. As mentioned in the section on Gypaetus, what is known 10 about the Neophrontops humerus (its distal end) is nothing like that of Gypaetus and 11 Neophron. 12 13 14 Ulna: Only one of the ulnae retains its proximal end. Although it is crushed, the well- 15 marked humero-ulnar depression is clearly visible, as in the extant Gypaetinae. The 16 tuberculum lig. collateralis ventralis is not distinctive, as in N. percnopterus. This 17 tuberculum is quite patent in Gypaetus, Aegypiinae and Anchigyps voorhiesi. The early 18 Miocene Palaeoborus rosatus has a prominent and developed tuberculum carpale, as 19 seen in Aegypiinae and Gypaetinae, but not in the other accipitrids. The characteristic 20 21 deep fossa radialis in Aegypiinae (Manegold et al. 2014) is not observed in P. rosatus. 22 Also in this species, this fossa is shallower, as in Gypaetinae -including the two new 23 species reported in this work-, and is less marked in the other accipitrids. The carpal 24 tuberosity is more pointed and relatively larger in Gypaetinae and Aegypiinae than in 25 the rest of Accipitridae. One such a robust carpal tuberosity is present in N. lolis n. sp. 26 and in P. rosatus. In fact, this tuberosity is more prominent in N. lolis n. sp. than in N. 27 percnopterus. The edge of the condylus dorsalis ulnae is more developed proximally in 28 29 the former than in the latter species. One of the features included in the diagnosis of 30 Mioneophron longirostris is a much longer ulna than tibiotarsus (ratio=1.45, Li et al. 31 2016). However, this same ratio is obtained in Neophron percnopterus. The length of 32 this bone in Mioneophron longirostris is similar to that of Neophron lolis n. sp. and N. 33 percnopterus (Table 1). 34 35 Radius: In size and overall shape of the bone, including curvature and twisting of the 36 37 diaphysis, the fossil is almost indistinguishable from Neophron percnopterus, and is 38 very similar to Gypohierax angolensis. In the new species, the tuberculum aponeurosis 39 ventralis is less pronounced than in N. percnopterus, and looks alike in G. angolensis. 40 Morphological differences with Necrosyrtes monachus are sharped. In particular, the 41 depressio ligamentosa on the distal end is smooth in the fossil and very deep in N. 42 monachus. Likewise, the fossil shares with these three living vultures a cotyla humeralis 43 44 round, one feature exclusive among the accipitrids compared. 45 46 Carpometacarpus: The three carpometacarpi found are very damaged. Nevertheless, 47 some features can be seen. The proximal junction between the ossa metacarpi majus and 48 minus takes place relatively distally, as observed in Gypaetinae and Aegypiinae. The 49 ventral edge of the proximal end of the trochlea carpalis is round, like in Neophron 50 percnopterus, Gypohierax and Necrosyrtes. In Gypaetus barbatus and G. georgii n. sp., 51 52 this edge is flattened. Necrosyrtes presents a pneumatic fossa between the os 53 metacarpale minus and the fovea carpalis caudalis that N. percnopterus, Gypohierax and 54 the fossil do not. 55 56 Manus, phalanx proximalis digiti majoris: This element in the new species is hardly 57 distinguishable in overall shape and size from N. percnopterus. In both taxa, the 58 proximal articular surface is triangular-like shaped whereas it is elongated in 59 60 Necrosyrtes and Gypohierax. Neophron percnopterus differs from the fossil because in Page 10 of 24 1 2 3 cranial view this bone is straight in the former, and slightly curved dorsally in the new 4 taxon. 5 6 7 Pelvis: It is similar in size and proportions to N. percnopterus (Figure 3). In lateral 8 view, the ala ischii is shorter and more rounded in the new species than in N. 9 percnopterus. The ilioschiatic, obturator and acetabulum foraminae are larger in the 10 extant Neophron, but in the fossil the pubis is more robust and is fused with the 11 ischium. The iliosynsacral sulcus is intermediate in width between N. percnopterus and 12 Mioneophron longirostris (Li et al. 2016, fig. 4). Vertebrae of the caudal region of the 13 14 synsacrum are higher in N. percnopterus. Batallones, the trochlea metatarsi III is relatively strong and robust and its outline is 15 16 nearly circular in lateral view, trait also observed in G. barbatus.. In Gypohierax, this 17 trochlea is relatively much smaller. The sulcus extensorius is shallow like in Gyps and 18 Aegypius, although not as smooth like in Gypaetus, Neophron percnopterus and 19 Necrosyrtes. This sulcus is deep in Gypohierax, Circaetus, Aquila, Haliaeetus and 20 Hieraaetus. Unlike in G. barbatus, the tuberositas musculi tibialis cranialis is little 21 developed and is situated close to the foramina vascularia proximalia. The crista 22 23 lateralis hypotarsi is very small, like in Gypohierax and Neophron. In distal view, the 24 trochleae metatarsorum are alligned in Gypohierax, whereas they form a curve in the 25 fossil, Necrosyrtes, G. barbatus and N. percnopterus. With regard to the other trochleae, 26 the trochlea metatarsi III is small in Gypohierax, very large in G. barbatus, Necrosyrtes, 27 and intermediate size in N. percnopterus and in the new species. The tarsometatarsus is 28 larger than in the extant Gypaetus G. barbatus, Neophron percnopterus, Gypohierax 29 Tarsometatarsus: This bone is slender as in Circaetus, Neophron, Necrosyrtes, 30 31 Gypohierax and Anchigyps voorhiesi (Table 1), not robust like in Gyps, Aegypius, 32 Haliaeetus, Gypaetus, and in some species of Aquila, as A. chrysaetos and A. heliaca. 33 Size and robustness of the fossil are similar in the extant Circaetus gallicus. In most 34 accipitrids trochlea metatarsi II protrudes distally from trochlea metatarsi III. On the 35 contrary, trochlea III is far more protruding in half of the Neophron percnopterus 36 specimens examined (n = 6), as well as in those of N. lolis n. sp., in Gypaetus barbatus 37 (the tarsometatarsus of Gypaetus georgii n. sp. is not recorded) and in Necrosyrtes. 38 39 Following the pattern of most Accipitridae, trochlea metatarsi II exceeds trochlea 40 metatarsi III distally in Neogyps, Neophrontops (Miller 1916; Olson and Rasmussen 41 2001), Anchigyps voorhiesi (Zhang et al. 2012a) and Gypohierax. In the specimens from 42 43 44 angolensis and Necrosyrtes monachus. The tarsometatarsus of Palaoeborus umbrosus 45 shows a more robust appearance than in N. lolis n. sp. In distal view, the trochleae 46 describe a more curved line in N. lolis and N. percnopterus than in P. umbrosus and P. 47 howardae (Storrs Olson, comm. pers. 2008). 48 49 Remarks: A more prominent distally metatarsal III is not autapomorphic within 50 Gypaetinae because such a trait is present in Necrosyrtes and not in Gypohierax. 51 52 However, this feature seem to be only seen in Neophron (N. lolis n. sp. included), 53 Gypaetus and Necrosyrtes among accipitrids. The bones of the new species bear a 54 greater general resemblance to the three living vultures gypaetines of small size: 55 Neophron N. percnopterus, Necrosyrtes N. monachus and Gypohierax G. angolensis. 56 The sulcus extensorius tarsometatarsi is deep in more predatory accipitrids, meanwhile 57 it is smooth in vultures and Gypaetus. In connection with this character, the facies 58 subcutanea lateralis is wide and flat in hunting species, and curved in scavengers. These 59 60 anatomical features may be ment Neophron lolis n. sp. had a more predatory behavior Page 11 of 24 1 2 3 than its extant congeneric species. However, since the beak of the new species is similar 4 in size and robustness to that of the current Neophron percnopterus, it can be assumed 5 6 that the kinds of food consumed by Neophron lolis n. sp. and its strategies for accessing 7 it were not very different from those performed by the extant Neophron N. 8 percnopterus. 9 10 11 Discussion 12 The late Miocene paleontological complex of Cerro de los Batallones is singular for 13 14 having yieded a wide variety of avian bones, mainly of accipitriforms. Not all 15 specimens have been preserved in good condition, but the richness of the deposits 16 allows characterizing and describing a certain number of the implied taxa. Two diurnal 17 raptors and other species are pending description. The birds from these sites bear 18 marked morphological similarities with their living relatives, which is indicative of their 19 respective close phylogenetic affinities, contrasting with what happens with the 20 21 mammal taxas found. The finding of sympatric fossil vultures is not unusual (Zhang et 22 al. 2012b) 23 Six species have been distinguished within Neophrontops. All of them are based 24 on scant and fragmentary remains, except N. americanus. An important differentiating 25 feature is size. Neophrontops N. dakotensis (early/middle Pliocene) and Neophrontops 26 N. vetustus (middle Miocene) are each represented by a single distal end of humerus 27 (Compton 1935; Wetmore 1943). Both specimens differ only in size, the former being 28 29 almost twice as large as the latter one, and both are considerably smaller than 30 Neophrontops N. americanus (early Miocene/late Pleistocene, Rich 1977). If the 31 paleospecies N. americanus had corresponded to one biological species, it would have 32 had an extraordinary vast existence. In the other extreme are two forms reaching similar 33 sizes, N. slaughteri (late Pliocene) and N. vallecitoensis (middle Pleistocene). N. 34 americanus reached intermediate dimensions (Wetmore 1943; Howard 1963; Feducia 35 1974; Rich 1977). From the above comparisons with the fossil bones from Batallones, it 36 37 follows that Palaeoborus and Neophrontops have anatomical features compatible with a 38 close relationships between them and to the Gypaetus and Neophron genera. However, 39 some of the characters observed in the humerus of Neophrontops and summarized by 40 Rich (1977) –such as a fossa m. brachialis triangular, the fossa pneumotricipitalis 41 ovoid-shaped with its tapering end dorsal, or sulcus transversus deep-, that it should 42 share with the rest of Gypaetinae, are neither exclusive to this group nor are they present 43 44 in all extant Gypaetinae. The finding of Neophron in the European Miocene reduces 45 support for the possibility that this genus had its origin in a stock of vultures from North 46 America. Furthermore, the shape of the tarsometatarsus and particularly the 47 arrangement of the trochlea metatarsi III with respect to the others trochleae not only 48 confers a more eagle-like appearance to Palaeoborus and Neogyps with respect to 49 Neophrontops, as noted by some authors (Miller and Crompton 1939), but also with 50 regard to Gypaetus and Neophron. The mandibular symphysis in Anchigyps voorhiesi is 51 52 strong and broad (Zhang et al. 2012a) which conflicts with the thin and slender aspect 53 that this region has in the extant Gypaetinae, Mionephron longirostris (Li et al. 2016), 54 Neophrontops americanus and surely in Neophron lolis n. sp., judging from its 55 maxillary jaws recovered –the mandible of Gypaetus georgii n. sp. is not preserved-. 56 The absence of pneumatic foramina at both ends of the ulna of Anchigyps voorhiesi and 57 regarded as a feature distinguishing Gypaetinae from Aegypiinae, is also present in both 58 subfamilies (on the impressio brachialis, the incisura radialis, and the distal radial 59 60 depression). If the published humerus length (Li et al., 2016) is right, this bone was Page 12 of 24 1 2 3 certainly very short, and would have consequences on the flight adaptations of this 4 species. 5 6 As regards Mioneophron longirostris, the part of the rostrum maxillare that is 7 exposed and the details of the pelvis, which preserves part of its integrity, undoubtedly 8 belong to a gypaetine closely related to Neophron. It has been considered parsimonious 9 to include the new species Neophron lolis n. sp. within de genus Neophron because the 10 anatomical differences between N. lolis n. sp. and N. percnopterus do not seem to 11 justify the naming of a new genus. Due to the poor state of preservation of the 12 Mioneophron M. longirostris specimen, it is difficult to assess the extent of the 13 14 anatomical differences between Mioneophron and Neophron. Referring the new species 15 to the genus Mioneophron because it has been described in the late Miocene and that 16 there was no evidence of Neophron prior to the Pleistocene is out of consideration 17 because taxonomic decisions have to be based only on intrinsic characteristics of the 18 organisms. Although it would be expected that late Miocene finds should be assigned to 19 genera other than the extant Gypaetus and Neophron, their respective morphological 20 21 differences with these are subtle, insufficient to erect new genera. 22 The forelimb to hindlimb ratio is lower in both new raptors described here than 23 in their extant congeners. Both Gypaetus barbatus and Neophron percnopterus perform 24 gliding flight, and presumably the extinct forms did too. Being likely Gypaetus georgii 25 n. sp. and Neophron lolis n. sp. worse fliers than their current close relatives, and since 26 flight capability is subjected to maximum stress during migration, it is possible that the 27 two new species were not migratory. The relative large size of the tarsometatarsus in 28 29 Neophron lolis n. sp. could point to a more predatory behavior than its extant 30 congeneric form. This would be strengthened if the femur and tibiotarsus, which have 31 not been recovered, show similar proportions with N. percnopterus. Apart from what 32 concerns its fligth capability, the morphological features of Gypaetus georgii shows no 33 evidence that such a species behaved very differently from the extant Gypaetus 34 barbatus. 35 36 37 38 Acknowledgements 39 My thanks to Storrs L. Olson for enabling my work in the USNM some time ago and 40 for sharing his notes on Neophrontops. Criticisms by Marcos Cenizo and another three 41 other anonymous reviewers have greatly improved the manuscript. I am also indebted to 42 Ana García for helping me with the Latin and to Arturo Morales for making it easier for 43 44 me to use his comparison collection at Universidad Autónoma de Madrid. 45 46 47 References 48 49 Baumel JJ, Witmer LM. 1993. Osteologia. In: Baumel JJ, King AS, Breazile JE, Evans 50 51 52 HE, Vanden Berge JC, editors. Handbook of avian anatomy. Publ Nuttall 53 54 Ornithol Club, 23:45-132. 55 56 Brodkorb P. 1964. Catalogue of fossil birds, part 2 (Anseriformes through Galliformes). 57 58 Bull Florida State Mus, Biol Sc, 8(3):195-335. 59 60 Page 13 of 24 1 2 3 Brodkorb P. 1978. Catalogue of fossil birds, part 5 (Passeriformes). Bull Florida State 4 5 6 Mus, Biol Sc, 23(3):139-228. 7 8 Calvo JP, Pozo M, Silva PG, Morales J. 2013. 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Gypaetus georgii n. sp.: scapula, (a) dorsal view; carpometacarpus (holotype, 11 Bat 3.2012.437), (b) ventral view; humerus, (e) caudal view; radius, (f) ventral view; 12 tibiotarsus, (g) cranial view, (h) distal view. Gypaetus barbatus: carpometacarpus, (c) 13 14 ventral view; humerus, (d) caudal view; tibiotarsus, (i) distal view. Abbreviations: ed – 15 epicondylus dorsalis, ii – incisura intercondylaris, pe – processus extensorius. Scale bar: 16 50 mm. 17 18 Figure 2. Neophron lolis n. sp.: cranium, (a) dorsal view, (b) lateral view; 19 tarsometatarsus (holotype, Bat 1.3087), (d) dorsal view; ulna, (f) dorsal view; humerus, 20 21 (g) caudal view. Neophron percnopterus: cranium, (c) lateral view; tarsometatarsus, (e) 22 dorsal view; humerus, (h) caudal view. Abbreviations: cb – crista bicipitalis, ev – 23 epicondylus ventralis, n – nostrils, tm II – trochlea metatarsi II, tm III – trochlea 24 metatarsi III. Scale bar: 50 mm. 25 26 Figure 3. Pelvis of Neophron lolis n. sp.: (a) dorsal view; (b) lateral view. Pelvis of 27 Neophron percnopterus: (c) dorsal view; (d) lateral view. Abbreviations: is – 28 29 iliosynsacral sulcus, iw – ala ischii. Scale bar: 50 mm. 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 Page 18 of 24 1 2 Table 1. Measurements (in mm) of bones of Gypaetus georgii n. sp. and Neophron lolis n. sp. compared with 3 other extinct and extant Gypaetinae taxa. 4 5 6 Maximum Width of the Width of the distal Width of the 7 length proximal end end diaphysis at 8 midpoint 9 Gypaetus georgii, n.sp. 10 scapulae 21.8, 23.1 11 12 humeri ca. 190, 190.7 39.6 32.7, 35.5 13 radii 214.5 9.6, 10.0 14.6 ca. 5.4 14 15 carpometacarpi 93.4 – 102.1 (3) 22.3 – 26.0 (3) 14.2, 16.0 5.9 – 6.9 (3) 16 phal. alar 1 dig. maj. 38.4, 38.9 17 18 tibiotarsi 154.8 24.8 19.8, 20.6 19 20 21 Gypaetus barbatus 22 scapulae MNCN-23330, 25.7 – 28.1 (4) 23 CAM-3, NMNH-17834, 24 345684 25 26 humeri MNCN-23330, 215.0 – 236.6 40.9 – 43.8 (3) 34.5 – 40.0 (4) 15.8 – 17.9 (4) 27 CAM-2, 3, NMNH- (4) 28 17834, 345684 29 30 radii MNCN-23330, 233.0 – 254.7 10.7 – 11.7 (3) 17.2 – 18.1 (3) 5.9 – 6.0 (3) 31 NMNH-17834, 345684 (3) 32 carpometacarpi MNCN- 110.1 – 123.6 28.0 – 31.3 (3) 19.1 7.2 – 8.3 (3) 33 23330, NMNH-17834, (3) 34 345684 35 36 phal. alar 1 dig. maj. 50.8 37 MNCN-23330 38 tibiotarsi MNCN- 157.0 – 171.6 10.9 – 12.5 (3) 21.0 – 21.5 (3) 10.9 – 16.6 (3) 39 23330, MNNH-17834, (3) 40 345684 41 42 43 Neophron lolis n. sp. 44 45 crania > 95 46 humeri ca. 151.7 28.2 24.7 47 48 ulnae ca. 180.0 14.2 – 14.3 (3) 7.1, 8.4 49 radii ca. 168.7 7.6 – 8.5 (4) 11.1 – 11.9 (4) 4.6 – 5.1 (5) 50 51 carpometacarpi ca. 81.0 17.5 5.3 52 53 phal. alar 1 dig. maj. 34.7 – 37.0 54 sacra 75.2, 77.0 55 56 tarsometatarsi 86.9, 93.5 16.1, 16.2 17.2, 17.7 8.2, 8.3 57 58 59 Neophron percnopterus 60 cranium MNCN-26843, 96.4 – 106.3 CAM-2, 3, 10 Page 19 of 24 1 2 humeri MNCN-26843, 118.8 – 151.6 29.9 – 31.5 (2) 25.1 – 25.7 (3) 10.6 – 11.6 (3) 3 CAM-2, 3 (2) 4 5 ulnae MNCN-26843, 175.5 – 179.0 16.9 – 17.5 (3) 14.6 – 14.7 (3) 8.0 – 8.5 (3) 6 CAM-2, 3 (3) 7 radii MNCN-26843, 168.2 – 171.9 7.7 – 8.2 (3) 12.1 – 12.9 (3) 3.9 – 5.4 (3) 8 CAM-2, 3 (3) 9 10 carpometacarpi MNCN- 76.7 – 83.3 (3) 19.6 – 21.2 (3) 12.0 – 12.7 (3) 5.0 – 5.3 (3) 11 26843, CAM-2, 3 12 phal. alar 1 dig. maj. 33.5 – 36.1 (3) 13 14 MNCN-26843, CAM-2, 15 3 16 sacra MNCN-26843, 77.2, 86.7 17 CAM-3 18 19 tibiotarsi MNCN-26843 129 20 tarsometatarsi MNCN- 76.7 – 83.5 (3) 16.4 – 17.1 (3) 16.9 – 17.9 (3) 7.3 – 8.1 (3) 21 26843, CAM-2, 3 22 23 24 Gypohierax angolensis 25 26 humeri NMNH-226143, 140.2 – 145.6 26.2 – 28.2 (4) 21.9 – 23.9 (4) 27 291316, 291078, (4) 28 224820 29 radii NMNH-226143, 150.1 – 155.5 6.8 – 8.1 (4) 9.7 – 10.7 (4) 3.4 – 4.1 (4) 30 291316, 291078, (4) 31 224820 32 33 carpometacarpi NMNH- 16.8 – 18.1 (4) 4.0 – 4.2 (4) 34 226143, 291316, 35 291078, 224820 36 37 phal. alar 1 dig. maj. 28.3 – 29.6 (4) 38 NMNH-226143, 39 291316, 291078, 40 224820 41 tarsometatarsi NMNH- 76.5 – 78.4 (4) 14.3 – 15.1 (4) 15.6 – 16.6 (4) 7.8 – 8.9 (4) 42 226143, 291316, 43 291078, 224820 44 45 46 Mioneophron 47 longirostris 48 49 cranium 113.2 50 51 humerus 146.0 52 ulna 180.9 53 54 radius 169.0 55 carpometacarpus 82.0 56 57 phal. alar 1 dig. maj. 33.5 58 sacrum 84.8 59 60 femur 80.4 tibiotarsus 124.7 Page 20 of 24 1 2 tarsometatarsus > 74.4 3 4 5 Necrosyrtes monachus 6 7 humeri NMNH-18894, 148.6 – 163.9 28.4 – 32.0 (5) 25.5 – 27.2 (5) 8 291440, 291441, (5) 9 291442, 291443 10 radii NMNH-18894, 180.2 – 193.8 6.6 – 9.0 (5) 11.8 – 13.5 (5) 4.1 – 5.2 (5) 11 291440, 291441, (5) 12 291442, 291443 13 14 carpometacarpi NMNH- 20.3 – 22.1 (5) 5.0 – 6.3 (5) 15 18894, 291440, 291441, 16 291442, 291443 17 18 phal. alar 1 dig. maj. 32.2 – 37.0 (5) 19 NMNH-18894, 291440, 20 291441, 291442, 21 291443 22 tarsometatari NMNH- 75.4 – 83.0 (5) 14.9 – 15.4 (5) 15.8 – 17.3 (5) 7.4 – 8.4 (5) 23 18894, 291440, 291441, 24 291442, 291443 25 26 27 Anchigyps voorhiesi 28 29 ulna (Zhang et al. 127.5 15.5 12.4 7.1 30 2012) 31 tibiotarsus (Zhang et al. ca. 118.3 13.3 7.4 32 33 2012) 34 tarsometatarsi (Zhang 84.5, 82.6 14.4, 13.9 15.8 6.5, 6.6 35 et al. 2012) 36 37 38 Neophrontops vetustus 39 40 humerus (Wetmore 17.1 41 1943) 42 humerus (taken on cast 19.2 43 USNM 16814, 44 unpublished) 45 46 47 Neophrontops 48 americanus 49 50 humerus (Howard 148 51 1932) 52 humeri (Rich 1977) 134 - 146 53 54 55 56 Palaeoborus umbrosus 57 tibiotarsus (in Howard 16.0 58 1966) 59 60 tarsometatarsus (Cope ca. 19 ca. 10 1874) Page 21 of 24 1 2 phal. alar 1 dig. maj. ca. 37 3 (Cope 1874) 4 5 6 Palaeoborus howardae 7 8 tarsometatarsus 20.4 9.7 9 (Wetmore 1936) 10 11 12 Neogyps errans 13 tibiotarsus (in Howard 18.0, 19.4, 20.8 14 1966) 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 URL: http://mc.manuscriptcentral.com/ghbi Page 22 of 24 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Fo 17 18 19 rP 20 21 22 23 ee 24 25 26 rR 27 28 29 ev 30 31 32 iew 33 34 35 36 37 Figure 1. Gypaetus georgii n. sp.: scapula, (a) dorsal view; carpometacarpus (holotype, Bat 3.2012.437), On 38 (b) ventral view; humerus, (e) caudal view; radius, (f) ventral view; tibiotarsus, (g) cranial view, (h), distal 39 view. Gypaetus barbatus: carpometacarpus, (c) ventral view; humerus, (d) caudal view; tibiotarsus, (i) 40 distal view. Abbreviations: ed – epicondylus dorsalis, ii – incisura intercondylaris, pe – processus 41 extensorius. Scale bar: 50 mm. ly 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Page 23 of 24 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Fo 17 18 19 rP 20 21 22 23 ee 24 25 26 rR 27 28 29 ev 30 31 32 iew 33 34 35 36 37 On 38 39 40 41 ly 42 43 44 45 Figure 2. Neophron lolis n. sp.: cranium, (a) dorsal view, (b) lateral view; tarsometatarsus (holotype, Bat 46 1.3087), (d) dorsal view; ulna, (f) dorsal view; humerus, (g) caudal view. Neophron percnopterus: cranium, 47 (c) lateral view; tarsometatarsus, (e) dorsal view; humerus, (h) caudal view. Abbreviations: cb – crista bicipitalis, ev – epicondylus ventralis, n – nostrils, tm II – trochlea metatarsi II, tm III – trochlea metatarsi 48 III. Scale bar: 50 mm. 49 50 51 52 53 54 55 56 57 58 59 60 Page 24 of 24 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Fo 17 18 19 rP 20 21 22 23 ee Figure 3. Pelvis of Neophron lolis n. sp.: (a) dorsal view; (b) lateral view. Pelvis of Neophron percnopterus: 24 (c) dorsal view; (d) lateral view. Abbreviations: is – iliosynsacral sulcus, iw – ala ischii. Scale bar: 50 mm. 25 26 rR 27 28 29 ev 30 31 32 iew 33 34 35 36 37 On 38 39 40 41 ly 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60