This is the accepted version of the article: Sánchez Marco, Antonio. «Old and new fossil birds from the Spanish Miocene». Journal of Iberian Geology, Vol. 47, Issue 4 (December 2021), p. 697-712. DOI 10.1007/s41513-021-00178-9 This version is available at https://ddd.uab.cat/record/250772 under the terms of the license 1 Old and new fossil birds from the Spanish Miocene 2 3 Antonio Sánchez-Marco. Institut Català de Paleontologia Miquel Crusafont, Campus UAB, 08193 4 Bellaterra, Spain, antonio.sanchez@icp.cat. 5 ORCID: 0000-0003-0654-1935 6 7 Abstract 8 Now a long time ago, avian remains from the Vallès-Penedès Basin and from other Miocene localities 9 inaugurated the study of the Tertiary birds from Spain. After so much time, it is suitable to review them. 10 Fossils unknown so far of the same or close taxa from the rest of the Iberian Peninsula are also presented 11 in this paper. The majority of species belong to Phasianidae. The present paper participates in a long 12 discussion about the systematic validity of the various fossil species of Tyto described in the European 13 Neogene. A considerable part of such discussion has as focal point Tyto balearica. Apparently, 14 discrepancies have been raised around the size range of this species, although a conviction of the 15 chronological and geographical distributions of this owl, in particular, if it were essentially an insular 16 species, has likely had an unwanted influence on a debate that should be restricted to anatomical aspects. 17 18 Keywords Palaeornithology, avian systematics, avian paleontology, Neogene birds, paleo-subspecies. 19 20 Resumen 21 Hace mucho tiempo, los restos de aves de la cuenca del Vallés-Penedés y de otras localidades del 22 Mioceno inauguraron el estudio de las aves del Terciario de España. Después de tanto tiempo, conviene 23 su revisión. También se presentan en este artículo restos desconocidos hasta ahora de los mismos taxones, 24 o próximos, procedentes del resto de la península ibérica. La mayor parte de las especies pertenecen a 25 Phasianidae. Este artículo participa en una larga discusión acerca de la validez sistemática de varias 26 especies fósiles de Tyto descritas en el Neógeno europeo. Una parte considerable de esta discusión tiene a 27 Tyto balearica como punto focal. En apariencia, la discusión se ha planteado en torno al rango de talla de 28 esta especie, si bien la convicción de las distribuciones cronológicas y geográficas de esta lechuza, en 29 particular, si fue una especie esencialmente insular, probablemente ha tenido una influencia indeseada en 30 un debate que debería estar restringido a aspectos anatómicos. 1 31 32 Palabras clave Paleornitología, sistemática aviar, paleontología de aves, aves del Neógeno, paleo- 33 subespecies. 34 35 Declarations 36 Funding: This work has been benefited from the grants: 2017SGR86, Generalitat de Calalunya, and the 37 project CGL2016-76431-P of the State Research Agency and the European Fund for Regional 38 Development. 39 Conflicts of interest/Competing interests: There is not any conflits of interest or competing interests. 40 Availability of data and material: Not applicable. 41 42 2 43 1 Introduction 44 45 In contrast to the abundant literature on Miocene mammals, proof of the interest in this animal 46 group and its extensive fossil record, the fossil birds from the Iberian Miocene have attracted little 47 attention from paleontologists. Most likely, this is due to the low number of fossil localities from this 48 epoch yielding large collections of avian remains. Thus, with the exception of a few relatively rich 49 deposits, Miocene avian remains from the Iberian Peninsula have come to light in small numbers and in a 50 fragmentary state, which sheds uncertainties to subsequent taxonomic tasks. Available data on fossil finds 51 and paleontological sites from the Iberian Miocene were summarized in three previous paleornithological 52 catalogs (Sánchez Marco 1995a, 1995b, 1999) and in a subsequent compilation paper (Sánchez and Sastre 53 2001). One aim of the present paper is to report new avian remains from the Miocene of the Iberian 54 Peninsula and critically revise previous studies on this topic. 55 Miocene avian remains from Iberian localities have been dealt with in a handful of works, 56 beginning with some of the earliest Spanish papers in this field (Villalta and Crusafont Pairó 1950; 57 Villalta 1963). After this time, there are only two publications of limited scope on this subject (Sánchez 58 Marco 2001, 2006). A number of misidentifications arisen in early works had settled in the specialized 59 literature, and such errors required a critical review. 60 The extinct owl Tyto balearica has been found in two new localities (this paper). This species 61 has been misinterpreted in some recent publications and requires additional clarification concerning the 62 understanding of what a taxon described on paleontological grounds means. Overall, this paper deals with 63 a considerable part of the current Iberian record of Miocene galliforms, which shows how meagre it is so 64 far. 65 66 2 Materials and Methods 67 68 For most of the mammal units and for the Neogene mammal ages, it has been followed de Bruijn 69 et al. (1992), and Calvo et al. (1993) for the continental stages. The osteological nomenclature follows 70 Baumel and Witmer (1993) and Livezey and Zusi (2006). Most of the measurents on phasianid coracoidea 71 follow Göhlich and Mourer-Chauviré (2005). The fossil remains studied in this work are housed in the 72 MNCN and the ICP. 3 73 Anatomical abbreviations: Gl, greatest length; Pw, proximal width; Pd, proximal depth; Wd, 74 smallest width of the diaphysis; Dw, distal width (abbreviated as Wd in Göhlich and Mourer-Chauviré 75 2005); Dd, distal depth; Lm, medial length; Wdf, width of distal facies; Lp, proximal length from distal 76 margin of facies articularis humeralis to the proximal end; Wpa, width of processus acrocoracoideus. 77 Institutional abbreviations: IPS, Museu del Institut Català de Paleontologia Miquel Crusafont 78 (Sabadell, Spain); MNCN, Museo Nacional de Ciencias Naturales (Madrid, Spain). 79 Age of the localities: The ages for the localities of provenance of the studied material have been 80 taken from the following published sources: Can Mas (el Papiol, Vallès-Penedès Basin; Agustí et al. 81 1985); Can Poncic (Sant Quirze, Vallès-Penedès Basin; Casanovas-Vilar et al. 2011a); El Fallol (Vallès- 82 Penedès Basin; Casanovas-Vilar et al. 2011b); Hostalets de Pierola indeterminate (els Hostalets de 83 Pierola, Vallès-Penedès Basin; Casanovas-Vilar et al. 2011a); Viladecavalls (Vallès-Penedès Basin; Alba 84 et al. 2010). Batallones 1 and 3 (Cerro de los Batallones, Torrejón de Velasco, Madrid; Morales et al. 85 2008); La Artesilla (Villafeliche, Zaragoza; de Bruijn et al. 1992); Toril 3 (Daroca, Zaragoza; de Bruijn et 86 al. 1992); Escobosa (Escobosa de Calatañazor village, Soria; de Bruijn et al. 1992); Valdecebro 5 87 (Valdecebro, Teruel Basin; Adrover et al. 1986; Sánchez Marco 1999); Los Mansuetos (Teruel, Teruel 88 Basin; de Bruijn et al. 1992; Calvo et al. 1999); Los Valles de Fuentidueña (Segovia; de Bruijn et al. 89 1992). 90 91 3 Systematic palaeontology 92 93 Order Galliformes Linnaeus, 1758 94 Family Phasianidae Vigors, 1825 95 96 Miophasianus altus (Milne-Edwards, 1869) 97 (Figure 1a–f) 98 99 Phasianus altus Milne-Edwards, 1869, vol. 2, p. 239, pl. 131, fig. 27-36 (original description). 100 p. Miophasianus altus Milne-Edwards, 1869: Villalta and Crusafont 1950:147, fig. 1. 101 p. Miophasianus altus Milne-Edwards, 1869: Villalta 1963: 271, pl. 4-5. 102 Miophasianus altus Milne-Edwards, 1869: Ballmann 1969: 175, pl. 15. 4 103 Miophasianus altus Milne-Edwards, 1869: Cheneval 2000: 351, fig. 9-11. 104 p.? Miophasianus altus Milne-Edwards, 1869: Göhlich 2002: 178, fig. 1-4. 105 106 Localities. Can Mas (el Papiol, MN4, early Aragonian); Hostalets de Pierola indeterminate (els 107 Hostalets de Pierola; MN7+8 to MN9, late Aragonian to early Vallesian). 108 Referred specimens. Can Mas: IPS12671 (Figure 1c), proximal end of left ulna; IPS12672 109 (Figure 1d), distal end of right ulna; IPS12670 (Figure 1e), proximal end of right femur; IPS12668 110 (Figure 1f), distal end of right tibiotarsus. Hostalets de Pierola indeterminate: IPS12649 (Figure 1b), 111 distal end of left humerus; IPS12651 (Figure 1a), distal end of right humerus. 112 Description. The humerus show a smooth sulcus humerotricipitalis, as it is common within the 113 family. However, this sulcus is deeper than in Gallus Brisson, 1766, and Phasianus Linnaeus, 1758. The 114 condylus ventralis is more protruding distally regarding the condylus dorsalis than in Gallus and 115 Phasianus. The cross-section of the diaphysis is elliptical. The fossa musculi brachialis is marked, but not 116 deep. The processus flexorius is rounded and scarcely protruding. With regard to the ulna, as it is typical 117 in Galliformes, the cotylae ventralis and dorsalis form a continuous shallow surface. The region of the 118 olecranon is curved dorsally, and the olecranon is slightly protruding. The cross-section of the diaphysis 119 is oval, cranially sharper. The proximal end has no distinctive characteristics. The condylus ventralis 120 ulnae is poorly developed and protrudes distally. This condylus is more protruding than in Gallus, and as 121 jutting as in Phasianus. In distal aspect, the ulna of Phasianus has a more flattened shape than in Gallus 122 and Miophasianus. The tuberculum carpale is small and rounded. The condylus dorsalis is large and 123 round, which gives a triangular outline to the bone in distal view. The single available femur lacks its 124 head, making it difficult to compare with other taxa. It has the typical wide and proximally protruding 125 trochanter of galliforms. The fossa trochanteris is large and curved. The section of the bone is elliptical. 126 In the tibiotarsus, both condyles are robust and well developed, being the lateral one the largest. In both 127 condyles, the distal edges are more flattened than in Gallus and Phasianus. The incisura intercondylaris is 128 wide and notably deeper than in the two mentioned genera. 129 Measurements. See Table 1. 130 Remarks. Miophasianus altus was originally described by Milne-Edwards (1869) based on fossil 131 remains from Sansan (MN6, Astaracian, France) as Phasianus altus. Lambrecht (1933) erected the genus 132 Miophasianus for the Miocene species previously allocated to Phasianus. Mlíkovský (2002) noticed that 5 133 the name Miophasianus was a nomen nudum because Lambrecht (1933) failed to indicate a type species 134 (required for names after 1930). Nevertheless, the name Miophasianus was made available by Brodkorb 135 (1952). At the mentioned work, Lambrecht (1933) erected the new genus Miogallus, with the type species 136 Miogallus longaevus (Ammon, 1918), on material from the middle Miocene of Dechbetten (Germany), a 137 taxon that was later synonymized by Mlíkovský (2002) with Miophasianus altus. However. Mlíkovský 138 (2002) did not examined directly the skeletal elements of both taxa. Therefore, its synonymy has to be 139 revised. 140 Villalta and Crusafont Pairó (1950) mentioned four species of galliforms from Hostalets de 141 Pierola: Miophasianus altus, M. medius, Palaeortyx miocaena and Palaeortyx edwardsi. Later, Villalta 142 (1963) incorporated Palaeoperdix sansaniensis to the record of this site. Likewise, Villalta (1963) 143 reported the find of two galliforms in Can Mas: Miophasianus altus and Palaeortyx edwardsi. Part of this 144 material was restudied by Sánchez Marco (2006), who agreed with most of the previous assignments by 145 Villalta and Crusafont Pairó (1950). Miophasianus altus has been also identified in some other sites 146 outside the Vallès-Penedès: La Grive-Saint-Alban (MN7+8; Déperet 1887; Lydekker 1893; Gaillard 147 1939; Ballmann 1969) in France; Attenfeld (MN7+8; Schlosser 1916), Dechbetten (MN5; Ammon 1918, 148 as Miophasianus augustus), Oehningen (MN7+8; Lydekker 1891), Sandelzhausen (MN5; Göhlich 2002), 149 and Steinheim (MN7+8; Heilzmann and Hesse 1995) in Germany; Děvinská Nová Ves (MN6; Švec 150 1986) in Slovakia; Gratkorn (MN7+8; Göhlich and Gross 2014) in Austria; and Toril 3 (Sánchez Marco 151 2006) in central Spain. Moreover, Cheneval (2000) mentioned that the identification of Miophasianus sp. 152 from the French locality of Vieux Collonges (MN5) by Ballmann (1972) might correspond to 153 Miophasianus altus, and the same could be said about the fossil material from the Spanish locality of 154 Córcoles (MN 4) (Alférez et al. 1982). 155 156 [PLACE FIGURE 1 ABOUT HERE] 157 158 Miophasianus medius (Milne-Edwards, 1869) 159 (Figure 1g,h) 160 161 Phasianus medius Milne-Edwards, 1869, vol. 2, p. 242, pl. 131, fig. 24-26 (original description). 162 p. Miophasianus altus Milne-Edwards, 1869: Villalta and Crusafont 1950:147. 6 163 Miophasianus medius Milne-Edwards, 1869: Villalta and Crusafont 1950:149, fig. 2. 164 p. Miophasianus altus Milne-Edwards, 1869: Villalta 1963:271. 165 Miophasianus medius Milne-Edwards, 1869: Villalta 1963:272, pl. V, fig. 3, 3a. 166 Miophasianus medius Milne-Edwards, 1868: Ballmann 1969:176, pl. 15. 167 Miophasianus medius Milne-Edwards, 1869: Bocheński 1987:71, pl. XVII. 168 Palaeoperdix medius Milne-Edwards, 1869: Cheneval 2000:349, fig. 8 (new combination). 169 Palaeoperdix medius Milne-Edwards, 1869: Sánchez Marco 2006:253. 170 “Miogallus” medius Milne-Edwards, 1869: Zelenkov 2017:74, fig. 3. 171 172 Localities. Hostalets de Pierola (MN7+8-MN9), Batallones 3 (MN 10). 173 Referred specimens. Hostalets de Pierola indeterminate: IPS12648 (Figure 1g), proximal end of 174 left humerus; without number, distal end of another –right- humerus, not seen for this study, although 175 Villalta and Crusafont Pairó (1950, fig. 1) figured it and gave its distal width (17.3 mm). Batallones 3: 176 2008-326 (Figure 1h), complete right radius. 177 Description. The head of the humerus is robust and rounded. The crista deltopectoralis is short 178 and slightly developed. It is curved ventrally. The crista bicipitalis is relatively small and round. The 179 fossa tricipitalis dorsalis is smooth. With regard to the radius, the cotyla humeralis is completely round. 180 The tuberculum bicipitale radii is elongated and conspicuous. The shaft is straight and has a round section 181 towards its cranial end, whereas it becomes flattened towards the distal end. The facies articularis 182 radiocarpalis is smooth and curved. The tuberculum aponeurosis ventralis protrudes laterally. The 183 humerus shows a deeper and narrower impressio coracobrachialis than in Gallus and Phasianus. In 184 proximal view, the caput humeri is cranio-caundally wider and laterally shorter than in both mentioned 185 species. Miophasianus medius does not seem to show distinctive morphological features with regard to 186 M. altus. Both taxa are only distinguishable on their respective size-ranges. 187 Measurements. See Table 1. 188 Remarks. The genus Palaeoperdix was originally described from Sansan by Milne-Edwards 189 (1869). This genus comprised P. longipes (type species by original designation), P. prisca and P. 190 sansaniensis. Ballmann (1969) wrote down that Palaeoperdix longipes was synonymous with Palaeortyx 191 phasianoides Milne-Edwards, 1869, which was also supported by Göhlich and Mourer-Chauviré (2005). 192 As to the two other species originally placed within Palaeoperdix, Cheneval (2000) synonymized 193 Palaeoperdix sansaniensis with Palaeoperdix prisca and transferred it to Palaeortyx Milne-Edwards, 7 194 1869. On the other hand, Cheneval (2000) transferred Miophasianus medius to Palaeoperdix. As a 195 consequence, for this author two species remain in the genus Palaeoperdix: P. longipes and P. medius. 196 However, I consider that Miophasianus medius must be placed within the same genus as Miophasianus 197 altus. I agree with the original description in that the distinction between medius and altus is the smaller 198 size of the former species (Milne-Edwards 1869-71). M. medius has been recorded in Olival da Susana 199 (MN-5, Portugal; Mourer-Chauviré and Antunes 2003), Przeworno II (MN6-7+8, Poland; Bocheński 200 1987) and La Grive-Saint-Alban (MN7+8; Déperet 1887; Ennouchi 1930; Gaillard 1939; Ballmann 201 1969). 202 203 [PLACE TABLE 1 ABOUT HERE] 204 205 Palaeortyx joleaudi Ennouchi, 1930 206 (Figure 1i) 207 208 Localities. La Artesilla (MN4). 209 Referred specimens. MNCN 74035, proximal end of right humerus. 210 Description. The specimen is a bone fragment. It has a very deep and large fossa 211 pneumotricipitalis dorsalis. The fossa pneumotricipitalis ventralis is well developed. It shows the 212 characteristic indentation of Palaeortyx between de caput humeri and the tuberculum ventrale (Göhlich 213 and Mourer-Chauviré 2005). 214 Measurements. See Table S1. 215 Remarks. The genus Palaeortyx was originally described by Milne-Edwards (1869-71) based on 216 fossil material from Saint-Gérand-le-Puy (MN1-MN2, France). This author erected three new species for 217 this genus: Palaeortyx gallica, Palaeortyx brevipes and Palaeortyxa phasianoides. Later, the smallest- 218 sized species within the genus known so far, Palaeortyx joleaudi, was described from La Grive-Saint- 219 Alban (MN7+8) by Ennouchi (1930). Mlíkovský (2002) synonymized Palaeortyx with the extant genus 220 of quails Coturnix Bonnaterre, 1791, an opinion that has been revised and discussed by Göhlich and 221 Mourer-Chauviré (2005). Regarding the few features that can be observed in the small fragment of bone 222 from La Artesilla, the fossa pneumotricipitalis dorsalis is deeper and relatively larger in Palaeortyx than 223 in Coturnix, whereas in Coturnix the fossa pneumotricipitalis ventralis is relatively wider than in 8 224 Palaeortyx. In Megalocoturnix cordoni, from the Ruscinian outcrop of Layna (Spain), the fossa 225 pneumotricipitalis dorsalis is likewise fairly smoother than in Palaeortyx (Sánchez Marco 2009). This 226 find in Artesilla constitutes the second record for this species, the first being that from La-Grive-Saint- 227 Alban, on which basis this taxon was originally described. 228 229 Palaeortyx brevipes Milne-Edwards, 1869 / Palaeortyx grivensis Lydekker, 1893 230 (Figure 1j–p) 231 232 Coturnix (?) miocenica Villalta, 1963:271, pl. IV: fig. 1, 1a, 1b (original description). 233 p. Coturnix gallica Milne-Edwards, 1869: Mlíkovský 2002:154. 234 235 Localities. Toril 3 (MN7+8), Escobosa (MN 7+8), Can Poncic (MN9), Batallones 1 (MN10), 236 Valdecebro 5 (MN12), Los Mansuetos (MN12). 237 Referred specimens. Toril 3: MNCN 74030, complete left coracoideum (Figure 1m); MNCN 238 74031, cranial end of right coracoideum; MNCN 74029, distal end of left tarsometatarsus (Figure 1k). 239 Escobosa: MNCN 74032, cranial end of left coracoideum; MNCN 74033 (Figure 1l) and MNCN 74034, 240 two proximal ends of left humerus (one of them, MNCN 74034, very damaged and not useful for taking 241 measurements). Can Poncic: IPS87218, proximal end of left carpometacarpus (Figure 1j). Batallones 1: 242 without number, cranial end of right scapula (Figure 1o), cranial end of right coracoideum, proximal end 243 of right humerus (Figure 1n), distal end of left humerus, distal end of left ulna, and proximal end of left 244 carpometacarpus. Valdecebro 5: MNCN 74036, distal end of right tarsometatarsus. Los Mansuetos: 245 IPS12678 (Figure 1p), proximal end of right carpometacarpus (Figure 1p). 246 Description. In the scapula, the acromion is very large and crescent-shaped. The tuberculum 247 coracoideum is patent. The facies articularis humeralis has a triangular shape. In the coracoideum of some 248 phasianids the processus procoracoideus is very small, as in Alectoris Kaup, 1829, Perdix Brisson, 1760, 249 or Lagopus Brisson, 1760. A characteristic feature of Palaeortyx is the extreme reduction of this 250 processus (Cheneval and Adrover, 1993). Moreover, it forms a continuous surface with the facies 251 articularis scapularis. In Palaeocryptonyx Depéret, 1892, there is a step between both anatomical 252 elements. The facies articularis humeralis is large and fused with the cotyla scapularis. The extremitas 253 omalis is not medially inclined. There is no foramen pneumaticum in the area of the impressio musculi 9 254 sternocoracoidei. The angulus medialis protrudes distally. The facies articularis sternalis is very short and 255 more curved than in Palaeocryptonyx . With regard to the humerus, in Palaeortyx it is characterized by 256 two deep (dorsal and ventral) fossae pneumotricipitalis, the former being larger. The impressio 257 coracobrachialis is very deep and wide. The crista deltopectoralis is relatively more developed than in 258 other phasianids. The fossa musculi brachialis is well marked. The region of the processus flexorius is 259 very developed. In the ulna, the shaft is compressed laterally. The tuberculum carpale is small. The 260 outline of the condylus dorsalis ulnae is round. Regarding the carpometacarpus, the available bone 261 fragments show a typically galliform-shape. The dorsal edge of the trochlea carpalis is characteristically 262 round. The processus extensorius is curved upwards and little developed. The processus alularis is 263 scarcely developed, like the processus intermetacarpalis. The processus intermetacarpalis is not very 264 developed (contra Cheneval and Adrover 1993). The three trochleae of the tarsometatarsus conform a 265 semicircle. The trochlea for digit II protrudes distally less than the trochlea of digit IV and is likewise less 266 developed. 267 Measurements. See Tables S1 and S2. 268 Remarks. The specimen from Los Mansuetos is the holotype and the only attributed specimen to 269 ?Coturnix miocenica Villalta, 1963. This taxon was synonymized with Coturnix gallica Milne-Edwards, 270 1869, by Mlíkovský (2002), who in the same work also transferred some species of Palaeortyx to the 271 extant genus Coturnix. As explained above, Palaeortyx brevipes was erected by Milne-Edwards (1869- 272 71) based on some bones from Saint-Gérand-le-Puy (MN1-MN2, France). The author noted that this 273 taxon was smaller than Palaeortyx gallica, another species also described by himself from the same 274 locality. Paleortyx brevipes has been noticed from the Eocene and Oligocene of Quercy (France) by 275 Gaillard (1908: Palaeortyx cayluxensis) and Mourer-Chauviré (1992) up to Aljezar B (Spain) (Cheneval 276 and Adrover 1993: Palaeortyx cf. brevipes). The small-sized Palaeortyx (P. gallica, P. brevipes, P. 277 grivensis, P. joleaudi) were synonymized by Mlíkovský (2002) and transferred to Coturnyx gallica. 278 Palaeortyx grivensis, was described by Lyddeker (1893) on material from La Grive (MN7+8). From the 279 same locality, Ennouchi (1930) described Palaeortyx depereti as being of intermediate in size between 280 Palaeortyx grivensis and Palaeortyx edwardsi Depéret, 1887. However, Ballmann (1969) wondered 281 whether the differences between Palaeortyx grivensis and Palaeortyx depereti were enough to justify the 282 recognition of two distinct taxa. Subsequent reviewers (Mlíkovský 2002; Göhlich and Mourer-Chauviré 283 2005; Göhlich and Pavia 2011) have supported such an opinion. The remains from Gargano (Italy) 10 284 previously assigned to Palaeortyx grivensis by Ballmann (1973, 1976) have been subsequently 285 considered to correspond to a new taxon, Palaeortyx volans Göhlich and Pavia, 2011. Ballmann (1973) 286 synonymyzed Palaeocryptonyx depereti (Ennouchi, 1930) with Palaeortyx brevipes, of the same size 287 range. The finds from the fossil localities studied in this paper have to be attributed to the genus 288 Palaeortyx, within the size ranges of Palaeortyx grivensis and Palaeortyx brevipes. Zelenkov (2017) has 289 transferred one coracoid from Rudabànya (MN9, Hungary) previously assigned to Palaeortyx grivensis 290 by Jánossy (1993) and to Palaeortyx brevipes by Kessler (2009) to Rollulinae indet. 291 292 Palaeortyx phasianoides Milne-Edwards, 1869 293 (Figure 1q,r) 294 295 Localities. Can Mas (MN4), Toril 3 (MN7+8). 296 Referred specimens. Can Mas: IPS12647 (Figure 1q), distal end of left humerus. Toril 3: MNCN 297 74028 (Figure 1r), proximal end of right carpometacarpus. 298 Descriptions. In the humerus, the sulcus humerotricipitalis is relatively deep. The condylus 299 ventralis protrudes distally. The condylus dorsalis is long in cranial view. The processus flexorius shows 300 a pointed outline in ventral view. The cross section of the diaphysis near the end of the bone is ovate and 301 flattened antero-posteriorly. In the carpometacarpus, the processus intermetacarpalis is well developed. 302 The trochlea carpalis ends distally (towards the processus extensorius) abruptly. The ventral outline of the 303 trochlea is very rounded. The processus extensorius is stout. The processus alularis is prominent. 304 Measurements. See Table S1. 305 Remarks. The specimen from Can Mas was attributed by Villalta (1963) to Palaeortyx edwardsi, 306 which was originally described by Depéret (1887) based on some bones from La Grive-Saint-Alban. 307 Ballmann (1969) pointed out the wrong attribution to the genus, and transferred it to Palaeocryptonyx 308 Depéret, 1892, resulting in the new combination Palaeocryptonyx edwardsi (Depéret, 1887). In a later 309 review, Mlíkovský (2002) synonymized the genus Palaeocryptonyx with Alectoris. Cheneval (2000) 310 referred the humerus from Can Mas to Palaeortyx prisca. Göhlich and Mourer-Chauviré (2005) founded 311 the distinction between Palaeortyx phasianoides and Palaeortyx prisca in their respective sizes, reaching 312 Palaeortyx phasianoides the highest size range. The condylus ventralis is more prominent distally than in 313 Alectoris and Gallus. In distal view, the processus flexorius juts from the bone more than in Numida 11 314 Linnaeus, 1766, Alectoris, Lagopus, Gallus, Phasianus and Perdix. This processus is little developed in 315 Numida and Coturnix. The fossa musculi brachialis is wider in the humerus of Palaeortyx phasianoides 316 than in Palaeocryptonyx edwardsi, giving to this part of the bone a more flattened appearance in 317 Palaeortyx phasianoides. The proximal end of the carpometacarpus is the same size in Palaeocryptonyx 318 edwardsi and Palaeortyx phasianoides. Here, the processus extensorius is curved and directed 319 proximally, as in Alectoris and Perdix. In Gallus, Phasianus, Lagopus, Coturnix and Palaeocryptonyx 320 this process is directed more laterally. This process is greatly reduced in Numida. In Palaeortyx 321 phasianoides, the dorsal edge of the trochlea carpalis does not stand out from the ventral edge as in 322 Palaeocryptonyx edwardsi. 323 324 Palaeortyx gallica Milne-Edwards, 1869 325 (Figure 1s–u) 326 327 Phalacrocorax ibericum Villalta, 1963:267, pl. II: fig. 4, 4a (original description). 328 Palaeoperdix sansaniensis Milne-Edwards, 1869: Villalta 1963:270, pl. IV: fig. 3, 3a, 3b. 329 p. Palaeortyx prisca (Milne-Edwards, 1869): Cheneval 2000:345 (new combination). 330 Phalacrocorax ibericus Villalta, 1963: Mlíkovský 2002:72 (ending corrected). 331 332 Localities. Toril 3 (MN7+8), Hostalets de Pierola indeterminate (MN7+8-MN9), Los Valles de 333 Fuentidueña (MN9). 334 Referred specimens. Hostalets de Pierola indeterminate: IPS73336 (Figure 1s), distal end of right 335 tibiotarsus. Toril 3: MNCN 74027 (Figure 1t), partial left tibiotarsus (missing its distal end). Los Valles 336 de Fuentidueña: IPS12650 (Figure 1u), distal end of right humerus. 337 Description. The humerus is badly damaged: both condylus dorsalis and processus flexorius are 338 partially lost. The fossa musculi brachialis is deep and well marked. The epicondylus dorsalis as well as 339 the tuberculum supracondylaris ventrale are scarcely developed. This distal end of humerus is quite 340 different from that of Phalacrocorax Brisson, 1760. A very small processus flexorius is a characteristic 341 feature of the latter genus. The specimen from Los Valles de Fuentidueña has the overall shape of the 342 Phasianidae. The two incomplete tibiotarsi have a characteristic overall galliform shape. The bone from 343 Hostalets de Pierola has a wide pons supratendineus, and both condyles are round and similar in size. The 12 344 facies articularis lateralis of the bone from Toril 3 is relatively small. In proximal view, it has an 345 elongated shape. The crista cnemialis is short. 346 Measurements. See Table S1. 347 Remarks. The distal tibiotarsus from Hostalets was identified by Villalta (1963) as Palaeoperdix 348 sansaniensis Milne-Edwards, 1869. This species was described based on a distal end of tibiotarsus from 349 Sansan, and considered by Milne-Edwards (1869) to be smaller than Palaeoperdix prisca Milne-Edwards, 350 1869, likewise described based on material from the same locality. However, owing to the fragmentary 351 state of the holotype, Milne-Edwards (1869) had some doubts on the allocation of this species to genus 352 Palaeoperdix, being originally spelled as “Palaeoperdix (?) sansaniensis”. As mentioned above, 353 Cheneval (2000) synonymized Palaeoperdix sansaniensis with Palaeoperdix prisca, and subsequently 354 transferred the latter species to genus Palaeortyx. The bones of Palaeortyx prisca reach larger sizes than 355 Palaeortyx gallica, but there is some overlap in the measurements. However, the bones from Hostalets de 356 Pierola and Toril 3 fit the tibiotarsus size ranges of Palaeortyx gallica, and are therefore attributed to this 357 species. In turn, the specimen from Los Valles de Fuentidueña was designated as the holotype of 358 Phalacrocorax ibericum Villalta, 1963. No other specimen has been subsequently assigned to this taxon. 359 Mlíkovský (2002) corrected the ending of the specific epithet (i.e., Phalacrocorax ibericus) and indicated 360 that the fragment was too eroded to allow for an exact identification. 361 362 Order Strigiformes (Wagler, 1830) 363 Family Tytonidae Ridgway, 1914 364 365 Tyto balearica Mourer-Chauviré et al., 1980 366 (Figure 1v,w) 367 368 Tyto campiterrae Jánossy, 1991:25-26, fig. 3, tab. 2 (original description) 369 p. Tyto sanctialbani Lydekker, 1893: Mlíkovský 1998:253. 370 p. Tyto sanctialbani Lydekker, 1893: Mlíkovský 2002:209. 371 Tyto campiterrae Jánossy, 1991: Pavia and Mourer-Chauviré 2001:1095, fig. 4. 372 373 Localities. Hostalets de Pierola indeterminate (MN7+8-MN9), Los Mansuetos (MN12). 13 374 Referred specimens. Hostalets de Pierola indeterminate: IPS12652 (Figure 1v), distal end of 375 right humerus. Los Mansuetos: IPS12673 (Figure 1w), pedal phalanx 3 of digit III. 376 Description. Regarding the phalanx from Los Mansuetos, its proximal articular facet has a 377 pyriform-rounded shape. The apophysis flexoris is large and fairly exceeds the apohysis extensoris, which 378 is relatively small and pointed. Its plantar surface is slightly concave. With regard to the humerus, the 379 distal humeral morphology of strigiforms is fairly distinguishable from that of accipitriforms. Among 380 some of the most conspicuous features, the condylus dorsalis is more developed and distally protruding in 381 the diurnal raptors. In Strigidae the epicondylus dorsalis is larger than in Tyto and is more protruding 382 dorsally. The zone of the processus flexorius and the epicondylus ventralis is more robust in Strigidae. 383 The fossa musculi brachialis is more marked in Tyto than in Strigidae. The humeral fragment from 384 Hostalets is indistinguishable from Tyto balearica. 385 Measurements. For the humerus, see Table 2. There are few data on phalangeal measurements. 386 The maximum length of the phalanx from Los Mansuetos is 16.4 mm, which is between the values for the 387 extant Tyto alba (11.4–12.3 mm, N=17; Langer, 1980) and Tyto balearica from the Middle Pleistocene of 388 Corsica and Sardinia (18.1 and 18.3 mm; Louchart 2002), but is very close to the measurements of Tyto 389 balearica from the Pliocene of Sète (16.7 mm; Mourer-Chauviré and Sánchez Marco 1988). 390 391 [TABLE 2 ABOUT HERE] 392 393 Remarks. Villalta (1963) attributed with doubts the phalanx from Los Mansuetos to the 394 strigiforms. Indeed, this phalanx belongs to Tyto. The articular facet is distinctively more elongated than 395 in Strigidae. In this family, the processus flexorius does not exceed the extensorius process, or exceeds it 396 slightly. However, in Tyto that happens in phalanges 3 III and 4 IV, as it can be seen in the fossil from 397 Los Mansuetos. In diurnal raptors, this phalanx shows a subtriangular articular facet. There are no 398 published data for this phalanx in Tyto sanctialbani, and hence it may not be excluded that the phalanx 399 from Los Mansuetos might be attributable to such a taxon. 400 The humerus from Hostalets de Pierola, in turn, was first identified as Aquila cf. minuta Milne- 401 Edwards, 1871 by Villalta (1963), and later assigned to Hieraaetus edwardsi (Sharpe, 1899) by 402 Mlíkovský (2002). The distal width of this humerus is 16.9 mm, which exceeds the variation ranges of 403 the extant barn owls Tyto alba (see Langer 1980) and Tyto sanctialbani (see Sánchez, 2001; Mlíkovský 14 404 1998; Pavia and Mourer-Chauviré 2011) , as well as the recently described Miotyto montispetrosi 405 (Göhlich and Ballmann 2013) from Steinberg and Goldberg (MN6). Tyto robusta and Tyto gigantea from 406 Gargano (Ballmann 1973, 1976) reach larger sizes. There are no published measurements of the distal 407 end of this bone for T. gigantea, but its total length corresponds to a bone fairly larger than the humerus 408 from Hostalets. Tyto robusta and T. gigantea were synonymized by Mlíkovský (1998), who considered T. 409 gigantea as a larger-sized chronospecies of the earlier T. robusta, and consequently synonymized Tyto 410 robusta Ballmann, 1973 with Tyto gigantea Ballmann, 1973. The size of the specimen from Hostalets fits 411 into the corresponding range of values of Tyto balearica (Table 2). 412 Bones attributed to Tyto sanctialbani have been reported from Toril 3 (MN7+8) and La Grive- 413 Saint-Alban (MN7+8) up to the upper Miocene (MN10) of Kohfidisch (Austria) (Mlíkovský 1998; 414 Sánchez Marco 2001; Pavia and Mourer-Chauviré 2011). Mlíkovský (1998) synonymized Tyto 415 campiterrae Jánossy, 1991 with T. sanctialbani. The former was described by Jánossy (1991) on the basis 416 of bones from the Hungarian locality of Polgárdi (MN13), with the larger size T. campiterrae as the sole 417 declared difference from T. sanctialbani. In a recent work on the remains of T. sanctialbani from La 418 Grive-Saint-Alban, Pavia and Mourer-Chauviré (2011) supported the taxonomic validity of T. 419 campiterrae as a valid species on the basis of a photograph, and against the view by Mlíkovský (1998) 420 that these two nominal taxa are synonymous. However, Pavia and Mourer-Chauviré (2011) did not 421 provide any measurements to support their view. The measurements of T. campiterrae provided by 422 Jánossy (1991) fit into the size ranges of T. balearica (see Table 2). Pavia and Mourer-Chauviré (2011) 423 further concluded that T. balearica is not recorded in the Miocene, as previously pointed out by Louchart 424 (2002), but did not based their conclusion on the consideration of size and morphology. In fact, these 425 authors did not deal throughout their paper with the Miocene fossil remains previously assigned to Tyto 426 balearica, with the only exception of those attributed to Tyto campiterrae. Louchart (2002), in contrast, 427 did discuss the bones allocated to T. balearica from Mio-Pliocene localities, and therefore deserves 428 further discussion. On the remains from Aljezar B (MN12) assigned to T. balearica by Cheneval and 429 Adrover (1993), Louchart (2002: 72) asserted (my translation from the French original): they “prove to be 430 different from T. balearica (by the lower dimensions of certain bones, especially the posterior phalanges). 431 This will not be detailed here, but separately (Louchart, in preparation). These data are therefore 432 considered not related to T. balearica.” Louchart (2002) said nothing about the descriptions of the bones 433 or the fact that the measurements given by Cheneval and Adrover (1993: table 4) fit the size ranges of T. 15 434 balearica from the localities where such a species was originally described. In that study, the 435 reconsideration of the measurements of the fossils of T. balearica from other Miocene and Pliocene 436 French and Spanish localities (Sète, Balaruc II, Layna, Valdecebro 5, Moreda; see Mourer-Chauviré and 437 Sánchez Marco 1988; Sánchez Marco 2001) follows the same reasoning: some values fall above or below 438 the original sample from the Balearic original localities plus the Pleistocene Corsican remains reported in 439 that same work, apparently meaning that the samples from Balearic, Corsica and Sardinia islands 440 encompass the whole size ranges of Tyto balearica. On this basis, it is difficult to understand why 441 Louchart (2002) agreed the remains from the Spanish site of Almenara-Casablanca 1 (a distal end of 442 tarsometatarsus and a pedal phalanx) were attributed to T. balearica, but not the bones from Sète (see 443 Table 2), for instance. It is noteworthy that the osteometric data from the original localities are very 444 scarce (the best represented measure being available only from three specimens), whereas data from 445 Corsica and Sardinia are based on larger sample sizes (although even there only a few measures reach 446 high numbers enough for statistical purposes). It is even more remarkable that some bones of the owl 447 from Corsica and Sardinia reach values considerably different from those of the Balearic islands as well 448 as from mainland localities. The Tyto described in Corsica and Sardinia is in general small-sized than the 449 Balearic taxon. Only the femur, some measures of the tarsometatarsus and one of the humerus fit with 450 Tyto balearica, which could be indicative of a singular process of insular differentiation, as in the cases of 451 Bubo insularis Mourer-Chauviré and Weesie, 1986, and Athene angelis Mourer-Chauviré et al., 1997, on 452 the same islands. After re-examining again the bones attributed to T. balearica from Iberian localities, 453 and reviewing the corresponding descriptions and measurements (see Table 2) provided by Mourer- 454 Chauviré and Sánchez Marco (1988), Cheneval and Adrover (1993) and Sánchez Marco (2001), it is 455 concluded here that these bones are indistinguishable from the holotype and the associated material of T. 456 balearica (Mourer-Chauviré et al. 1980). On the other hand, the diagnostic characters of the genus 457 Miotyto Göhlich and Ballmann, 2013, are not present in the fossil material of T. balearica. Thus, in the 458 tarsometatarsus of the latter species, there is no vestigial arcus extensorius (see dorsal views in Figure 459 4.2) and the tuberculum musculi fibularis brevis is not directed plantolaterally, unlike in Miotyto. Tyto 460 balearica (see proximal views in Figure 4.2) shares both features with T. alba, which shows that the 461 Neogene remains attributed to T. balearica cannot be alternatively assigned to a species of Miotyto. 462 463 [PLACE FIGURE 2 ABOUT HERE] 16 464 465 Order Accipitriformes Vieillot, 1816 466 Family Accipitridae Vieillot, 1816 467 (Figure 1x, y) 468 469 Localities. Hostalets de Pierola indeterminate (MN7+8-MN9), Los Valles de Fuentidueña 470 (MN9), Viladecavalls (MN10). 471 Referred specimens. Hostalets de Pierola indeterminate: IPS12653, pedal (ungual) phalanx. 472 Viladecavalls: IPS12674 (Figure 1x), pedal phalanx 1 of digit III; IPS12675, proximal end of pedal 473 phalanx 1 of digit I. Los Valles de Fuentidueña: IPS12676 (Figure 1y), pedal (ungual) phalanx. 474 Description. The phalanx 1 digitis I from Viladecavalls lacks its distal end. Its articular cotyla is 475 laterally elongated. The proximal end is compressed cranio-plantarly. A groove runs along the plantar 476 side of the phalanx. The articular cotyla of the phalanx 1 digitis III from Viladecavalls adopts a rounded 477 shape. Its plantar groove is deep at the proximal area of the bone. Fovea subtrochlearis absent, and foveae 478 ligamenti collateralis marked. 479 Measurements. Viladecavalls—phalanx 1 digitis I: maximum length, 15.9 mm; maximum width 480 of the proximal end, 5.7 mm. Viladecavalls—phalanx 1 digitis III: maximum width of the proximal end, 481 8.5 mm. Hostalets de Pierola indeterminate—pedal (ungual) phalanx: maximum length, ca. 26.2 mm; 482 width of the proximal end, 4.8 mm. Los Valles de Fuentidueña—pedal (ungual) phalanx: maximum 483 length, 21.0 mm; width of the proximal end, 5.0 mm. 484 Remarks. The two phalanges from Viladecavalls were attributed to Aquila sp. by Villalta (1963), 485 and this assignment was retained by Mlíkovský (2002). Although lacking the distal end of the phalanx 1 486 digitis I, the bone did not stretch much more because the preserved remain retains the curve at the end of 487 the bone. By its size and general shape, both phalanges from Viladecavalls are reminiscent of Buteo 488 Lacépède, 1799, Accipiter Brisson 1760, and Hieraaetus pennatus Gmelin, 1788. A more detailed study 489 of these phalanges is the subject of an ongoing separate work. 490 The two ungual phalanges from Hostalets de Pierola and Los Valles de Fuentidueña were 491 attributed to Aquila cf. minuta Milne-Edwards, 1871 by Villalta (1963), although Lambrecht (1933) had 492 previously established that this taxon was a junior synonym of Aquila edwardsi (Sharpe, 1899). Later, 493 Mlíkovský (2002) proposed the new combination Hieraaetus edwardsi, in which the material from 17 494 Hostalets was included. The item from Los Valles de Fuentidueña was not mentioned in that work. 495 However, due to the lack of diagnostic features, little can be said on the taxa to which these ungual 496 phalanges belong, other than concluding that they belong to an accipitriform of intermediate size. 497 498 Order Charadriiformes (Huxley, 1867) 499 Family Recurvirostridae Bonaparte, 1831 500 501 Recurvirostra teruelensis (Villalta, 1963) new combination 502 (Figure 1z) 503 504 Totanus teruelensis Villalta, 1963:268, pl. II: fig. 5, 5a, 5b, pl. III: fig. 1, 1a, 1b (original description). 505 Tringa teruelensis (Villalta, 1963): Bocheński 1997:316 (new combination). 506 Larus teruelensis (Villalta, 1963): Mlíkovský 2002:137 (new combination). 507 508 Localities. Los Mansuetos (MN12). 509 Referred specimens. IPS12677, distal end of right humerus. The distal end of ulna reported by 510 Villalta (1963) has not been found. 511 Description. This is the emended description of the fragment of humerus (holotype). The 512 processus supracondylaris dorsalis is partially broken, but a wing-like surface in the dorsal edge of the 513 distal end of the humerus (between the dorsal condyle and the dorsal epicondylar process), which is 514 characteristic of several charadriiforms, is discernible. Its contour is sinuous. The sulcus 515 humerotricipitalis is deep. The processus flexorius is relatively small and the epicondylus ventralis is 516 round and very pronounced. The condylus dorsalis protrudes dorsally. 517 Measurements. Smallest width of the diaphysis, 3.4 mm; distal width, 7.6 mm; distal depth, 4.8 518 mm. 519 Remarks. Villalta (1963) erected the species Totanus teruelensis based on a distal end of 520 humerus and a distal end of ulna. Both items were described and figured (Villalta 1963), but the ulnar 521 fragment has not been found in the collection by the author of the present work. None of them was 522 designated as the holotype. Although, whereas the humeral fragment was described first, this one should 523 be considered the holotype. Olson (1985) indicated that this taxon had been based on the humeral 18 524 fragment, and noted down that the deep fossa musculi brachialis was more like to that seen in the 525 humerus of gulls. Bocheński (1997) synonymized Totanus Bechstein, 1803 with Tringa Linnaeus, 1758. 526 Finally, Mlíkovský (2002) considered Olson’s (1985) opinion on the fossa musculi brachialis to be 527 correct, and transferred the taxon to Larus Linnaeus, 1758. However, the fossa musculi brachialis is 528 considerable deeper in Larus, Sterna Linnaeus, 1758 and Gelochelidon CL Brehm, 1830 than in the fossil 529 from Los Mansuetos. The condylus dorsalis is prominent in the studied specimen, as in Recurvirostra and 530 in Gelochelidon, but not as prominent in Tringa, Limosa Brisson, 1760, Himantopus Brisson, 1760, 531 Calidris Pallas, 1764, Arenaria Brisson, 1760, Actitis Illiger, 1811, and Lymnocryptes F. Boie, 1826. A 532 sinuous contour of the dorsal edge of the bony wing is seen in Recurvirostra, whereas in Tringa 533 Linnaeus, 1758, Limosa and Himantopus such an edge is smooth. The condylus ventralis protrudes more 534 distally than the condylus dorsalis in the fossil. This feature is only seen in Recurvirostra, and 535 Haematopus Bonaparte, 1838, and slightly in Limosa. The processus supracondylaris dorsalis is more 536 developed in Gelochelidon, Arenaria, Tringa, Sterna and Limosa. The fossa musculi brachialis is less 537 deep in: Haematopus, Himantopus and Limosa.The fossil specimen from Los Mansuetos is considerably 538 smaller than the corresponding element in the extant Recurvirostra avosetta Linnaeus, 1758. 539 540 Charadriiformes indet. 541 (Figure 1a’) 542 543 Localities. El Fallol (MN3). 544 Referred specimens. IPS56, distal end of left tarsometatarsus. 545 Description. This item is a very small bone fragment. The trochleae of digits II and IV are 546 eroded and their corresponding plantar ends are absent. Both of them reach the same level distally. The 547 trochlea of digit III is fairly larger and distally more protruding than the other two. In Cathartidae, the 548 trochlea of digit IV reaches distally as far as the trochlea of digit III, or exceeds this one, which is not 549 seen in the fossil from El Fallol. In distal view, the trochleae do not form a marked curvature. Its overall 550 appearance points to the charadriiforms, but this small piece of bone lacks diagnostic traits to pinpoint its 551 identification. Sterna paradisaea Pontoppidan, 1763 is the recent species to which the fossil bears a 552 greater resemblance, although the former is smaller. 553 Measurements. Distal width, 6.6 mm; distal depth, ca. 4.2 mm. 19 554 Remarks. Based on this tiny distal end of tarsometatarsus, Villalta Comella and Crusafont Pairó 555 (1955) erected the species ?Plesiocathartes gaillardi. This genus allocation was owed to the 556 morphological similarities with the Eocene/Oligocene species Plesiocathartes europaeus Gaillard, 1908 557 from Quercy. However, according to Gaillard (1908), Plesiocathartes europaeus was in its turn even 558 smaller than any other known Cathartidae. In the original description, Plesiocathartes gaillardi was 559 considered to be even smaller than the type species of the genus (Crusafont and Villalta 1955). The 560 taxonomic validity of the former taxon was maintained by Villalta (1963). However, Jollie (1977) 561 dismissed any link between Plesiocathartes europaeus and the Cathartidae, and Mlíkovský (2002) placed 562 both Plesiocathartes europaeus and Plesiocathartes gaillardi as Aves incertae sedis. Plesiocathartes 563 gaillardi should be taken as nomen dubium. 564 565 Order Passeriformes (Linnaeus, 1758) 566 Parvorder Passerida Sibley et al., 1988 567 568 Localities. Los Mansuetos (MN12). 569 Referred specimens. IPS73335, distal end of left humerus. 570 Description. This bone shows a typical passeriform morphology. The processus flexorius is well 571 developed and protrudes disto-ventrally. Brachial depression absent. The impression of the anterior 572 articular ligament is deep and large. The fossa olecrani is deep. 573 Measurements. Smallest width of the diaphysis, 1.7 mm; distal width, 4.4 mm; distal depth, 2.2 574 mm. 575 Remarks. Villalta (1963) attributed this humeral fragment to Fringilla sp. However, several 576 families of Passeriformes resemble the fossil in both size and shape, so that a more precise taxonomic 577 attribution is not warranted. The bone size prevents it to be attributed to Corvidae. 578 579 4 Discussion and conclusions 580 581 A summary of the taxa discussed in this study provides is given in Table 3. Most of prior 582 identifications gathered from the literature are wrong. The item from Can Mas assigned to Palaeortyx 583 edwardsi is now attributed to Palaeortyx phasianoides. Three taxa have been found to be taxonomically 20 584 invalid: Phalacrocorax ibericum Villalta, 1963 is a junior subjective synonym of Palaeortyx gallica; 585 Coturnix miocenica Villalta, 1963 is a junior subjective synonym of Palaeortyx brevipes/P. grivensis , 586 and ?Plesiocathartes gaillardi Crusafont et al., 1955 is a nomen dubium, being attributed to an 587 indeterminate charadriiform. The species Totanus teruelensis Villalta, 1963, later transferred to Larus 588 teruelensis by Mlíkovský (2002), is now transferred to Recurvirostra teruelensis (Villalta, 1963) comb. 589 nov. The fragment of humerus from Los Mansuetos assigned to Fringilla sp. shares features with several 590 families of Passerida and should be thus considered an indeterminate representative of this taxon. There is 591 no evidence of Aquila minuta Milne-Edwards, 1871—transferred to Aquila edwardsi by Lambrecht 592 (1933) and to Hieraetus edwardsi by Mlíkovský (2002)— in the Iberian record. The distal end of 593 humerus from Hostalets de Pierola formerly attributed to Aquila cf. minuta is now attributed to Tyto 594 balearica, whereas the two ungual phalanges from Hostalets and Los Valles de Fuentidueña are assigned 595 to Accipitridae indet. Two pedal phalanges from Viladecavalls, previously assigned by Villalta (1963) to 596 Aquila sp., are now attributed to Accipitridae indet. 597 A large phasianid, Miophasianus altus, is recorded from the Early to the Late Miocene of the 598 Iberian Peninsula (Sánchez Marco 2006; and the present study). Another large-sized taxon of this group, 599 Miophasianus medius, was already kown from the Middle to Late Miocene of Hostalets de Pierola 600 (Sánchez Marco 2006), and now it is also reported from the Late Miocene of Batallones. Also, a number 601 of skeletal remains belonging to smaller-sized forms have been found along the Miocene. The smallest is 602 Palaeortyx joleaudi, from the MN 4 of La Artesilla. Other small-sized skeletal elements from the Middle 603 to the Late Miocene could be assigned to either Palaeortyx brevipes or Palaeortyx grivensis. Some bones 604 slightly larger, from the Middle and Late Miocene, belong to Palaeortyx gallica. Finally, Palaeortyx 605 phasianoides has been found from the Early and Middle Miocene. 606 607 [TABLE 3 ABOUT HERE] 608 609 Villalta and Crusafont (1950) attributed a distal end of humerus from Hostalets de Pierola to 610 Palaeortyx miocaena. That work yielded a poor-quality drawing of it. The same taxonomic assignment 611 was kept by Villalta (1963), in which all the studied bones were figured with photographs, with the 612 exception of this humeral fragment. That is why it is likely that the bone was already lost in 1963. 21 613 In the Neogene of Europe, there are owl skeletal remains referable to the family Tytonidae. 614 Apart from Miotyto montispetrosi, these remains are morphologically indistinguishable from the extant 615 species Tyto alba, or discernible only by minor details. Tyto sanctialbani has a similar size to Tyto alba 616 and Miotyto montispetrosi, and has been recorded from the late Aragonian to the Vallesian. Larger bones 617 than those of the extant barn owl, and of similar size than those of Tyto balearica from its original 618 localities, have been reported likewise from the Middle Miocene to the Middle Pleistocene. The bones 619 attributed to Tyto campiterrae can not be distinguished from those of Tyto balearica, the former thus 620 being a junior subjective synonym of the latter. In contrast, Tyto robusta and Tyto gigantea, from the 621 Mio-Pliocene of Gargano island, are remarkably larger than their congeneric species. 622 Louchart (2002) found abundant material of a species of Tyto in the Middle Pleistocene of 623 Sardinia and Corsica. These bones are larger than those of the extant Tyto alba, and some of its 624 measurements are similar to those of Tyto balearica, but most of them correspond to a smaller-sized form 625 of barn owl. Rather than describing a new species, Louchart (2002) attributed this material to a new 626 subspecies, Tyto balearica cyrneichnusae. The subspecies notion is useful when it refers to populations 627 bearing few different features, but capable of interbreeding with other populations with the result of 628 fertile offspring. A fossil or paleontological species designates a collection of morphological features 629 observed on fossil items. Predictably, many species described on fossils correspond to extinct biological 630 species, but such an assumption is not free of uncertainty in each particular case. A subspecies within a 631 paleontological species raises a controversial issue because the theoretical criterion of required 632 interbreeding among individuals of different subspecies cannot be tested. Two sets of morphological 633 characters that differ in a few of them have to be designated with different specific names because it 634 cannot be assumed that they formed in the past a single breeding community. The fossil material in itself 635 must be the only base for describing new taxa. Geographical and temporal distributions of the findings 636 should be viewed as extrinsic features, not valid for the decision of erecting a new taxa. 637 Methodologically, it is incorrect to take the size ranges of the bones of Corsica and Sardinia as those of 638 Tyto balearica, and then discard the entire peninsular record with the exception of Almenara-Casablanca 639 1, a site from the last Pliocene and located on the Iberian Mediterranean coast. Indeed, the measurements 640 on the mainland remains of Tyto balearica overlap with those from Sardinia and Corsica. 641 The rejection of the continental record of Tyto balearica is based on the interpretation consisting 642 in an owl, larger than the extant Tyto alba, spread over the west of the Mediterranean, appearing in the 22 643 Plio-Pleistocene boundary and becoming extinct in Corsica around the middle to late Pleistocene. Implicit 644 in this reasoning is to take Tyto balearica as a biological species. An alternative explanation, knowing 645 that Tyto alba was widely extended over the continent from the early Pleistocene onwards (see Tyrberg 646 1998), would be that the fossil remains from Sardinia and Corsica might represent a process of insular 647 differentiation from populations of Tyto alba, which avoids the chronological and geographical gaps 648 between the localities in the Balearic Islands as well as in Sardinia and Corsica. 649 650 Acknowledgments 651 I appreciate very much the criticism of David M. Alba, Hanneke Meijer and two anonymous 652 reviewers to a previous version of the manuscript, which has greatly improved this paper. 653 654 References 655 656 Adrover, R., Alcalá, L., Mein, P., Moissenet, E. & Orrios, J. (1986). Mamíferos del Turoliense medio en 657 la rambla de Valdecebro (Teruel). Estudios Geológicos 42, 495–509. 658 Agustí, J., Cabrera, L. & Moyà-Solà, S. (1985). Sinopsis estratigráfica del Neógeno de la fosa del Vallés- 659 Penedés. 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(j–p) Palaeortyx brevipes / Palaeortyx grivensis: 810 IPS87218, left carpometacarpus in ventral view (j); MNCN 74029, left tarsometatarsus in dorsal view (k); 811 MNCN 74033, left humerus in caudal view (l); MNCN 74030, left coracoideum in dorsal aspect (m); 812 without number (Batallones 1), left humerus in caudal view (n); without number (Batallones 1), right 813 scapula in lateral view (o); IPS12678, right carpometacarpus in ventral view (p). (q, r) Palaeortyx 814 phasianoides: IPS12647, left humerus in caudal view (q); MNCN 74028, right carpometacarpus in 815 ventral view (r). (s–u) Palaeortyx gallica: IPS73336, right tibiotarsus in cranial view (s); MNCN 74027, 816 left tibiotarsus in caudal view (t); IPS12650, right humerus in cranial view (u). (v, w) Tyto balearica: 817 IPS12652, right humerus in cranial aspect (v); IPS12673, pedal phalanx in lateral view (w). (x) 818 Accipitridae indet.: IPS12674, pedal phalanx in plantar view. (y) Accipitridae indet.: IPS12676, pedal 28 819 (ungual) phalanx in lateral view. (z) Recurvirostra teruelensis: IPS12677, right humerus in cranial view. 820 (a’) Charadriiformes indet.: IPS56, left tarsometatarsus in dorsal aspect. Scale bar equals 1 cm.[planned 821 for page width] 822 823 Figure 2. Tarsometatarsi of Tyto balearica from Valdecebro 5 (a–f) and Layna (g–j), in dorsal (a, c, e, g, 824 i) and proximal (b, d, f, h, j) views. Scale bar equals 1 cm. [planned for column] 825 826 Table 1. Measurements (in mm) of forelimbs of large-sized European Miocene galliforms. References are 827 indicated as superscripts: 1, Göhlich (2002); 2, Bocheński (1987); 3, Sánchez Marco (2006); 4, this study. 828 Notes: *, Miophasianus altus for Göhlich (2002); **, Miophasianus altus for Villalta and Crusafont Pairó 829 (1950). Number of specimens in brackets. 830 831 Table 2. Measurements (in mm) of strigiforms. References are indicated as superscripts: 1, Langer 832 (1980); 2, Louchart (2002); 3, Mourer-Chauviré et al. (1980); 4, Mourer-Chauviré and Sánchez Marco 833 (1988); 5, Sánchez Marco (2001); 6, Ballmann (1973); 7, Jánossy (1991); 8, Mlíkovský (1998); 9, Pavia 834 and Mourer-Chauviré (2011); 10, Göhlich and Ballmann (2013); 11, this study. Number of specimens in 835 brackets. 836 837 Table 3. Some taxa found in Miocene Iberian localities. Localities: 1, Fallol; 2, Can Mas; 3, La Artesilla; 838 4, Hostalets de Pierola indeterminate; 5, Toril 3; 6, Escobosa; 7, Los Valles de Fuentidueña; 8, Can 839 Poncic; 9, Viladecavalls; 10, Batallones 1 and 3; 11, Valdecebro 5; 12, Los Mansuetos. 840 841 Table S1. Measurements (in mm) of scapulae and humeri of medium and small-sized European Miocene 842 galliforms. References are indicated as superscripts: 1, this study; 2, Göhlich (2002); 3, Göhlich and 843 Mourer-Chauviré (2005); 4, Cheneval (2000); 5, Göhlich and Pavia (2008); 6, Ballmann (1969); 7, 844 Cheneval and Adrover (1993). Number of specimens in brackets. 845 846 Table S2. Measurements (in mm) of coracoidea of medium and small-sized European Miocene 847 galliforms. References are indicated as superscripts: 1, Göhlich and Mourer-Chauviré (2005); 2, 29 848 Ballmann (1969, as Palaeortyx depereti); 3, Cheneval and Adrover (1993); 4, Göhlich and Pavia (2008); 849 5, this study. Number of specimens in brackets. 850 30 Table 3. Some taxa found in Miocene Iberian localities. Localities: 1, Fallol; 2, Can Mas; 3, La Artesilla; 4, Hostalets de Pierola indeterminate; 5, Toril 3; 6, Escobosa; 7, Los Valles de Fuentidueña; 8, Can Poncic; 9, Viladecavalls; 10, Batallones 1 and 3; 11, Valdecebro 5; 12, Los Mansuetos. 1 2 3 4 5 6 7 8 9 10 11 12 MN7+8- MN3 MN4 MN4 MN7+8 MN7+8 MN9 MN9 MN10 MN10 MN12 MN12 MN9 Miophasianus altus • • Miophasianus medius • • Palaeortyx joleaudi • Palaeortyx • • • • • • brevipes/grivensis Palaeortyx • • phasianoides Palaeortyx gallica • • • Tyto balearica • • Accipitridae indet. • • • R. teruelensis • Charadriiformes indet. • Passerida indet. • 16 17 ck 18 19 20 21 Table 1. Measurements (in mm) of forelimbs of large-sized European Miocene galliforms. References are indicated as superscripts: 1, Göhlich (2002); 2, Bocheński 22 23 (1987); 3, Sánchez Marco (2006); 4, this study. Notes: *, Miophasianus altus for Göhlich (2002); **, Miophasianus altus for Villalta and Crusafont Pairó (1950). 24 25 26 Number of specimens in brackets. 27 28 29 30 Humerus Age Gl Pw Pd Wd Dw Dd 31 32 M. altus (La Grive)1 MN7+8 103.2 24.3–26.1 (4) — 9.9 20.7–21.1 (2) — 33 34 M. altus (Steinheim)1 MN7+8 100.5 ca. 25.9–27.0 (2) — 9.6–10.0 (2) 19.7–21.0 (2) — 35 36 M. medius* (Sandelzhausen)1 MN5 — ca. 18.0 — — ca. 17.5 — 37 38 M. medius (La Grive)1 MN7+8 — 17.9 7.4–7.6 (2) — — — 39 40 MN6- 41 M. medius (Przeworno II)2 — >15.5–16.4 (2) 6.9 (2) — — — 42 MN7+8 43 44 M. medius (Toril 3)3 MN7+8 — — — — 17.2 8.7 45 46 M. altus (Hostalets de Pierola, MN7+8- 47 — — — — 19.0–19.5 (2) – 48 IPS12649, IPS12651)4 MN9 49 50 M. medius (Hostalets de Pierola, IPS MN7+8- 51 — 17.6 — — — — 52 12648) 4 MN9 53 54 M. medius** (Hostalets de Pierola, IPS MN7+8- 55 — — — — 17.3 — 56 12649) 4 MN9 57 58 Ulna 59 60 61 62 63 64 65 16 17 18 19 20 21 22 M. altus (Sandelzhausen)1 MN 5 — 13.7 — — — — 23 24 MN7+8- 25 M. altus (Can Mas, IPS 12671) 4 — 13.6 — — — — 26 MN9 27 28 MN7+8- 29 M. altus (Can Mas, IPS 12672) 4 — — — — 12.2 — 30 MN9 31 32 Radius 33 34 M. medius (Batallones 3) MN10 62.5 5.7 5.6 2.8 7.5 3.5 35 36 Femur 37 38 M. altus (Steinheim)1 MN7+8 101 21 — — 19 — 39 40 M. altus (Dechbetten)1 MN5 100 20 — — 20 — 41 42 M. altus (Sandelzhausen)1 MN5 116 24.0 14.3 9.5 21.7 17.5 43 44 M. altus (Toril 3)3 MN7+8 87.3 20.0 11.9 8.2 17.4 — 45 46 MN7+8- 47 M. altus (Can Mas, IPS12670)4 — — 13.7 8.7 — — 48 MN9 49 50 Tibiotarsus 51 52 M. altus (Sansan)1 MN6 — — — 7.8 ca. 16 15.1 53 54 M. altus (La Grive)1 MN7+8 — — — 8.3 13.3–15.8 (2) 12.2–15.8 (3) 55 56 M. altus (Steinheim)1 MN7+8 — — — — 15–16 (3) 15–16 (3) 57 58 59 60 61 62 63 64 65 16 17 18 19 20 21 ca.12.6–ca.16.3 22 M. altus (Sandelzhausen)1 MN5 ca.188 — — 8.8 12.9–17.5 (3) 23 (4) 24 25 26 M. altus (Toril 3)3 MN7+8 147.5 17.9 16.2 8.7 11.8–14.9 (3) 11.6–15.0 (3) 27 28 M. medius (La Grive)1 MN7+8 — — — — ca.10 ca.10.1–10.3 29 30 MN6- 31 M. medius (Przeworno II)2 — — — — 10.7 ca.10.5 32 MN7+8 33 34 M. altus (Can Mas, IPS 12668)4 MN4 — — — — 16.0 15.0 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 Table 2. Measurements (in mm) of strigiforms. References are indicated as superscripts: 1, Langer (1980); 2, Louchart (2002); 3, Mourer-Chauviré et al. (1980); 4, Mourer-Chauviré and Sánchez Marco (1988); 5, Sánchez Marco (2001); 6, Ballmann (1973); 7, Jánossy (1991); 8, Mlíkovský (1998); 9, Pavia and Mourer-Chauviré (2011); 10, Göhlich and Ballmann (2013); 11, this study. Number of specimens in brackets. Humerus Ulna Tibiotarsus Tarsometatarsus Biozone max. length distal width prox. width distal width distal width max. length prox. width distal width 73.7–85.3 55.4–64.6 Tyto alba1 — 11.6–13.5 (28) 8.1–9.4 (26) 6.3–7.1 (26) 8.5–10.0 (29) 8.4–9.9 (29) 9.5–11.2 (28) (28) (29) Middle Tyto balearica (Corsica Pleistocen 110.6 15.2–16.6 (4) 11.1 8.3–9.2 (8) 10.5–12.7 (12) 72.5 11.8 13.4 and Sardinia)2 e Tyto balearica (Canet MN17 102.0 16.4–16.9 (2) — — — — — 14.4 and S’Ònix)3 Tyto balearica (Almenara-Casablanca MN17 — — — — — — — 13.1 1)4 Tyto balearica (Balaruc MN16 — — >10.4 — — — — — II)4 Tyto balearica MN16 — — — — 10.5 — — 11.6 (Moreda)5 Tyto balearica (Sète)4 MN15 — — >9.7 >7.2 — — — 12.9 Tyto balearica MN15 — 15.2–16.9 (4) 10.1 — — — 10.6–12.1 (4) 11.5–13.0 (2) (Layna)5,11 MN14– Tyto robusta (Gargano)6 — ca. 24.0 — ca. 11.0 ca. 15.0 — ca. 15.0 — MN15 Tyto balearica (=T. campiterrae Jánossy, MN13 — 14.0–14.5 (3) — — — 64.5–65.7 (2) 10.2–12.5 (11) 11.2–11.4 (3) 1991) (Polgárdi)7 Tyto balearica MN12 — — — 7.9 11.6 — 10.6–11.6 (3) — (Valdecebro 5)5 Tyto sanctialbani MN10 — — — — — 10.1 — — (Kohfidisch)8 Tyto sanctialbani (La MN7+8 — — — — ca. 8.8 — ca. 9 10.8–10.9 (2) Grive)8 Tyto sanctialbani (La ca. 8.6–10.3 9.4–ca. 10.5 MN7+8 — 11.7–14.8 (12) — 6.8–7.9 (3) 58.1–60.0 (2) 10.4–11.9 (18) Grive) 9 (13) (7) Tyto sanctialbani (Toril MN7+8 86.9 13.0 — — — — — — 3)5 Miotyto montispetrosi ca. 9.5–9.7 MN6 — 14.4 — 10.9–11.0 (2) 55.1–55.1 (2) 10.4–10.5 (2) 11.8–11.8 (2) (Steinberg)10 (2) Tyto balearica MN7+8– (Hostalets de Pierola, — 16.9 — — — — — — MN9 IPS12652)