C. Mourer-Chauviré would like to thank Pierce Brodkorb, Helen James, and Storrs Olson for their hospitality during her stays in the United States. The authors thank the reviewers Marco Pavia and Antoine Louchart for their useful comments which greatly improved the manuscript.
1The Ceyssaguet locality is situated in the French Massif Central, near the Loire valley, about 1000 m above sea-level. Several Early Pleistocene sites are situated in the same area, such as Senèze, Sainzelles, and Soleilhac (fig. 1) (Bonifay & Brugal, 1996). At Ceyssaguet the bones were found in an ancient volcano. After its explosion, a maar lake was formed inside the crater. Then, the outlet of the lake dug a small gorge in the surrounding basalts in which loess were deposited making possible the preservation of the fossil bones (Argant & Bonifay, 2011) (fig. 2). The fossils with CEY-2 acronym come from the inside of the crater.
2The age of this locality is relatively well established. The basalts of the oldest flow have been dated in 1985 by J.-M. Cantagrel (Department of Geology-Mineralogy, University of Clermont-Ferrand) at 1.4 Ma. This dating has been communicated in litteris and included into several publications but has not been formally published. The faunas, which come from the external side of the volcano were clearly deposited after its explosion, but during the existence of the lake. This period of time can be estimated at about 200,000 years at most, which implies that these faunas are about 1.2 Ma old (Bonifay & Brugal, 1996). This dating is in good agreement with the biostratigraphical profile of the whole of the fauna which corresponds to the latest Villafranchian (Bonifay, 1996). The different groups of mammals which have been published so far are the elephants (Aouadi & Bonifay, 1998), the Equidae (Aouadi, 2001), the Cervidae (Croitor & Bonifay, 2001), and the Carnivores (Tsoukala & Bonifay, 2004). The Ceyssaguet mammalian faunas belong to the group of Western European faunas known as “Tasso/Farneta group” (Alberdi et al., 1998). It is a transitional group which includes Villafranchian forms which do not persist in the galerian faunas which succeed them. At Ceyssaguet, these Villafranchian forms are associated to rare taxa such as Panthera gombaszoegensis, Canis falconeri, Lynx lynx and Palaeoloxodon antiquus, which announce the following times.
Fig. 1: Location map of the Ceyssaguet locality
Fig. 2: Ceyssaguet (Haute-Loire). Location of the fossil sites at the time of their formation on the slopes of the volcano and inside the crater
3The anatomical terminology follows Baumel and Witmer (1993). The material is deposited in the Musée National de Préhistoire, at Les Eyzies, Dordogne. Abbreviations: dig., digit ; maj., majus ; MN, Mammal Neogene (Guérin, 1982) ; MNQ, Mammal Neogene and Quaternary (Guérin, 2007) ; mtc., metacarpus ; mtt., metatarsus ; phal., phalanx ; post., posterior ; tr., trochlea ; USNM, United States National Museum of Natural History.
4Family Phalacrocoracidae (Bonaparte)
5Genus Phalacrocorax Brisson
6Phalacrocorax carbo (Linnaeus)
7Material: CEY 92 9933 Left coracoid, omal part and shaft (pl. 1, fig. 3-4) ; CEY 92 9938 Left scopus claviculae ; CEY 92 9936 Left scapula almost complete (pl. 1, fig. 5-6) ; CEY 92 9939 Left humerus, almost complete (pl. 1, fig. 1-2) ; CEY 92 9935 and 9941 Left ulna, proximal and distal parts (pl. 1, fig. 7) ; CEY 92 9934 Left radius, proximal part ; CEY 92 9941 bis Left radius, distal part ; CEY 92 9932 Left os carpi ulnare ; CEY 92 9940 Left carpometacarpus, shaft of the mtc. maj. ; CEY 92 9937 Left phal. I dig. maj. ; CEY 92 9942 8th cervical vertebra and 9th, 10th, and 11th associated cervical vertebrae. This material comes from the same individual.
8Measurements: table 1
Pl. 1: Phalacrocorax carbo.
Left humerus, CEY 92 9939, 1/ caudal view ; 2/ cranial view. Left coracoid, omal part and shaft, CEY 92 9933, 3/ dorsal view ; 4/ ventral view. Left scapula, CEY 92 9936, 5/ lateral view ; 6/ coastal view. Left ulna, CEY 92 9935 + 9941, 7/ dorsal view. Accipiter gentilis. Left carpometacarpus, CEY 96 14911, 8/ ventral view. Corvus pliocaenus. Left coracoid, omal part, CEY-2 7722, 9/ dorsal view ; 10/ ventral view. Right ulna, proximal part, CEY-2 8384, 11/ cranial view. Right ulna, distal part, CEY-2 7722, 12/ ventral view. Scale bars: 2cm.
Pl. 2: Aquila claudeguerini n. sp.
Left tarsometatarsus, CEY-2 5726, holotype, 1/ dorsal view ; 2/ plantar view ; 3/ medial view ; 4/ lateral view ; 5/ distal view ; 7/ distal view. Left phalanx 1 post. dig. III, CEY-2 8218, paratype, 5/ dorsal view. Left phalanx 4 post. dig. III, CEY-2 5249, paratype, 6/ dorsal view. Scale bars: 2 cm
9The material has been compared with the largest possible number of recent cormorant species from the collections of the USNM and of Pierce Brodkorb, University of Florida. All the morphological characteristics of the Ceyssaguet form agree with Phalacrocorax carbo, the Great Cormorant. This species has a very large geographic distribution, from Arctic to Tropics (del Hoyo et al., 1992). It includes six different-sized subspecies: the largest one is P. carbo carbo (East Canada, Greenland, Iceland, Norway, British isles), and the smallest one is P. carbo lucidus (Coastal West and South Africa, inland East Africa). By its dimensions the Ceyssaguet form is included in the variation range of P. carbo sinensis (Central and South Europe to India and China) but this does not imply that it belongs to this subspecies. It differs simply by the humerus which is slightly more robust (ratio width of shaft in the middle x 100 / total length: 6.1) (tab. 1).
Tab. 1. Phalacrocorax carbo from Ceyssaguet, measurements of the main long bones, in mm, compared to recent P. carbo sinensis.
1/ For the distal width and depth of humerus, see figure 3; 2/ Width measured on the ventral face; 3/ Height from the top to the base of the facies articularis clavicularis; 4/ From the caudal edge of condylus dorsalis to tip of tuberculum carpale.
10A very large number of fossil forms of Phalacrocoracidae have been described, at least from the Early Oligocene to the Pleistocene (Mayr, 2009 ; Göhlich & Mourer-Chauviré, 2010). Some taxa are smaller than the Ceyssaguet form, others are larger. The forms comparable in size to the Ceyssaguet form are Phalacrocorax marinavis Shufeldt, 1915, P. brunhuberi (von Ammon, 1918), P. idahensis (Marsh, 1870), P. filyawi Emslie, 1995, P. chapalensis Alvarez, 1977, P. rogersi Howard, 1932, and P. gregorii De Vis, 1905.
11Phalacrocorax marinavis, from the Early Miocene of Oregon, is the size of small individuals of recent P. carbo but differs in some morphological details (Shufeldt, 1915). Phalacrocorax brunhuberi, from the Middle Miocene of Germany, was described as Ardea brunhuberi and was transferred to the genus Phalacrocorax by Brodkorb (1980), who also placed the species Phalacrocorax praecarbo von Ammon, 1918, from the same locality, in synonymy with it. P. brunhuberi is known by a proximal part of carpometacarpus and P. praecarbo by an omal part of coracoid. Both specimens have the same dimensions as European forms of the recent species P. carbo. A third taxon from the same locality, Botaurides avitus von Ammon, 1918, known by a cervical vertebra, has also been placed in synonymy with P. brunhuberi (Olson, 1985). Mlíkovský (1992) has placed these three taxa in synonymy with Phalacrocorax intermedius (Milne-Edwards, 1867) from the Middle Miocene of France. We disagree with this last synonymy because we think that P. intermedius is an Anhingidae; however this is beyond the scope of the present paper. For P. idahensis from the Pliocene of Idaho and Florida, and P. filyawi, from the Pliocene of Florida, some of the measurements (Murray, 1970 ; Emslie, 1995) are comparable to those of a small specimen of a recent P. carbo, but the relative proportions of the long bones are different. P. idahensis is slightly smaller but its tibiotarsus is more elongated, while in P. filyawi the femur is shorter but the tibiotarsus and tarsometatarsus are longer. P. chapalensis, from the Plio-Pleistocene of Mexico, is quite comparable in size with P. carbo, although the author writes that it generally exceeds P. carbo in size (Alvarez, 1977, p. 212). The holotype is a proximal part of tarsometatarsus some morphological characteristics of which differ from P. carbo, in particular the shape of the intercondylar prominence. According to the author, P. chapalensis is more closely related to the recent P. auritus, the Double-Crested Cormorant, but it is larger. P. rogersi, from the Early Pleistocene of California, is known by an almost complete coracoid. Its size is included in the size variation range of P. carbo but it differs from this species and from the Ceyssaguet form by morphological characteristics (Howard, 1932): in P. rogersi the shaft is dorso-ventrally deeper ; the tip of the processus acrocoracoideus, in internal view, is rounded, while it is more pointed in the Ceyssaguet form and P. carbo ; the processus acrocoracoideus does not overhang the sulcus m. supracoracoidei, while it overhangs in P. carbo and even forms a small tubercle, on the dorsal side, in the Ceyssaguet form ; the ventral face, distally to the facies articularis clavicularis, is smoothly convex, while it is flattened in the Ceyssaguet form and in P. carbo. An important morphological characteristic is the shape of the intermuscular line on the ventral face, but unfortunately it is not visible on the Ceyssaguet form. According to Howard (1932) P. rogersi is closer to the recent species P. pelagicus, the Pelagic Cormorant, and P. perspicillatus, the extinct Spectacled Cormorant. Lastly P. gregorii, from the Pleistocene of Australia, has been placed in synonymy with P. carbo by Boles (2010).
12Forms closely related to the recent species and designated as Phalacrocorax cf. carbo, have been reported from the Late Miocene of Ethiopia, age ca. 5.8-5.6 Ma (Louchart et al., 2008), the Early Pliocene of Chad, age ca. 5 Ma (Louchart et al., 2004), and the Early Pleistocene of Turkey, age ca. 1 - 0.9 Ma (Louchart et al.,1998).
13The recent species P. carbo has been reported in the Early Pleistocene of Olduvai Gorge, Tanzania, from the upper part of Bed II (Harrison and Walker, 1979 ; Harrison, 1980), the age of which is about 1.15 Ma (Hay, 1976). In the Palearctic it has been reported in the Early Pleistocene of ‘Ubeidiya, in Israel (Tchernov, 1980), dated at ca. 1.4 Ma based on biochronological and archaeological arguments (Tchernov et al. 1986 ; Tchernov, 1988 ; Guérin, 2007), then it has been reported in a few Middle Pleistocene localities (Tyrberg, 1998). Ceyssaguet is therefore among the oldest localities where this recent species was found.
14Family Accipitridae (Vieillot)
15Aquila claudeguerini n. sp.
16Holotype: CEY-2 5726 Left tarsometatarsus, almost complete (pl. 2, fig. 1-4, 7)
17Paratypes: CEY-2 8218 Left phalanx 1 post. dig. III (pl. 2, fig. 5) ; CEY-2 5249 Left phalanx 4 post. dig. III (claw) (pl. 2, fig. 6) ; CE-2 7545 sacrum.
18Etymology: This species is dedicated to our late colleague and friend, Claude Guérin, in recognition of his numerous works on Pleistocene faunas.
19Type-locality: Ceyssaguet, Lavoûte-sur-Loire, Haute-Loire, France.
20Age: Late Early Pleistocene, Late Villafranchian, 1.2 Ma.
21Diagnosis: Species of Aquila with a tarsometatarsus larger than in the recent species of this genus. Tarsometatarsus robust, as in A. chrysaetos, while in the other recent or extinct species of Aquila the tarsometatarsus is more slender. In plantar view sulcus flexorius shallow and fossa mtt. I very elongate.
22Measurements: tables 2, 3 and 4
Tab. 2: Aquila claudeguerini n. sp., measurements of the tarsometatarsus, in mm, compared to recent and extinct species of eagles.
1/ After Louchart et al. (2005), except for the width in the middle which is not given by these authors; 2/ After Ballmann (1973), some measurements have been taken directly on figure 2.
23The Ceyssaguet tarsometatarsus has been compared with all the recent genera of Accipitridae present in the collection of the USNM, Smithsonian Institution, in Washington D.C. These comparisons have shown that it can be referred to the recent genus Aquila.
24CEY-2 5726 Left tarsometatarsus, almost complete (pl. 2, fig. 1-4, 7)
25The proximal articular surface and a part of the bone situated below are not preserved. The cristae lateralis and medialis hypotarsi and the crista medianoplantaris are missing. The foramina vascularia proximalia are present. They are situated at the same level and visible on both faces, dorsal and plantar. The tuberositas m. tibialis cranialis is situated on the lateral side, next to the facies subcutanea lateralis. In A. chrysaetos, the tubercle is well individualized and clearly separated from the dorsal edge of the facies subcutanea lateralis, whereas in CEY-2 5726 it is very slightly individualized and merges with the dorsal edge of this facies.
26On the distal part, the cortical zone of the tr. mtt. II is missing, and so it seems slightly shorter than the tr. mtt. III, whereas in A. chrysaetos, both trochleae are the same length. The wing of this tr. is not preserved. On the medial side there is a very deep fovea lig. collateralis. As in A. chrysaetos, the tr. mtt. III shows two parallel ridges, and it is slightly oblique compared to the longitudinal axis of the bone. Also as in A. chrysaetos, the tr. mtt. IV is slightly shorter than the tr. mtt. III. The wing of this trochlea is plantarly directed and more elongate than the wing of the tr. mtt. IV of A. chrysaetos.
27The plantar face is almost flat in its proximal part, then it extends in a shallow sulcus flexorius, whereas this sulcus is clearly deeper in A. chrysaetos. The fossa mtt. I is very elongate. Its proximal part is situated directly in the prolongation of the crista plantaris medialis, and it is pointed. In A. chrysaetos, the fossa metatarsi I is shorter, there is an interval between the fossa and the crista plantaris medialis and its proximal part is rounded. Lastly, on the plantar face, the foramen vasculare distale is very small whereas it is much larger in A. chrysaetos.
28CEY-2 8218 Left phalanx 1 post. dig. III (pl. 2, fig. 5)
29The proximal part of this phalanx is missing. There are no morphological differences compared to A. chrysaetos.
30CEY-2 5249 Left phalanx 4 post. dig. III (claw) (pl. 2, fig. 6)
31A part of the proximal articular surface and the distal part are missing. On the medial side there is a longitudinal ridge, as in A. chrysaetos. The tuberculum flexorium is medio-laterally wide.
32CEY-2 7545 Synsacrum
33This synsacrum includes the whole of the vertebrae, except for the last sacral vertebra. The total length of this element is 123 mm, while the same length in a recent A. chrysaetos is only 93 mm.
34The Ceyssaguet form differs mainly from the recent species A. chrysaetos by its larger size. The tarsometatarsus is incomplete but, measuring the distance between the lateral proximal vascular foramen and the distal end, compared to the same distance in a recent specimen of A. chrysaetos, the total length can be estimated at 132 mm. In a recent population of 26 individuals, the maximal length of the tarsometatarsus is 112.1 mm (Louchart et al., 2005) (tab. 2). These large dimensions are also found in the pedal phalanges (tab. 3 and 4).
Tab. 3: Aquila claudeguerini n. sp., measurements of the phalanx 1 post. dig. III, in mm, compared to recent and extinct species of eagles.
1/ After Ballmann (1976).
Tab. 4: Aquila claudeguerini n. sp., measurements of the phalanx 4 post. dig. III, in mm, compared to recent and extinct species of eagles.
1/ After Louchart et al. (2005).
35Most of the other recent species of Aquila are smaller than A. chrysaetos, with the exception of A. audax (Latham), which lives in Australia, and of A. verreauxii Lesson, which lives in Africa (del Hoyo et al., 1994, vol. 2). However, in these two species, the tarsometatarsi are smaller than in A. claudeguerini, and they are more slender and elongate. Among the different recent species of Aquila, it is A. chrysaetos which has the stouter tarsometatarsi and for that reason A. claudeguerini is more similar to it. Its ratio DW/TL (0.237) is close to that of A. chrysaetos (0.236) and it is larger than the ratios of A. audax (0.223) and A. verreauxii (0.209) (tab. 2).
36The recent species, Aquila chrysaetos, and indeterminate species of the genus Aquila, have been reported in a few Early Pleistocene localities of the Palearctic (Tyrberg, 1998, 2008), but none of them can be attributed to the same species as the Ceyssaguet form, whose main characteristic is its larger size than that of Aquila chrysaetos.
37Aquila kurochkini Boev, 2013, from the Early Pleistocene (MN 17) of Varshets, Bulgaria, is the same size as the recent species Aquila fasciata, and smaller than A. chrysaetos (Boev, 2013). The elements attributed to this species had been previously attributed to different taxa (Boev, 2002). The genus Aquila was also reported in the Early Pleistocene (first half of the MN 18 zone) locality of Slivnitsa, Bulgaria (Boev, 1998, 2000), but later it was designated as cf. Aquilinae gen. (Boev 2002). Aquila cf. chrysaetos is reported from the locality of Villany 3, Hungary, dated of 1.8 Ma (MN 18) and this form is the same size as the recent A. chrysaetos (Jánossy, 1977). In Cava Sud, Soave, Italy (MNQ 19), is found an Aquila sp. of small size (Pavia & Bedetti, 2013). At ‘Ubeidiya, Israel, age 1.4 Ma, are found an Aquila chrysaetos, the remains of which show similarity with the recent form, an Aquila sp. approaching A. pomarina in size, and another Aquila sp. known by a terminal phalanx which “resembles that of a slightly larger eagle”, slightly larger than the previous one (Tchernov, 1980, p. 25). In China, in the Early Pleistocene locality of Zhoukoudian 18, Aquila sp. is known by a distal part of tarsometatarsus, the distal width of which is ca. 18 mm (Hou, 1993, pl. 2, fig. 9) while in recent A. chrysaetos it varies from 21.8 to 27.3 mm. Lastly, in Monte Peglia, Italy, age ca. 1.0 Ma (MNQ 20), there is an Aquila sp. the dimensions of which are comparable to those of Aquila clanga (Bedetti, 2003).
38Aquila nipaloides Louchart et al., 2005, from the Pleistocene of Corsica and Sardinia, is close to the Steppe eagle, Aquila nipalensis, but larger. Its tarsometatarsus is smaller than that of A. claudeguerini and more gracile (ratio DW/TL 0.213) (tab. 2). Aquila chrysaetos simurgh Weesie, 1988, from the Pleistocene of Crete, has been described as a subspecies of A. chrysaetos, but, according to Louchart et al. (2005), it cannot be attributed to the genus Aquila. Its tarsometatarsus is unknown. The species Aquila borrasi Arredondo, 1970, from the Pleistocene of Cuba, has been transferred by Suárez and Olson (2007) to the genus Buteogallus. The tarsometatarsi of this species are very long and elongate. In Aquila bivia Emslie and Czaplewski, 1999, from the Late Pliocene of Florida and Arizona, the size of the known elements is larger than the maximal size of the same elements in A. chrysaetos, but “the tibiotarsus is longer and proportionally more gracile compared to A. chrysaetos, suggesting that the fossil had relatively longer and more slender legs compared to other eagles of this genus” (Emslie & Czaplewski, 1999, p. 192). The species Spizaetus willetti Howard, 1935, from the Pleistocene of Nevada, only known by a distal part of tarsometatarsus, has been referred to the genus Spizaetus, but shows similarities with the genus Aquila (Louchart et al., 2005). Aquila claudeguerini also shows similarities with Spizaetus willetti, in particular in the very elongate shape of the fossa met. I and the very small opening of the distal vascular foramen on the plantar face. However A. claudeguerini differs from S. willetti in the following characteristics: tr. met. III wider ; tr. met. II slightly shorter than tr. met. III (longer in S. willetti) ; tr. met. II without articular furrow in distal view (articular furrow well expressed in S. willetti) ; tr. met. II with a deeply hollowed fovea ligamenti collateralis (not clearly visible in S. willetti) ; shaft narrowing markedly above trochleae (narrowing slightly in S. willetti). In Garganoaetus freudenthali Ballmann, 1973, from the Late Miocene of Gargano, Italy, the tarsometatarsus length is similar to that of A. claudeguerini, but the shaft and the distal end are clearly narrower (Ballmann, 1973, 1976) (tab. 2).
39Other genera and species of very large Accipitridae have been described in the Pleistocene of Northern America, Bahamas, Cuba, and New Zealand. Wetmoregyps daggetti Miller, 1915, from Southern California and Northern Mexico, has been transferred to the genus Buteogallus (Olson, 2007). Its tarsometatarsi are still larger than those of Buteogallus borrasi. Titanohierax gloveralleni Wetmore, 1937, from Little (see Olson & Hilgartner, 1982) Exuma Island, Bahamas, is known by an incomplete tarsometatarsus. Wetmore (1937) indicates that the total length of this tarsometatarsus may have been near that of Buteogallus daggetti, but it was much heavier and more robust. According to Olson & Hilgartner (1982, p. 27) “Titanohierax is not related to the eagles of the genera Aquila or Haliaeetus”. Gigantohierax suarezi Arredondo and Arredondo, 2002, from Cuba, has been described after a slightly incomplete femur, but the material also includes an incomplete tarsometatarsus 200 mm long. Lastly Harpagornis moorei Haast, 1872, from New Zealand, is known from an abundant material. A cladistics analysis brought to the conclusion that the genus Harpagornis was closely related to the genus Aquila (Holdaway, 1991). However the study of ADN has shown that it was closer to the genus Hieraaetus and that it should be designated as Hieraaetus moorei (Haast, 1872) (Bunce et al., 2005). The tarsometatarsus is from 131.5 to 166.4 mm long, with a mean value of 148.2 (n = 21).The distal width (33.1-44.6, mean 38.7, n = 22) is proportionally much larger than in A. claudeguerini (DW/TL 0.261) (Holdaway, 1991).
Fig. 3: Diagram showing the way of measuring the distal width (A) and distal depth (B) of the Phalacrocorax humerus.
40The species Aquila claudeguerini was probably of Pliocene origin and became extinct during the MPR episode, or Middle Pleistocene Revolution. This event occurred between 1.25 and 0.7 Ma, and corresponds to an increasing of the severity and duration of the Pleistocene cold stages. During this episode several species of Villafranchian mammals, mainly herbivores, vanished and were replaced by new forms coming from Africa or from Asia (Madurell‑Malapeira et al., 2014). It is possible that this climate change has led to the scarcity of the prey on which it was feeding, and then to its extinction and replacement by the recent species Aquila chrysaetos.
41Accipiter gentilis (Linnaeus), female size
42Material: CEY 96 layer2 N11 14911 Left carpometacarpus (pl. 1, fig. 8)
43This carpometacarpus is made up of two fragments which cannot be joined because a small part is missing between them. Its morphological characteristics make it possible to ascribe it to the species Accipiter gentilis, the Northern Goshawk. In this species females are conspicuously larger than males. The total length of this bone can be estimated at 66 mm and its proximal width is 16.2 mm. In the recent females the total length of the carpometacarpus varies from 59.8 to 64.5 mm (n=12) and the proximal width from 14.4 to 15.9 mm (Otto, 1981). The Ceyssaguet Goshawk is thus slightly larger than the recent forms.
44A. gentilis has rarely been reported in the Early Pleistocene. The subspecies A. gentilis brevidactylus, described in the site of Grotte de l’Escale, at Saint‑Estève‑Janson (MNQ 22 zone) (Mourer-Chauviré, 1975), has been reported as A. gentilis cf. brevidactylus in several other Middle Pleistocene localities (Tyrberg, 1998). This subspecies is characterized by a size generally smaller than that of the recent forms. Thus the Ceyssaguet form corresponds to a different form.
45Family Corvidae Vigors
46Genus Corvus Linnaeus
47Corvus pliocaenus (Portis, 1887)
48Material: CEY-2 7722 Group of bones from the same individual, including a left coracoid, omal part and shaft, a shaft of left humerus, a right humerus, distal part, incomplete, a right ulna, distal part, and eight pedal phalanges more or less complete (pl. 1, fig. 9-10, 12) ; CEY-2 8384 Right ulna, proximal part and shaft. The proximal articular surface is incomplete, the olecranon is missing (pl. 1, fig. 11).
49The species Corvus pliocaenus, described in the Villafranchian of Valdarno, Italy, is a form intermediate in size between the recent species Corvus corone and Corvus frugilegus on one side, and Corvus corax, on the other side. This species, including the subspecies Corvus pliocaenus janossyi, has been reported from a great number of Early and Middle Pleistocene localities (zones MN 18 to MNQ 24) in France, Spain, Italy, Greece, Germany, and the Czech Republic (Tyrberg, 1998). The measurements are given in table 5.
Tab. 5: Corvus pliocaenus, measurements of the coracoid and ulnae of, in mm, compared to recent corvids and to Corvus pliocaenus janossyi from Saint-Estève-Janson.
1/ After Tomek and Bocheński (2000); 2/ Measurements of adults only; 3/ From caudal edge of condylus dorsalis to tip of tuberculum carpale.
50Pollen analyses performed on hyaena coprolites have indicated a predominance of tree pollens. These pollens indicate a juxtaposition of biotopes, which can be explained by the topography, “under temperate conditions, locally humid and fresh, with forest formations then possibly occupying the slopes of the volcanic relief” (Argant & Bonifay, 2011, p. 9, our translation). The occurrence of a cormorant is probably related to the presence of a lake, but this recent species, P. carbo, has a very wide geographic distribution and can adapt itself to a large number of biotopes. The extinct species, Aquila claudeguerini and Corvus pliocaenus, do not provide information about the palaeoenvironment. Accipiter gentilis, the recent Northern Goshawk, lives in wooded areas, particularly in coniferous forests, but also in deciduous or mixed forests (del Hoyo et al., 1994).
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