Corvids from the Neogene and Quaternary of the Carpathian Basin (1 Viewer)

Kessler, J. (2020)
Evolution of corvids and their presence in the Neogene and the Quaternary in the Carpathian Basin
Ornis Hungarica 28: 121-168
doi: 10.2478/orhu-2020-0009
http://www.ornis.hu/?download&aid=414&volume_id=39&lang=eng

Corvids are the largest songbirds in Europe. They are known in the avian fauna of Europe from the Miocene, the beginning of the Neogene, and are currently represented by 11 species. Due to their size, they occur more frequently among fossilized material than other types of songbirds, and thus have been examined to the largest extent. In the current article, we present their known evolution and their fossilized taxa in Europe and examine the osteology of extant species.

Systematic paleontology - 1

Systematic paleontology
Fam. Corvidae (Vigors, 1825) Corvids are the largest songbirds in Europe, hence they are unmistakable with any other members of songbird families. They live from plains to snowy mountains. They have sedentary,
as well as migratory, species. They nest in crevices of rocks or on trees. They are omnivores. Their species shift occurred mainly in the Pleistocene. They are well represented in fossilized materials both in species and in numbers. This is not only due to their large population, seasonal constancy and relatively slower flight, but mostly due to their remains that are large enough to be detected via traditional collection methods.

Genus Corvus Linnaeus, 1758
Corvus † pliocaenus (Portis, 1889) / syn. C. †betfianus Kretzoi, 1962; (Table 1)
Site and era: Polgárdi, Upper Miocene (MN 13) (Hungary) (Kessler 2010); Beremend 26, Lower Pliocene (MN 15) (Kessler 2010); Beremend 15, 18, Upper Pliocene (MN 16) (Jánossy 1992, 1996); Q1: Beremend 16, 17 (Jánossy 1992, 1996) (all in Hungary); Betfia 9 (Gál 2002) (Romania); Q2: Betfia „Aven” (Kessler 1975, Jánossy 1979, Gál 2002), Betfia 5 (Kretzoi 1962, Kessler 1975, Jánossy 1979, Gál 2002) (all in Romania); Q3: Tarkő 2 (Jánossy 1979) (Hungary). From sites in Europe outside the Carpathian Basin Q1-2: Czech Republic, France, Germany, Spain; Q3: France (Tyrberg 1998).
We described a similar species, Corvus simionescui (Kessler, 1979) (Mălușteni-Berești, Romania, Lower Pliocene, MN 15) next to the Carpathian Basin, Mlíkovskỳ (2002) classified it to the recent Corvus corone species as well. A common characteristic of taxa described with different names is that they range in size between crows and ravens. Mlikovskỳ (2002) classifies certain taxa (C. antecorax, C. pliocaenus janossyi) to ravens, while others (Corvus pliocaenus, C. betfianus) to the recent Hooded Crow (C. corone). In our opinion, it is a transitional species that had gone extinct in the Middle Pleistocene. Due to the age and dimensions of the finds, we deem the species Corvus pliocaenus as valid. This is the oldest corvid of this size in Europe.

Corvus † hungaricus Lambrecht, 1916 /syn. C. antecorax Mourer-Chauviré, 1975; C. pliocaenus janossyi Mourer-Chauviré, 1975 Site and era: Q1: Beremend 17 (Kessler 2010), Nagyharsányhegy 2 (all in Hungary) (Lambrecht 1916, Kessler 2010).
Material: distal end of right humerus, right tarsometatarsus (as holotype) (Nagyharsányhegy); distal end of humerus, 2 proximal end of carpometacarpus, proximal end of femur, 3 fragments of tarsometatarsus (1 proximale, 2 distale), (Beremend 17). Dimensions: tarsometarsus A = 67 mm, C = 10 and 11.29 mm, E = 4.0 and 4.51, 4.27 mm, F = 8.67 and 7.94 mm, G = 5.26 and 4.56 mm; humerus C = 18.69 mm, D = 10.08 mm, F = 13.5 mm; carpometacarpus C = 12.68 and 12.36 mm, D = 6.91 and 6.94 mm. It is a corvid with a lean skeleton and a size between crows and ravens. Mlikovskỳ (2002) suggests reevaluation of the taxon, which we conducted, and the results confirmed the Original diagnosis. In addition, we found the proximal fragment of a humerus as well at the site of Nagyharsány mountain. Its identification from Beremend also confirms the reality of the taxon, and since other raven-sized species known from the fossilized and recent avian fauna of Europe were described later, we suggest keeping the taxon based on the principal of priority as well. From Late Pliocene – Early Pleistocene and Middle Pleistocene of France (Senéze, Saint-Estéve-Janson, Lunel Viel, MN 17-18, Q2 and Q3) the Corvus antecorax (Mourer-Chauviré, 1975) and Corvus pliocaenus janossyi (Mourer-Chauviré, 1975) finds of similar sizes are known (Mourer-Chauviré 1975).

Corvus † harkanyensis Kessler, 2010
Site and era: Csarnóta 2, Upper Pliocene (MN 15-16) (Hungary) (Kessler 2010).
Material: distal end of right humeus – as holotype; distal end of left tibiotarsus – as paratype.
Dimensions: humerus: E = 4.90 mm; F = 11.53 mm; G = 5.89 mm; H = 6.20 mm; tibiotarsus E= 3.10 mm; F = 5.60 mm; G = 5.71 mm;
A corvid with the size of a Jackdaw, at the distal epiphysis of which in cranial view (Figure 17A) the tuberculum supracondylare ventrale (a) is more prominent than the present Corvus species, the condylus ventralis (c) is not ovoid but round in shape, the epicondylus ventralis (e), though damaged, is much wider and stouter, the processus flexorius (b) is curled up. On the distal end of the tibiotarsus (Figure 17B), in cranial view the end of the sulcus extensorius (c) above the pons supratendineus (b) is not rounded but has a pointed end (Kessler 2010). Its name stems from the nearby town of Harkány. The fossilized species bearing characteristics and dimensions of the recent jackdaw was probably the ancestor of the form we see today in the Carpathian Basin. Smaller Corvus species are only known from the Upper Pliocene (C. cf. monedula: Văršec Bulgária, MN 17; Boev 1995, 2000) among the avian fauna of Europe. From the Carpathian Basin and its immediate vicinity from the early Pleistocene of Stránská skála, Czech Republic, from where the Corvus moravicus (Mlikovskỳ 1995) extinct species was described, but the person describing it subsequently revoked it and reclassified the material as C. monedula (Mlikovskỳ 2002). The latter had been identified from several Middle and Upper Pleistocene sites (Jánossy 1979, Gál 2002).

Corvus monedula Linnaeus, 1758 / syn. Corvus cf. † moravicus Mlíkovskỳ, 1996 Q1: Betfia 2, 9 (Kormos 1913, Čapek 1917, Lambrecht 1933, Kessler 1975, Jánossy 1979, Gál 2002); Q1-2: Betfia „Aven” (Kessler 1975, Jánossy 1979, Gál 2002) (all in Romania); Q2: Nagyharsányhegy 1-4 (Lambrecht 1916, 1933, Jánossy 1979) (Hungary); Q3: Vindija (M. Malez 1961, M. Malez & Rukavina 1975, V. Malez 1973, 1986, 1988) (Croatia); Q4/I: Merkenstein (Wettstein & Mühlhofer 1938), Mixnitz – Drachenhöhle (Lambrecht 1933) (all in Austria); Velika Pecina (V. Malez 1984, 1986, 1988) (Croatia); Bajót-Baits Cave, Bajót-Hóman Cave (Jánossy 1979), Csákvár-Eszterházy Cave (Lambrecht 1933, Kretzoi 1954, Jánossy 1979), Felsőtárkány-Peskő Cave (Lambrecht 1912, 1933, Jánossy 1979, 1986), Hámor-Puskaporos Cave (Lambrecht 1912a, 1912b, 1916, 1933, Jánossy 1979, 1986), Pilisszántói I. Cave (Lambrecht 1915, 1933, Jánossy 1979, 1986), Szilvásvárad-Istállóskői Cave (Lambrecht 1912, 1933, Jánossy 1952, 1955, 1979, 1986), Tata-Kálváriahegy no. 4. Cave (Gál 2004, 2005b), Tokod-Nagyberek (Jánossy 1979) (all in Hungary); Homoródalmási-Orbán Balázs Cave (Vîrghiş-Peștera Mare) (Kessler 1977, Gál 2002), Körösmart (Rîpa), (Jánossy in Hamar & Csák 1969, Kessler 1974b, Gál 2002), Nándor-Nándori Cave (Nandru-Peştera Curata) (Jánossy 1965, Fischer & Stephan 1977, Kessler 1985, Jurcsák & Kessler 1988, Gál 2002, 2003), Ohábaponor-Bordu Mare Cave (Ohaba Ponor-Peştera Bordu Mare) (Kessler 1985, Jurcsák & Kessler 1988, Gál 2002, 2003) (all in Romania); Q4/ II: Bodajk-Rigólyuk (Kordos 1984), Budapest-Sas György Place, Csepel Vízművek (Gál 2015), Csákvár-Esterházy Cave (Kretzoi 1954), Felsőtárkány-Petényi Cave (Jánossy 1979), Legény-Cave (Lambrecht 1914), Szendrő-Felsővár (Gál 2015), Széchény (Gál 2015), Székesfehérvár-Sziget (Gál 2015) (all in Hungary); Bégakalodva (Cladova) (Gál 2004), Kazánszoros-Töröklik Cave (Cazanele Mari, Peştera Cuina Turcului) (Kessler 1974a, Fischer & Stephan 1977), Kisbács-Bácsitorok (Baciu, Gura Baciului) (Kessler 2013a), Körösbánlaki Cave (Peştera din Bălnaca) (Kessler 1982), Peterd-Tordai-Hasadék-Magyar Cave (Petrești, Cheile Turzii-Peștera Ungurească) (Kessler & Gál 1998, Gál 2004), Remetelórév-Bólyikői Cave (Lorău-Peştera din Piatra Boiului) (Kessler 1982); Révi caves (Peşterile din Vadu Crișului) (Kessler 1982), Székelykeresztúr (Cristuru-Secuiesc) (Gál 2008, 2015), Szkerisoara-Sasok Cave (Scărişoara, Peştera Vulturilor) (Kessler 1982, Jurcsák & Kessler 1986, 1988), Vársonkolyos-Izbîndiş Cave, Vársonkolyos-Kis Magyar Cave (Şuncuiuş, Peştera Izbîndiş; Peştera Napiştileu) (Kessler 1977, Gál 2002) (all in Romania). From sites in Europe outside the Carpathian Basin Q1-2: Bulgaria, Czech Republic, France, Ukraine; Q3: Czech Republic, France, Germany, Spain, Ukraine; Q4: Austria, Belgium, Bosnia-Herzegovina, Bulgaria, Croatia, Czech Republic, France, Germay, Greece, Ireland, Italy, Montenegro, Moldova, Poland, Portugalia, Russia, Schwitzerland, Spain, Ukraine, United Kingdom (Tyrberg 1998).
[Note: Gál (2002) identifies the species as C. † moravicus from the Betfia 9 site, but Mlikovskỳ (2002) reclassifies the fossil taxon as C. monedula based on the Corvidae size database published by Kessler and Moldvai (1993). We find it necessary to distinguish between present jackdaws from the Upper Pliocene-Lower Pleistocene and Upper Pleistocene at least on a subspecies level, named C. monedula † moravicus.]

Systematic paleontology - 2

Corvus corax Linnaeus, 1758
Q3/I: Hundsheim (Mlikovskỳ 2009) (Austria); Q3/II: Vindija (M. Malez 1961, M. Malez & Rukavina 1975, V. Malez 1973, 1986, 1988, Musil 1980) (Croatia); Q4/I: Velika Pecina (M. Malez & Rukavina 1975, V. Malez 1984, 1986, 1988) (Croatia); Bajót-Baits Cave, Bajót-Jankovich Cave (Jánossy 1979a, 1979b); Budapest-Remetehegyi Cave (Kormos 1914, Lambrecht 1933, Jánossy 1979, 1986), Felsőtárkány-Peskő-Cave (Lambrecht 1912a, 1912b, 1933, Jánossy 1979, 1986), Hámor-Puskaporos Cave (Lambrecht 1912a, 1912b, 1916, 1933, Jánossy 1979, 1986), Kesztölc-Bivak Cave (Jánossy 1979), Pilisszántói I. Cave Lambrecht 1915, 1933, Jánossy 1979, 1986), Répáshuta-Balla Cave (Lambrecht 1912a, 1912b, 1933), Tatabánya-Szelim Cave (Jánossy 1979a, 1979b) (all in Hungary); Nándor-Nándori Cave (Nandru-Peştera Curata) (Jánossy 1965, Fischer & Stephan 1977, Kessler 1985, Jurcsák & Kessler 1988, Gál 2002, 2003), Ohábaponor-Bordu Mare Cave (Ohaba Ponor-Peştera Bordu Mare) (Kessler 1985, Jurcsák & Kessler 1988, Gál 2002, 2003) (all in Romania); Q4/II: Felsőnyék-Várhegy (Gál 2004, 2015), Felsőtárkány-Petényi Cave (Jánossy 1979), Mélyvölgy (Jánossy 1979), Pilisszentkereszt, Szendrő-Felsővár (Gál 2015) (all in Hungary), Kazánszoros-Climente Cave (Cazanele Mari-Peştera Climente I) (Kessler 1981, Gál 2002); Kazánszoros-Töröklik Cave (Cazanele Mari, Peştera Cuina Turcului) (Kessler 1974a, Fischer & Stephan 1977), Kazánszoros-Icoana Cave (Cazanele Mari, Peştera Icoana) (Kessler 1985, Jurcsák & Kessler 1986, 1988), Kovászna (Covasna) (Bindea 2008), Körösbánlaki Cave (Peştera din Bălnaca) (Kessler 1982) (all in Romania). From sites in Europe outside the Carpathian Basin Q1-2: Greece; Q3: Azerbaijan, Croatia, Czech Republic, France, Ukraine; Q4: Armenia, Austria, Belgium, Bosnia-Herzegovina, Bulgaria, Croatia, Czech Republic, France, Georgia, Germany, Greece, Italy, Montenegro, Moldova, Norway, Poland, Portugalia, Russia, Switzerland, Spain, Ukraine, United Kingdom (Tyrberg 1998).

Corvus corone Linnaeus, 1758
Q3: Vindija (M. Malez 1961, M. Malez & Rukavina 1975, V. Malez 1973, 1986, 1988, Musil 1980) (Croatia); Q4/I: Budapest-Remetehegyi Cave (Kormos 1914, Lambrecht 1933, Jánossy 1979, 1986), Szilvásvárad-Istállóskői Cave (Lambrecht 1912, 1933, Jánossy 1952, 1955, 1979, 1986), Varbó-Lambrecht Kálmán Cave (Jánossy 1964, 1979) (all in Hungary); Körösmart (Rîpa) (Jánossy in Hamar & Csák 1969, Kessler 1974b, Gál 2002), Ohábaponor-Bordu Mare Cave (Ohaba Ponor-Peştera Bordu Mare) (Kessler 1985, Jurcsák & Kessler 1988, Gál 2002, 2003), Rév-Pince Cave (Vadu Crişului, Peştera Pincelului) (Gál 2002) (all in Romania); Q4/II: Teufelslucke (Söergel 1966) (Austria); Bajcsa-Castle (Gál 2015), Balatonlelle-Kenderföldek (Gál 2005b), Felsővadász-Várdomb, Hajdúnánás (Gál 2015), Nagysomlyói Fosse (Kessler 2010) (all in Hungary); Kazánszoros-Töröklik Cave (Cazanele Mari, Peştera Cuina Turcului) (Kessler 1974a, Fischer & Stephan 1977), Körösbánlaki-Cave (Peştera din Bălnaca) (Kessler 1982), Peterd-Tordai-hasadék – Magyar Cave (Petrești, Cheile Turzii-Peștera Ungurească) (Kessler & Gál 1998, Gál 2005a), Remetelórév-Bólyikői Cave (Lorău-Peştera din Piatra Boiului) (Kessler 1982), Révi caves (Peşterile din Vadu Crișului) (Kessler 1982), Szkerisoara-Sasok Cave (Scărişoara, Peştera Vulturilor) (Kessler 1982, Jurcsák & Kessler 1986, 1988), Vársonkolyos-Kis Magyar Cave, Vársonkolyosi caves (Şuncuiuş-Peştera Napisteleu-Peşterile din Şuncuiuş) (Kessler 1977, Gál 2002) (all in Romania); Padina (Classon 1980, Gál 2004) (Serbia). From sites in Europe outside the Carpathian Basin Q3: Azerbaijan, Croatia, Czech Republic, France, Germany, Greece, Spain; Q4: Austria, Belgium, Bosnia-Herzegovina, Croatia, Czech Republic, France, Georgia, Germany, Italy, Malta, Portugalia, Russia, Switzerland, Spain, Ukraine, United Kingdom (Tyrberg 1998).

Corvus frugilegus Linnaeus, 1758
Q4/I: Felsőtárkány-Peskő Cave (Lambrecht 1912a, 1912b, 1933, Jánossy 1979, 1986), Pilisszántói I. Cave (Lambrecht 1915, 1933, Jánossy 1979, 1986) (all in Hungary); Q4/II: Bajcsa-Castle (Gál 2002, 2015), Balatonkeresztúr-Réti dűlő (Gál 2004), Bodajk-Rigólyuk (Kordos 1984), Endrőd 39 (Gál 2005a), Pilismarót-Malompatak (Jánossy 1985), Széchény (Gál 2015), Szendrő (Gál 2005b, Tassi 2006), Tác-Gorsium (Bökönyi 1984, Jánossy 1985), Visegrád-Palace (Gál 2015) (all in Hungary); Kazánszoros-Töröklik Cave (Cazanele Mari,
Peştera Cuina Turcului) (Kessler 1974a, Fischer & Stephan 1977), Körösbánlaki Cave (Peştera din Bălnaca) (Kessler 1982), Vársonkolyosi caves (Kessler 1977, Gál 2002) (Şuncuiuş-Peşterile din Şuncuiuş) (all in Romania); Padina (Classon 1980, Gál 2004) (Serbia). Q3: Azerbaijan, Germany, Greece; Q4: Belgium, Bosnia-Herzegovina, Croatia, France, Germany, Greece, Ireland, Italy, Netherlands, Ukraine, United Kingdom (Tyrberg 1998).

Corvus corone/frugilegus [sic] = Corvus corone cornix [FR]
Q4/II: Kaposújlak-Várdomb, Paks-Gyapa, Dombóvár-Tesco (Gál 2017); [Note: there are only very small morphological differences between the skeletal parts of the Hooded Crow (Corvus corone cornix) and the Rook (C. frugilegus), but the bones of the former species are slightly more robust. Incidentally, the subspecies – Carrion Crow, which is widespread in the western and southern parts of Europe, is also pure black, like the juvenile crow. The latter is rarely reported from fossil finds, probably because of the great
similarity. Presumably, it occurs in many materials because the two species could not be separated. All indications are that a separation of the two species may have occurred recently, probably in the Holocene.]

Corvus sp. foss. indet.
Site and era: Litke 2 (MN 5) (Kessler & Hír 2011), Polgárdi 4 (MN 13) (Jánossy 1991, 1995), Beremend 26 (MN 15) (Kessler 2010), Villány 3 (MN 16) (Kessler 2010) (all in Hungary).
[Note: material from the Neogene mostly consists of claws. Claws of corvids have a typical shape, but since there is a size difference between the four claws, and those belonging to species of more or less similar sizes cannot be reliably distinguished, we can only classify them to the genus level.]

Corvus sp. indet.
Q1: Villány 5 (Kessler 2010), Beremend 17 (Jánossy 1991, 1992) (all in Hungary); Q4/I: Szárazgerence (Jánossy 1979, 1986), Varbó-Lambrecht Kálmán Cave (Jánossy 1964, 1979) (all in Hungary), Detrekőszentmiklós-Pálffy Cave (Dzeráva Skála-Plavecky Mikulas) (Lambrecht 1913, 1933) (Slovakia); Q4/II: Ecsegfalva (Pike-Tay et al. 2004, Gál 2007), Tatabánya-alsó – Törekvés Cave (Kessler 2010), Maroslele-Pana (Bökönyi 1964, Jánossy 1979, 1985, Gál 2005a, 2005b), Szolnok-Szanda (Jánossy 1985, Gál 2005a, 2005b) (all in Hungary).
Spread: From the Late Pliocene of Spain and Bulgaria (Puebla de Valverde, as well as Slivnica, MN 17-18) Corvus finds described to the genus level are known from the Neogene of Europe (Mlíkovskỳ 2002).

† Miocorvus Lambrecht 1933
Miocorvus † larteti (Milne-Edwards, 1871)
Site and era: Tasádfő (Tăṣad, Romania), Middle Miocene (MN 7) (Gál & Kessler 2006, Kessler 2010); Mátraszőlős 3, Middle Miocene (MN 7/8); Rudabánya, Upper Miocene (MN 9) (Kessler 2010, Kessler & Hír 2012); Polgárdi 4, Upper Miocene (MN 13); Csarnóta 2, Beremend 26, Lower Pliocene (MN 15) (Kessler 2010) (all in Hungary).
Material: distal end of humerus (Mátraszőlős 3), distal end of carpometacarpus, phalanx 1. digiti II. alae (Csarnóta 2), 4 distal fragments of tibiotarsus (Rudabánya 3, Polgárdi 4, Beremend 26), 4 phal. pedis (Tasádfő).
Dimensions: humerus F = 10.11 mm; carpometacarpus F = 5.6 mm, G = 3.3 mm; phal. alae C = 3.36 mm, E = 4.69 mm, F = 3.43 mm; tibiotarsus E=2.49, 2.80 and 2.92 mm, F=5.44, 5.51, 5.65 and 5.78 mm, G=5.72 and 6.89 mm; tarsometatarsus F = 5.6 mm, G = 3.3 mm.
The small (jay-sized) crow described by Milne-Edwards (Corvus larteti Milne-Edwards, 1871) was renamed by Lambrecht (1933). We classified the material described by Jánossy (1979, 1997) from the Csarnóta 2 site here as well, but this is highly questionable due to the age difference. Outside of the typical site (the Middle Miocene of France) (Sansan, MN 6) (Milne-Edwards 1871), it was also reported in 2006 from the Middle Miocene of Tăṣad, Romania (MN 7) (Kessler & Venczel 2009) and the Middle Miocene of Dobrogea as well, also in Romania (Credinţa, MN 8) (Gál & Kessler 2006).

Systematic paleontology - 3

Garrulus Vieillot, 1816
Garrulus glandarius (Linnaeus, 1758)
Q1: Németóvár 4B (Deutsch-Altenburg, Austria) (Jánossy 1981), Beremend 17 (Q1) (Jánossy 1992) (Hungary); Betfia 2, 9 (Kormos 1913, Čapek 1917, Lambrecht 1933, Kessler 1975, Jánossy 1979, Gál 2002) (Romania); Q1-2: Betfia „Aven” (Kessler 1975, Jánossy 1979, Gál 2002), Kiskóh-Medvék Cave (Chişcău-Peştera Urşilor) (Kessler 1982) (all in Romania); Q3/I: Hundsheim (Mlíkovskỳ 1998, 2002) (Austria); Vindija (M. Malez 1961, V. Malez & Rukavina 1975, V. Malez 1973, 1986, 1988, Musil 1980) (Croatia); Q4/I: Mixnitz-Drachenhöhle (Lambrecht 1933) (Austria); Krapina, Velika Pecina, Veternica (V. Malez 1973, 1984, 1986, 1988, V. Malez-Bačić 1979) (all in Croatia); Bajót-Baits Cave, Bajót-Hóman Cave (Jánossy 1979), Budapest-Remetehegyi Cave (Kormos 1914, Lambrecht 1933, Jánossy 1979, 1986), Felsőtárkány-Peskő Cave (Lambrecht 1912a, 1912b, 1933, Jánossy 1979, 1986), Hámor-Puskaporos Cave (Lambrecht 1912a, 1912b, 1916, 1933, Jánossy 1979, 1986), Pilisszántói I. Cave (Lambrecht 1915, 1933, Jánossy 1979, 1986), Szilvásvárad-Istállóskői Cave (Lambrecht 1912a, 1912b, 1933, Jánossy 1952, 1955, 1979, 1986), Varbó-Lambrecht Kálmán-Cave (Jánossy 1964, 1979) (all in Hungary); Hidegszamos-Csont Cave (Someşul Rece) (Lambrecht 1915), Ohábaponor-Bordu Mare Cave (Ohaba Ponor-Peştera Bordu Mare) (Kessler 1985, Jurcsák & Kessler 1988, Gál 2002, 2003) (all in Romania); Q4/II: Balatonkeresztúr-Réti dűlő (Gál 2004, 2015), Ecsegfalva (Pike-Tay et al. 2004, Gál 2007), Felsőtárkány-Petényi Cave (Jánossy 1979), Legény Cave (Kormos 1914), Miskolc-Felső-forrás, Anonym Cave (Kessler 2010), Rezi (Kessler 2009), Tatabánya-alsó – Törekvés Cave (Kessler 2010), Vác-Széchenyi street, Visegrád-Várkert (Gál 2015) (all in Hungary); Remetelórév-Bólyikői Cave (Lorău-Peştera din Piatra Boiului) (Kessler 1982), Révi caves (Peşterile din Vadu Crișului) (Kessler 1982), Szegyestel-Drăcoiaia Cave (Sighiştel, Peştera Drăcoaia) (Kessler 1982); Székelykeresztúr (Cristuru-Secuiesc) (Gál 2008, 2015), Vársonkolyos-Kis Magyar Cave (Şuncuiuş, Peştera Napiştileu) (Kessler 1977, Gál 2002) (all in Romania). From sites in Europe outside the Carpathian Basin Q1-2: France, Germany; Q3: Czech Republic, France, Germany, Italy, Ukraine, United Kingdom; Q4: Austria, Belgium, Bosnia-Herzegovina, Bulgaria, Croatia, Czech Republic, France, Georgia, Germany, Greece, Ireland, Italy, Luxemburg, Moldova, Poland, Portugalia, Russia, Spain, Ukraine, United Kingdom (Tyrberg 1998).

Nucifraga Vieillot, 1816
Nucifraga caryocatactes Linnaeus, 1758 [sic] = Nucifraga caryocatactes (Linnaeus, 1758)
Q1: Betfia 9 (Gál 2002) (Romania); Q3: Vindija (M. Malez 1961, M. Malez & Rukavina 1979, V. Malez 1973, 1986, 1988, Musil 1980) (Croatia); Q4/I: Merkenstein (Wettstein & Mühlhofer 1938), Mixnitz-Drachenhöhle (Lambrecht 1933) (Austria); Bajót-Öregkő (Kormos 1914), Balla-Cave, Budapest-Remetehegyi Cave (Kormos 1914, Lambrecht 1933, Jánossy 1979, 1986), Felsőtárkány-Peskő Cave (Lambrecht 1912a, 1912b, 1933, Jánossy 1979, 1986), Galgóc (Lambrecht 1915, 1933), Hámor-Puskaporos Cave (Lambrecht 1912a, 1912b, 1916, 1933, Jánossy 1979, 1986), Pilisszántói I. Cave (Lambrecht 1915, 1933, Jánossy 1979, 1986) (all in Hungary); Barcarozsnyó (Peṣtera Gura Cheii-Râşnov) (Gál 1998, 2002), Szegyestel-Măgura Cave (Sighiştel, Peştera Măgura) (Kessler 1982, 1985, Gál 2002) (all in Romania); Detrekőszentmiklós-Pálffy Cave (Dzeráva Skála-Plavecky Mikulas) (Lambrecht 1913, 1933) (Slovakia); Q4/II: Teufelslucke (Soergel 1966) (Austria); Felsőtárkány-Petényi Cave (Jánossy 1979) (Hungary); Kazánszoros-Töröklik Cave (Cazanele Mari, Peştera Cuina Turcului) (Kessler 1974a Fischer & Stephan 1977), Révi caves (Peşterile din Vadu Crișului) (Kessler 1982), Szkerisoara-Coiba Mare Cave (Scărişoara, Peştera Coiba Mare) (Kessler 1982), Jurcsák & Kessler 1986, 1988), Vársonkolyos-Izbîndiş Cave, Vársonkolyos-Kis Magyar Cave (Şuncuiuş, Peştera Izbîndiş, Peştera Napiştileu) (Kessler 1977, Gál 2002) (all in Romania). From sites in Europe outside the Carpathian Basin
Q1-2: Spain; Q3: Czech Republic, France, Germany, Italy, Ukraine; Q4: Austria, Bulgaria, Croatia, Czech Republic, France, Georgia, Germany, Italy, Poland, Portugalia, Russia, Switzerland, Spain (Tyrberg 1998).

Pica (Linnaeus, 1758) [sic] = Pica Brisson, 1760
Pica pica † major Jánossy, 1979 (Table 2)
Site and era: MN 15: Beremend 26 (Kessler 2010), Csarnóta 2 (Jánossy 1979, Kessler 2010) (all in Hungary); Q1: Beremend 16, 17 (Jánossy 1992) (Hungary); Betfia 2, 9 (Kormos 1913, Čapek 1917, Lambrecht 1933, Kessler 1975, Jánossy 1979, Gál 2002) (all in Romania); Q2: Betfia „Aven” (Kessler 1975, Jánossy 1977, 1979, Gál 2002) (Romania); Q2: Nagyharsányhegy 1-4 (Lambrecht 1916, 1933, Jánossy 1979) (Hungary); Q3/I: Hundsheim (Lambrecht 1933, Jánossy 1979) (Austria); Dorog-Hungáriahegy (Jánossy 1953, 1986, Jánossy & Vörös 1987), Vértesszőlős 2 (Jánossy 1979, 1990) (all in Hungary). Mlikovskỳ (2002) classifies the fossil magpie species to the recent Western Jackdaw (Corvus monedula) based on the material from Stránská Skála classified by Jánossy (1972). When examining the fossilized material, we determined that this does not apply to the material from the Carpathian Basin, as the dimensional and morphological characteristics both refute this. Based on examination of the Csarnóta 2, Beremend 26 and numerous Lower Pleistocene materials, the validity of the fossilized subspecies is evident. The magpie characteristics, as well as the larger sizes than that of the present species, can be clearly shown. The present species is only known from the Middle Pleistocene of Europe, and is probably the direct descendant of the fossil subspecies. Mourer-Chauviré (1975) also describes the fossil subspecies (Pica pica major) from the Middle Pleistocene (Saint-Estéve Janson, Lunel Viel, Q3), while the present species is only known from the Upper Pleistocene. Another magpie find described to the genus level (Pica sp.) is known from the late Pliocene of Bulgaria (Văršec MN 17) (Mlikovskỳ 2002).

Pica pica (Linnaeus, 1758)
Q3/II: Vindija (M. Malez 1961, M. Malez & Rukavina 1975, V. Malez 1973, 1986, 1988, Musil 1980) (Croatia); Q4/I: Mixnitz-Drachenhöhle (Lambrecht 1933) (Austria);Bajót-Öregkő (Lambrecht 1914), Budapest-Remetehegyi Cave (Kormos 1914, Lambrecht 1933, Jánossy 1979, 1986), Cserépfalu-Subalyuk Cave (Jánossy 1979); Felsőtárkány-Peskő Cave (Lambrecht 1912a, 1912b, 1933, Jánossy 1979a, 1986), Hámor-Puskaporos Cave (Lambrecht 1912a, 1912b, 1916, 1933, Jánossy 1979, 1986), Kesztölc-Bivak Cave (Jánossy 1979), Pilisszántói I. Cave (Lambrecht 1915, 1933, Jánossy 1979, 1986), Répáshuta-Balla Cave (Lambrecht 1912a, 1912b, 1933), Szilvásvárad-Istállóskői Cave (Lambrecht 1912a, 1912b, 1933, Jánossy 1952, 1955, 1979, 1986), Tatabánya-Kálváriahegy Cave no. 4. (Gál 2004, 2005b), Varbó-Lambrecht Kálmán Cave (Jánossy 1964, 1979) (all in Hungary); Körösmart (Rîpa) (Jánossy in Hamar & Csák 1969, Kessler 1974b, Gál 2002), Ohábaponor-Bordu Mare Cave (Ohaba Ponor-Peştera Bordu Mare) (Kessler 1985, Jurcsák & Kessler 1988, Gál 2002, 2003) (all in Romania); Q4/II: Teufelslucke (Soergel 1966) (Austria); Budapest-Sas György square – Teleki Palace (Gál 2015), Ecsegfalva (Pike-Tay et al. 2004, Gál 2007), Felsőtárkány-Petényi Cave (Jánossy 1979), Ludas-Budzsák (Bökönyi 1974, Gál 2005a), Pilismarót-Malompatak (Jánossy 1985) (all in Hungary); Bégakalodva (Cladova) (Gál 2005a), Kazánszoros-Töröklik Cave (Cazanele Mari, Peştera Cuina Turcului) (Kessler 1974a, Fischer & Stephan 1977), Körösbánlaki Cave (Peştera din Bălnaca) (Kessler 1982), Remetelórév-Bólyikői Cave (Lorău-Peştera din Piatra Boiului) (Kessler 1982), Révi Cave (Peşterile din Vadu Crișului) (Kessler 1982), Vársonkolyosi caves (Peşterile din Şuncuiuş) (Kessler 1977, Gál 2002) (all in Romania). From sites in Europe outside the Carpathian Basin Q1-2: Spain; Q3: Azerbaijan, Croatia, Czech Republic, France, Germany, Italy, Spain, Ukraine; Q4: Austria, Belgium, Bosnia-Herzegovina, Croatia, Czech Republic, France, Georgia, Germany, Ireland, Italy, Montenegro, Moldova, Poland, Portugalia, Russia, Switzerland, Spain, Ukraine, United Kingdom (Tyrberg 1998).

Pyrrhocorax Vieillot, 1816
Pyrrhocorax graculus † vetus Kretzoi, 1962 (Table 3)
Site and era: MN 15: Beremend 26 (Kessler 2010), Csarnóta 2 (Jánossy 1972) (all in Hungary); MN 16: Villány 3 (Kessler 2010 as Pyrrhocorax pyrrhocorax) (Hungary); Q1: Beremend 17 (Jánossy 1991, 1992) (Hungary); Betfia 2, 9 (Kormos 1913, Čapek 1917, Lambrecht 1933, Kessler 1975, Jánossy 1979, Gál 2002) (Romania); Q1-2: Betfia „Aven” (Kessler 1975, Jánossy 1979, Gál 2002) (Romania); Q2: Betfia 5 (Kretzoi 1962, Kessler 1975, Jánossy 1979, Gál 2002), Kiskóh-Medvék-Cave 2 (Chişcău-Peştera Urşilor) (Kessler 1982, Jurcsák & Kessler 1988, Gál 2002) (all in Romania); Méhész (Vcelare) (Jánossy 1979) (Slovakia); Q3/I: Hundsheim (Lambrecht 1933, Jánossy 1979, Mlikovskỳ 1998, 2002) (Austria); Tarkő 3, 4 (Jánossy 1979) (Hungary); Gombaszög (Gombasek) (Kessler 2009) (Slovakia). Mlikovskỳ (2002) classified the subspecies with different sizes and especially ratios to those of the present alpine chough. This is countered by the facts that on the one hand, the differences indicated by the diagnosis are clearly visible, and on the other hand, its subsequent characteristics can be associated with much earlier materials (Csarnóta 2, Beremend 17, and numerous other Lower and Middle Pleistocene sites). All this supports our opinion that this is a fossil subspecies, as well as the direct ancestor, of the present species.

Pyrrhocorax graculus (Linnaeus, 1766)
Q3/II: Vindija (M. Malez 1961, V. Malez 1973, 1986, 1988, M. Malez & Rukavina 1979, Musil 1980) (Croatia); Q4/I: Merkenstein (Wettstein & Mühlhofer 1938), Mixnitz-Drachenhöhle (Lambrecht 1933) (all in Austria); Velika Pecina (M. Malez & Rukavina 1975, V. Malez 1984, 1988) (Croatia); Bajót-Öregkő (Kormos 1914), Bajót-Hóman Cave (Jánossy 1979), Budapest-Remetehegyi Cave (Kormos 1914, Lambrecht 1933, Jánossy 1979, 1986), Cserépfalu-Subalyuk Cave (Jánossy 1979); Csobánka-Kiskevélyi Cave (Lambrecht 1912a, 1912b, 1915, 1933, Jánossy 1979), Kesztölc-Bivak Cave (Jánossy 1979), Pilisszántói I. Cave (Lambrecht 1915, 1933, Jánossy 1979, 1986), Szilvásvárad-Istállóskői Cave (Lambrecht 1912a, 1912b, 1933, Jánossy 1952, 1955, 1979, 1986), Vaskapu Cave (Mottl 1941) (all in Hungary); Barcarozsnyó (Gura Cheii-Cave, Râşnov) (Gál 1998, 2002), Hidegszamos-Csont Cave (Peştera cu Oase, Someşul Rece) (Lambrecht 1915), Măgura-Valea Coacazei Cave (Măgura-Peştera din Valea Coacăzei) (Gál 2002), Nándor-Nándori Cave (Nandru-Peştera Curata) (Jánossy 1965, Fischer & Stephan 1977, Kessler 1985, Jurcsák & Kessler 1988, Gál 2002, 2003), Ohábaponor-Bordu Mare Cave (Ohaba Ponor-Peştera Bordu Mare) (Kessler 1985, Jurcsák & Kessler 1988, Gál 2002, 2003), Peterd-Tordai-hasadék – Binder Cave (Cheile Turzii-Peștera Binder) (Kessler 1985, Gál 2002) (all in Romania); Q4/II: Knochenhöhle (Bocheński & Tomek 1994), Grosse Offenbergerhöhle (Bocheński & Tomek 1994), Hohlensteinhöhle (Bocheński & Tomek 1994), Tropfsteinhöhle, Tunnelhöhle (Fladerer 1993) (all in Austria); Felsőtárkány-Petényi Cave (Jánossy 1979), Hosszúhegyi Cave (Jánossy 1979b) (all in Hungary); Herkulesfürdő-Rablók Cave (Băile Herculane, Peştera Hoţilor) (Kessler 1981, Gál 2002), Kazánszoros-Climente Cave (Cazanele Mari, Peştera Climente I) (Kessler 1981, Gál 2002), Kazánszoros-Töröklik Cave (Cazanele Mari, Peştera Cuina Turcului) (Kessler 1974a, Fischer & Stephan 1977) (all in Romania). From sites in Europe outside the Carpathian Basin Q1-2: Bulgaria, Spain, Ukraine; Q3: Azerbaijan, Czech Republic, France, Georgia, Germany, Spain, Ukraine; Q4: Austria, Bulgaria, Croatia, Czech Republic, France, Georgia, Germany, Greece, Italy, Montenegro, Poland, Russia, Switzerland, Spain, Ukraine, United Kingdom (Tyrberg 1998).
The present species is also known from the Late Pliocene of Bulgaria and Spain (Văršec and Meda Gran, MN 17) – these, however, supposedly belong to the fossil subspecies above – as well as from numerous sites in France and a few in Greece from the Lower and Middle Pleistocene. It can nowadays be found in the Alps, Pyrenees and the Balkan Peninsula (as well as the Upper Pleistocene and Holocene sediments of the caves located there) (Mlikovskỳ 2002).

Pyrrhocorax pyrrhocorax (Linnaeus, 1758)
Q2: Nagyharsányhegy 1-4 (Kessler 2010) (Hungary); Q3/I: Hundsheim (Mlikovskỳ 2009) (Austria); Q3/II: Vindija (M. Malez 1961, M. Malez & Rukavina 1975, V. Malez 1973, 1986, 1988, Musil 1980) (Croatia); Solymár-Ördöglyuk (Jánossy 1979) (Hungary); Q4/I: Luegloch (Mottl 1953) (Austria); Felsőtárkány-Peskő Cave (Lambrecht 1912a, 1912b, 1933, Jánossy 1979, 1986), Hámor-Puskaporos Cave (Lambrecht 1912a, 1912b, 1916, 1933, Jánossy 1979, 1986), Répáshuta-Balla Cave (Lambrecht 1912a, 1912b, 1933) (all in Hungary); Körösmart (Rîpa) (Jánossy in Hamar & Csák 1969, Kessler 1974b, Gál 2002), Rév-Pince Cave (Vadu Crişului, Peştera Pincelului) (Gál 2002) (all in Romania); Q4/II: Grosse Offenbergerhöhle (Bocheński & Tomek 1994) (Austria); Kazánszoros-Töröklik Cave (Cazanele Mari, Peştera Cuina Turcului) (Kessler 1974a, Fischer & Stephan 1977) (Romania). From sites in Europe outside the Carpathian Basin Q1-2: Spain, Ukraine; Q3: Azerbaijan, Czech Republic, France, Georgia, Spain, Ukraine; Q4: Austria, Bosnia-Herzegovina, Bulgaria, Croatia, France, Georgia, Greece, Italy, Luxemburg, Portugalia, Romania, Russia, Switzerland, Spain, Ukraine, United Kingdom (Tyrberg 1998).

Perisorius Bonaparte, 1831
Perisorius infaustus Bonaparte, 1831
Q4/I: Répáshuta-Balla-Cave (Lambrecht 1912, 1933) (Hungary).

Corvidae gen. et sp. foss. indet.
MN 15: Beremend 26 (Kessler 2010: as Nucifraga caryocatactes) (Hungary).

Corvidae gen. et sp. indet.
Q2: Ürömhegy (Jánossy 1961, 1986); Q4/I: Tatabánya-Kálvária no. 4. Cave (Gál 2004, 2005b) (all in Hungary).

Fred