Modelo de los Refugios Mediterráneos Estables Avian zoogeographical patterns during the Quaternary in the Mediterranean region and paleoclimatic interpretation.
Antonio Sánchez Marco.
Este artículo se publicó en el año 2004, en la revista Ardeola, 51 (1): 91-1
An overview of the avian geographical distribution patterns in the Mediterranean region during the Quaternary is presented. Seventy−two sites distributed along the region are analysed. Most of the sites bear avian assemblages rich enough to offer an insight on the main features of local paleornithocaenoses. The approach of the steady Mediterranean refugia is used to explain the avian geographical distribution drawn from the Mediterranean fossil records. As a consequence, we increase the accuracy of our knowledge on the climatic changes during this period.
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Se ofrece un estudio sintético sobre los patrones de distribución geográfica de las aves durante el Cuaternario en la región Mediterránea. A este fin, se ha hecho uso de 72 yacimientos repartidos por toda el área de estudio. La mayor parte de estos yacimientos poseen asociaciones fósiles con suficiente entidad para aportar datos fiables sobre las paleornitocenosis locales. Se propone un modelo explicativo para interpretar la distribución geográfica que parece indicar el registro fósil, al que se ha denominado Modelo de los refugios mediterráneos estables. Como consecuencia, se obtienen algunos rasgos de los cambios climáticos que han afectado a la región mediterránea durante este periodo.
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The scenario of a climatically unstable late Neogene is supported on several sets of evidences. The most precise among those may be the temperature changes recorded in the ice cores of the Antarctic and Greenland, as well as the δ O18/O16 record on foraminifera from the oceanic bottoms. Periglacial tilt formations and deposits of loess in medial latitudes indicate that there were phases in which the north of Eurasia and North America were covered in several occasions by an ice cap. During these stages (stadials), tempered species took refuge in the most southern zones of their respective areas of distribution, or even further south. At the same time, the fossil record demonstrates the blossoming of immigrant species, which are adapted to the colder conditions.
The shifts of the geographical distribution of taxa as well as biocaenoses are explained by two major theories: the dispersal theory (Simpson, 1965; Matthew, 1915) and the vicariance theory (Croizat et al., 1974). These theories have had different methodological developments, and are usually considered as alternative approaches. The Refugia theory does not contradict the isolation of populations secondary to the fragmentation of the territory, neither the dispersal of some taxa with the subsequent arrival into new areas.
The character of refuge assigned to the Mediterranean region (MR) during the Quaternary, specially during the last glacial phase (Würm), the role it played in the phylogenetic differentiation, as well as in the current west Palearctic distribution of birds, have been the subject of a large number of studies. Many of them are summarized in Moreau (1954, 1972), Blondel (1985, 1987), Tyrberg (1991a, 1991b) and Mourer-Chauviré (1993). Related to this, some species which are assumed to have different climatic and habitat singficances since they display at present disjunct geographic areas, appear together in the same levels of several Pleistocene deposits in southern Europe. This phenomenon has been referred in the literature as “mixed faunas”.
The Refugia theory, applied to the Quaternary, does not propose that all speciations have taken place in ecological refuges, but tries to explain many differentiation events which contributed to the development of the modern biotas (Haffer, 1982). The Mediterranean Refugia theory (MRT) (Moreau, 1954, 1972), particular derivation from this theory, has proved to be limited in its explanatory capacity. Moreover, an uncritical actualistic notion of the “indicator species” (a species that manifests by its presence the existence of certain environmental conditions) has overcome in fact the MRT, leading to misinterpretations. The Pleistocene avian assemblages from the south of Europe are, but few exceptions, characteristic of tempered climatic conditions, similar to the actual ones. Nevertheless, when the remains of currently northern breeding or resident species, as the Snowy owl (Nyctea scandiaca) or the Pine grosbeak (Pinicola enucleator), are found in outcrops of the south of the continent, even in fossil assemblages of Mediterranean type, it continues to be interpreted as a proof of regional cool conditions in the past. Thus, the notion of “indicator species”, with the meaning of “local or regional conditions indicator” leads to misinterpretate the paleoecological and paleoclimatical implications of the avian fossil record (Sánchez Marco, 1996). In this way, the indicator value of individual taxa prevails in the literature on the ornithological community character. The increasing number of studies of European Pleistocene avian localities modifies our understanding of this period. Thus, this paper proposes an alternative interpretation: the approach of the steady Mediterranean refugia (ASMR).
The ASMR was outlined in Sánchez Marco (1996). It leans on avian community features and, as a second step, in the individual species composition. Thus, the ASMR emphasizes the necessity of using analytical and comparative approaches to paleornithological assemblages with current ornithocaenoses (Sánchez Marco, 1999a, 1999b). Therefore, there are several ecological patterns providing the guide to find out as much the paleoclimatic characteristics as the configuration of the landscape in the past. Some of these patterns have been studied in mammals and birds, as the habitat spectra of the community (Fleming, 1973; Evans et al., 1981; Sánchez Marco, 1999a, 1999b). The phenetic behaviour spectra has been used by Sánchez Marco (1999b). The ASMR attempts to apply in concrete terms the Refugia theory to the distribution of birds in the Mediterranean region during the Quaternary. It relies on the notion and description of the role of “Northern irruptive species” (see the paragraph “Material and methods”). Its main difference with the MRT (Moreau, 1954, 1972) consists of comprising the occurrence of Mediterranean avifaunas during the whole Quaternary in the south of the Mediterranean peninsulas, even in the cool phases.
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2. GEOGRAPHIC AND CLIMATIC CONDITIONS IN THE QUATERNARY
The MR is the contact area between two of the largest continental plates: the Eurosiberian and the African ones. This condition has led to very complex geological and paleogeographical histories (Maldonado, 1985). Tectonic movements of both plates in the early Oligocene drove to a closed sea in this area. Subsequent to the desiccation of great extensions of the Mediterranean sea at the fall of the Miocene, the Messinian episode –from 5.7 to 5.3 Ma (see Gautier et al., 1994)-, the opening of the strait of Gibraltar decisively transformed the regional geography, determining important biogeographical changes in mammal faunas from both the continent and the islands (Moyà-Solà et al., 1999; Alcover, 2000; Azanza et al., 2000). Two other major biotic events, occurred at the end of the Neogene, affected the mammal faunas in the western MR: the Equus-elephant event (Early Villafranchian, MN 16, c. 2.7 Ma) and the Galerian mammal pulse (1.0 Ma) (Azanza et al., 2000). These events are in accordance with two of the three major shifts to more arid and open conditions, occurred in Africa around 2.8 Ma, 1.7 Ma and 1.0 Ma and which are recorded in marine eolian deposits (DeMenocal, 1995). The Mediterranean sea reaches a considerable extension and its influence on the climate constitutes one of the main features of this region, since the lands around it show more or less homogeneous climatic conditions. Faunistic successions in the western Mediterranean islands as well as mass extinctions episodes are the object of some works, such as Moyà-Solà et al. (1999), Alcover (2000) and Seguí & Alcover (1999).
A critical date in the recent history of the global climate is 2,7 Ma. This cooling episode in the Arctic hemisphere had begun towards 3.1 Ma (Raymo et al., 1989; Raymo, 1991), causing the growth of the polar cap. Previously, the temperate periods were warmer than the present one and the cold periods were warmer than some later warm interstages. Until that date, it is likely that the continents of the Northern hemisphere were free of considerable masses of ice. Coinciding with the beginning of the Matuyama, aproximately 2.4 Ma ago, the first event of massive formation of icebergs in the North Atlantic is well documented. Important amounts of materials transported by icebergs were deposited in the Atlantic ocean (Shackleton et al., 1984; Raymo et al., 1989). It was the beginning of the first Cenozoic glaciation in the Northern hemisphere.
The marine sediments 2.7 Ma old document accused drops of the oceanic temperatures (Raymo, 1992; Raymo et al., 1989). Since then, the oscillations between tempered and cold stages have acquired greater amplitude keeping a tendency towards a major cooling. From approximately 1.5 Ma, the thermal oscillations point to a rate of increasing amplification, exceeding themselves the previous extremes as much mild as cold. Comparing the values of δ 18O in oceanic sediments of the present interglacial stage with those of the last one million and a half of years, only in six occasions were reached such high temperatures. This happens in interstadials 43 (1.2 to 1.3 Ma), 35 (~1.1 Ma), 31 (0.9 to 1 Ma), 11 (~0.4 Ma), 9 (~0.3 Ma) and 5c (~0.13 Ma). In fact, the Holocene temperatures are surpassed in the four last ones. It is interesting to notice that most interglacials did not reach as high thermal values as in recent times.
In the other extreme of the oscillations, also the lowest temperatures of the cold episodes take place in the end of the Pleistocene. They are concentrated from approximately 0.5 Ma, and are episodes 16 (< 0.6 Ma), 12 (~0.4 Ma), 6 (~0.15 Ma) and 2 (0.02 Ma). A series of datings by the U/Th method in marine sediment grants to the Eemian interglacial a maximum span of 10 ka, and between 1 and 2 ka to the substage 5e (Slowey et al., 1996). The fast and short-duration climatic changes happened during the last interglacial and in the base of the Weichselean, as showed from the oxygen isotopes analysis of ice cores from Greenland, were interpreted as “artefacts” owing to changes of state at the deepest zone of the ice cap (Adkins et al., 1997). Although these climatic changes could be coincident with the above mentioned observations on the faunal changes of rodents in Central European deposits (Horáček & Ložek, 1988).
During the Würm period, the highest peaks in the Iberian peninsula were covered by glacial sheets (Uchupi, 1988). The loess accumulations stretched out from west of France to the east, along the edge of the southern end of the last European icecap (Horáček & Ložek, 1988). Currents of polar waters descended up to a latitude of 42º N during the Quaternary cool peaks, and the Gulf stream was turned aside towards the southwest. The Iberian peninsula was exposed to predominant winds from the west (Uchupi, 1988). Warmer temperatures giving rise to the present warm interstadial are noted in high latitudes 18 ka BP.
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The scope of this paper covers the Quaternary, although the lowest limit considered here is not the generally accepted 1.6 – 1.8 Ma (Aguirre & Pasini, 1985). The reason for this consideration is that this limit –as well as the divisions of this period- has been established on the shifts of mammal distributions (mainly in Europe) (for instance, Mein, 1976; Fahlbusch, 1976; Bruijn et al., 1992; Calvo et al., 1993), and these ones do not correspond to the changes of avian paleogeography. In this regard, Higueruelas (ca. 3 Ma) is the oldest locality in the MR with an association of avian taxa similar to the ones found during the Lower Pleistocene: the small passeriforms constitute an important fraction of the species assemblage; from 26 species present in Higueruelas, only one (new) species of Palaeocryptonyx seems not to go through the conventional limit of the Pleistocene; and it appears a Corvus of the pliocaenus/antecorax group (a typical corvid or corvids of the European Lower and Middle Pleistocene). And the uppermost limit chosen for the present work is the Epipaleolithic (after the last main cool pulse).
For this study I have analysed seventy-two fossil sites located in the current MR and in its peripheral fringe. The African side of this region is still a vacuum territory for the paleornithology of the Quaternary. When it has been possible, I have choosen fossil localities, geological layers or sedimentary complexes providing rich avian assemblages, so that they can approach to a certain extent the paleoclimatic and paleoenvironmental conditions. This comprises a total amount of 89 avian assemblages (Appendix 1). Some of the fossil associations used in this work can be excluded as samples of paleornithocaenoses, since constitute clear cases of associations containing the recent introduction in Europe of Gallus (Bacho Kiro, Fontéchevade, Ibex), where the mixture of layers is doubtless. Despite this, such localities have been chosen due to the scarcity of fossil assemblages in some areas or to cover some time spans. They are only used to illustrate de presence of individual species. The data have been taken directly from the original works to avoid possible mistakes, with the exception of one or two cases which are pointed out. Some publications have not been considered for this paper since they contain obvious mistakes. All the authors and publications do not merit the same degree of confidence, however, the aim of this paper is not to do a review of the published material. Some of the taxonomical identifications and the chronological attributions should be re-examined.
The geographical situation of the localities are given in the map of figure 1. The sites have been listed in two chronological tables (Tab. 1 and 2), although their respective positions are approximative for many of them.
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| Figure 1 – Map of a part of the Mediterranean region with the fossil localities used in this work. Current shorelines. Abbreviations: A – Arbreda, A-T – Ambrona and Torralba, A1 – Áridos 1, AC – Arene Candide, Ac – Acquedolci, Ak – Akrotiri, At – Atapuerca complex (Elefante, Dolina, Galería), BB – Bois-des-Brousses, Bi – Binigaus, BK – Bacho Kiro, C1 – Casablanca 1, C3 – Castiglione 3, Ca – Carnello, CB – Cau d’en Borrás, Ce – Cendres, CF – Contrada Fusco, CN – Cova Nova, Co – Coscia, CR – Ca na Reia, Ct – Canet, CV – Cingle Vermell, DT – Devil’s Tower, Du – Dursunlu, EP – Es Pouàs, F – La Fage, FB – Figueira Brava, Fn – Fontéchevade, Fo – Fontbrégoua, Fu – Fumane, G – Gumbes (B, C), Ge – Gegant, Go – Gorham, H1 – Huéscar 1, Ha – Hayonim, Hi – Higueruelas, Ho – Hortus, Ib – Ibex, J2 – Jarama II, K22 – K22, Kk – Kozarnika, La – Lazaret, Li – Liko, M5 – Montoussé 5, MA – Mas-d’Azil, N – Nerja, O2 – Ohalo 2, O3 – Orgnac 3, P – Palidoro, PO – Pedrera de S’Ònix, Pu – Punta di Calcina, Q – Quibas, Qa – Quartaccio, R – Radice, Rm – Romanelli, Ro – Romaní, Rs – Romains, Rz – Razhishkata, S – Salpêtre, Sa – Salpêtrière, Sj – Šandalja (I, II), Sp – Spinagallo, Te – Temnata, TN – Torre Nave, TP – Torre in Pietra, Tr – Trebački, Ub – Ubeidiya, Va – Varshets, Vb – Valdegoba, Vi – Victoria, Vn – Vindija. |
3.1. NO-MEDITERRANEAN IRRUPTIVE SPECIES
Some high dispersive species, which are nowadays distributed along northern Eurasia are recorded in Quaternary localities of MR. A case repeatedly noted in the literature is that of the Snowy owl (Nyctea scandiaca) (Lambrecht, 1933; Brodkorb, 1971; Boev, 1998a). According to their respective current distributions, such species are considered herein as irruptive in the whole or a part of the MR during the Quaternary (Appendix 2). In Appendix 1 each irruptive species are enlightened when the fossil locality falls in the corresponding “area of irruption” of that species (see Appendix 2). The current ranges of the Pygmy owl (Glaucidium passerinum) and Tengmalm’s owl (Aegolius funereus) cover approximately the north border of the MR. When these species appear in localities near the limits of the MR, they are not considered as irruptive into it, but as punctuating the spreading of Eurosiberian climatic conditions.
3.2. NO-MEDITERRANEAN LOW-DISPERSIVE SPECIES
The grouses (Lagopus, Tetrao, Lyrurus, Bonasa) are a low-dispersive group of birds whose respective geographical distributions are fairly outside the MR (Cramp, 1998). In this paper it is assumed that those species were not included in the past components of Mediterranean paleornithocaenoses neither. Thus, their presence in Mediterranean localities may be understood as: (a) the fossil association was deposited under no-Mediterranean conditions or (b) it is a mixture of paleornithocaenoses from periods with different climatic conditions. These species are underlined in Appendix 1.
3.3. EAST MEDITERRANEAN SURVIVING SPECIES
Appendix 3 lists the species which are nowadays distributed along the east part of the MR, but that are also recorded in fossil localities from the west part. In Appendix 1 these taxa are enlightened when appear in localities from the west area of the MR.
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4. SURVEY OF THE FOSSIL LOCALITIES
4.1. LAST PLIOCENE AND EARLY PLEISTOCENE
The Iberian locality of Higueruelas (MN 16) (Bruijn et al., 1992) has been dated between 3.18 and 3.46 Ma (Bonadonna & Villa,1984) and to 3.2 Ma (Aguirre & Morales, 1990). The presence of several shallow lakes, combinated with regional volcanism, provided suitable environments for the preservation of a large sample of the local paleornithocaenoses. In the species assemblage yielded by this outcrop, the larger ecological group was constituted by aquatic birds: Podiceps auritus, P. nigricollis, Cygnus cygnus, Anser sp., Tadorna sp., Aythya marila, Oxyura leucocephala and Mergus albellus, as well as for those species also linked to aquatic environments: Ardea cinerea, Nycticorax nycticorax, Ixobrychus minutus, Plegadis falcinellus, Actitis hypoleucos and Anthus pratensis.
The finding of C. cygnus, A. marila and M. albellus constitute an unexpected event concerning the age and geographic situation documented by this locality. At present, they are breeding species in northern zones of Eurasia, and they are rare in Iberian territory. In the case of the swan, this one only appears with occasion of hard winters (Díaz et al., 1996). Contingents of these three species move during the winter, not only to Central Europe and north of France, but also to more southern latitudes, like the Balkan peninsula and Anatolia.
Montoussé 5, dated to the Upper Pliocene, in the northern side of the Pyrenees, has yielded a small quantity of bird remains, including Corvus pliocaenus and Palaeocryptonyx sp. (Clot et al., 1976). The fauna is characteristic of temperate conditions and of forest environments. Each one of these mentioned taxa appear in two respective sites in Mallorca (Balearic islands), Palaeocryptonyx sp. in Ca na Reia (Upper Pliocene), together with a small quantity of other taxa (Alcover, 1989), and C. pliocaenus in Pedrera de S’Ònix (Plio-Pleistocene) (Mourer-Chauviré et al., 1977). Moreover, this locality situated near Manacor town has yielded a relatively large collection of birds (Mourer-Chauviré et al., 1977; Mourer-Chauviré in Alcover et al., 1981), most of them from woodland habitats, and two irruptive taxa Cygnus cf. cygnus and Bucephala cf. clangula. Further taxonomic revision of this fauna by Seguí (1996) has not changed its paleoenvironmental implications.
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| Table 1 – Late Pliocene and Pleistocene localities from the Mediterranean region used in this work. 1.- Chronology (x 106 aBP). 2.- Magnetostratigraphy. 3.- Epochs. 4.- Lithic intervals. 5.- Major divisions after micromammals. [Ampliación de la tabla] |
Although it is partially outside of the chronological framework of this paper, it is worth while to mention the Tyto balearica, a paleospecies of barn owl widely distributed in the West Mediterranean area throughout a notable span of time, which was classified on fossil material from three Balearic localities (Canet, Pedrera de S’Ònix and Binigaus) dated of the Plio-Pleistocene boundary (Mourer-Chauviré et al., 1980). This species was later identified in some older sites in France and Spain: Layna (Early Pliocene, MN 15), Sète (Early Pliocene, MN 15), Balaruc II (Late Pliocene, MN 16), Casablanca 1 (Late Pliocene, MN 17) (Mourer-Chauviré & Sánchez Marco, 1988; Mourer-Chauviré, 1995; Sánchez Marco, 1995a), Aljezar B (Late Miocene, MN 12) (Cheneval & Adrover, 1993), Valdecebro 5 (Late Miocene, MN 12), Moreda (Late Pliocene, MN 16) (Sánchez Marco, 2001). This owl has been also identified in two Middle Pleistocene sites: Castiglione 3 CG (Mourer-Chauviré in Salotti et al., 1997, Mourer-Chauviré et al., 1997) and probably Punta di Calcina (Corsica) (Pereira et al., 2001). Moreover, remains from the complex of outcrops of Gargano (Early Pliocene, Southern Italy) have been attributed to this species (Mlíkovský, 1998).
Varshets fossil locality is a ponor situated near the homonymous town, in north-west of Bulgaria. It has been assigned to the Late Pliocene, MN 17 (Boev, 2002). Its very rich association of birds has lead to a large bibliographic production (Boev 1995, 1997, 1998b, 1998c, 1999a, 1999b, 1999c, 1999d, 1999e, 2000a), summarized in Boev (2002). The specific composition of this fauna is different enough from the Pleistocene ones to establish confident comparisons. There has not been found Geronticus nor Tyto balearica nor Palaeocryptonyx. The presence of Lagopus and Tetrao in the Balkan peninsula will be constant along the whole Quaternary.
Located in the north of the Dead sea rift (Israel), and near the Tiberias or Kinneret lake, there is the site of Ubeidiya, composed of more than sixty archaeological horizons of an age of c. 1.4 Ma. This lower Pleistocene site reveals a complex geological structure in which tectonic movements have tilted the deposits. The cultural remains are assigned to the Acheulian tradition (Bar Yosef & Goren-Inbar, 1993). As evidenced by the abundant aquatic species (Tchernov fide Tyrberg, 1998), the outcrop was situated close to an area of marshes and bodies of water.
Quibas site has an age between 1.0 and 1.3 Ma after the rodents association. No avian wintering or northern species have been found in this outcrop. The group of taxa, where most of them are related with masses of water and parkland environments, points to moister climatic conditions than present ones (Montoya et al., 1999, 2001).
The early Pleistocene site of Victoria is located on the southern Iberian coast. Some remains of swan (Cygnus cf. olor) also appear in this outcrop, which should be considered as a wintering species. Prunella modularis, is currently a resident passeriform in mountain areas in the half north of the Iberian peninsula, but during the winter this bird makes in the whole of its area of distribution short displacements towards the south or to lower levels. Emberiza melanocephala does not live today in the western zone of the Mediterranean. It breeds in the Balkan region and in Anatolia.
The rich deposit of Dursunlu is placed in the eastern extreme of the MR, in the Konya basin, near Llgin (southern Anatolia), which is attributed an age between 0,9 and 1 Ma (Guleç et al., 1998). Louchart et al. (1998) offer a list of 50 species, the immense majority of which is of aquatic habitats. The set of taxa of resident and breeding species corresponds to tempered climatic conditions, with the presence of numerous wintering birds.
The study of Elefante site is still incomplete. This locality belongs to the sierra de Atapuerca karstic infillings complex, situated in the northern plateau of the Iberian peninsula. The lower layers contain a rodent assemblage from ca. 1 Ma. Bones from Corvus antecorax and Haliaeetus albicilla are the most abundant ones (Rosas et al., 2001). There are no fossil remains fairly attributable to wintering species.
There are other rich fossil localities in the Iberian peninsula in chronological range between 1 My and the beginning of the Middle Pleistocene. Huéscar 1 is a fossil locality situated in a present-day arid zone in the southern Iberian inland. The avian taxa are, with little exceptions, aquatic ones, most of them, anatids (Sánchez Marco, 1989). Two of the species that appears here, Melanitta nigra and Mergus serrator, currently are not inland wintering species (Díaz et al., 1996). This behaviour is associated to periods of storms and rough weather in the coasts.
The Lower Pleistocene Dolina outcrop belongs also to the Atapuerca complex. The layer 6 (TD-6) is older than 0.8 Ma and has yielded one of the richest avian assemblages in Iberia, where open country species constitute the best represented group, followed by bushland biotope and inland water birds (Sánchez Marco, 1999b). On the other hand, resident species reach the highest values, especially those residents in the modern Eurosiberian zone of the Iberian peninsula. Wintering and breeding taxa have a comparatively low incidence. The occurrence in TD 6 of such taxa as Anas, Melanitta, Porzana and Cinclus provide evidence for the presence of a body of water nearby, larger than the modern-day Arlanzón brook that runs about 3 km from the outcrops. In conclusion, the avian remains from this layer point to three main ecological traits: (a) the existence of a pool, lake or slow river in (b) an open country type of habitat, and (c) that climatic conditions were similar to modern Iberian winters or that most fossils were deposited in the cave during the winter season (Sánchez Marco, 1999b).
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The site Orgnac 3, at Orgnac-L’Aven (France), was constituted by the fulfilling of a doline. The stratigraphic profile shows 19 layers, among them the layer i is the richest in avian remains. By this reason as well as for the occurrence of N. scandiaca, the fossil assemblage from this unit has been chosen for the present study. The avian bones were studied by Mourer-Chauviré (1975a). After the rodent association, Jeannet (1974 fide Mourer-Chauviré, 1975a) attributed the layer i to the Mindel-Riss phase. An important fraction of the species are tree-dwelling birds, and it is also remarkable the abundance of galliforms. The rodents should indicate a temperate and dry climate (Chaline, 1972, fide Mourer-Chauviré, 1975a).
The Aven 1 of La Fage is a karstic infilling situated near Noailles town (France). It has yielded one of the largest list of species in the Middle Pleistocene, with more than 100 taxa (Mourer-Chauviré, 1975b; Mourer-Chauviré et al., 2003) distributed along the whole stratigraphy (Mourer-Chauviré & Philippe, 1972). Avian remains are more abundant in the stratigraphic unit CO (Mourer-Chauviré, 1975a), a subdivision of the layer 5. The climatic conditions for this layer have been interpreted (Mourer-Chauviré, 1975b; Mourer-Chauviré et al., 2003) as being slightly colder than today, but with colder and warmer periods, because terrestrial gasteropods, micro- and macromammals as well as some birds from this layer would indicate temperate climatic conditions, and, on the other hand, some other bird species are tied to cold climates.
Ambrona and Torralba are two localities 3 km apart, both related to acheulean occupations (Santonja, 1989; Santonja & Vila, 1990). The ten avian species found here are aquatic ones (Sánchez Marco, 1990, 1999c). With the exception of two rallids (Porphyrio porphyrio and Fulica cf. atra), the remaining species were identified as wintering species. Also attributable to this age, and located in the middle of Iberia is Áridos 1, an anthropic deposit formed in one bank of the Jarama river. The best ecological group represented here is the one tied to woodlands and forests. Likewise, there are aquatic and open country species (Mourer-Chauviré, 1980).
Castiglione 3 is a karstic cavity in the Oletta massif (north of Corse). The deposit Castiglione 3 CG is a fissure that contains a large quantity of fossil remains. It has been attributed to an age of 350 ka (Salotti et al., 1997). The identification of the birds is owed to Mourer-Chauviré (in Salotti et al., 1997). This Middle Pleistocene site records one of the last occurrences of T. balearica, which appears with T. alba. The major part of the collection is composed by diurnal and nocturnal raptors, with two insular paleoendemisms.
Fontéchevade cave is in the middle west of France, near Montbron town. The fossil assemblage corresponds to a woodland habitat type (Mourer-Chauviré, 1975a). It contains characteristic Mediterranean avian taxa as well as no-Mediterranean ones, and one or two species which occurrence could be interpreted as the result of irruptions from north Europe.
The site known as Galería also belongs to the Atapuerca complex. The three stratipraphic units containing the highest diversity of avian species are, from top to bottom: TG11, TG10A-TN7 and TG10B-TN6 (Sánchez Marco, 1995b, 1999a). These units are younger than an inmmediately below stalagmitic crust dated to 317.6 ± 60 ka (ESR, Grün & Aguirre, 1987), and older than the crust inmmediately above dated to 211 ± 32 ka (ESR, Falguères, 1986) and 177.3 ± 23 ka (ESR, Grün & Aguirre, 1987). The filling of this cavity does not coincide chronologically with the ones of Dolina and Elefante (Made et al., 2003).
The bird remains found in the TG10B-TN6 and TG10A-TN7 units, compose habitat and phenetic spectra which are charateristic of humid zones with open lands. They correspond to zones in which the fauna acquires its greater diversity in the wintering periods. Some considerable forest masses might have existed, as it can be deduced from the passeriforms related to landscapes with shrubs and trees. There are no evidences to afirm that these areas, with more or less open forests, constituted one of the main elements of the surroundings. The habitat and phenetic spectra of the superior stratigraphic unit, TG11, points to reductions of the temperature and the humidity for that period, in comparison to the underlying layers conditions. During the corresponding time interval the bodies of water continued to determine the local communities of birds and it seems that a regression of the forests took place. One trait of the landscape recorded in the Early and Middle Pleistocene of Atapuerca could be the existence of a humid zone that extended at the feet of the mountain range of Atapuerca. This humid zone was constituted by backwaters (pools or fluvial sections of slow waters) and streams. It changed in several occasions in extension, and probably in depth. It seems that this region was a wintering area, very important in the periods corresponding to lower layers of Elefante and upper ones of Galería (Made et al., 2003).
The Lazaret site is a 40 m long cavity, located in the Boron mountain, near of Nice and 100 m from the current coastline. During the Middle Pleistocene its distance to the sea was around 500 m (Vilette, 1993). The stratigraphic unit C.III is observed in the profile at the entrance of the cave. These sediments were deposited before the formation of a stalagmitic crust dated between 125 and 70 ka BP (Falguères et al., 1992). Thus, Lazaret C.III has been attributed to an age of 150 to 125 ka BP, OIS 6. All the species showed in table 1, except Cinclus cinclus, appears in the first layers of this stratigraphic unit (Vilette, 1993: table XII). The avian assemblage corresponds to a forest area with inland humid zones. The nearness of the sea is also noted in the fossil remains of birds. Some years before the work of the unit C.III, Mourer-Chaviré (1964, 1975a) studied two other units of this cave: Locus VIII and some layers from the bottom of the cavity. Both infillings (Lazaret Locus VIII and Lazaret bottom) are considered to be deposited during a large span of time of the Riss interval (Mourer-Chauviré, 1975a). Thus, none of both avian associations may be understood as samples of individual avifaunas, but as a mixture of different ones. In spite of which, their taxonomical compositions and their respective representation of habitats are very similar to those of the unit C.III, although the latter encompasses a shorter interval. These three associations include typical Eurosiberian and Mediterranean taxa, particularly the corresponding to Locus VIII and C.III. A sharp difference of this one is the appearance of several northern irruptive species as well as H. albicilla and C. macrourus, today restricted to more eastern areas.
Therefore, either the southern coast of France suffered fluctuating climatic conditions during the cool phases of this period or there was a strip alongside of the coast with a real “mixed” ornithofauna with no parallel today.
Approximately in the middle of the Italian peninsula, Quartaccio quarry (Vitinia, Rome), records an assemblage from the Vitinia Formation (Bedetti, 2001). The corresponding avian list of the Appendix 1 is the result of a revision by Bedetti (2001) of previous works. Most of the species correspond to aquatic-dwelling birds. The occurrence of the Eider (Somateria molissima) should be considered has an irruptive species into this region.
The avian remains from Spinagallo cave, near Siracusa (southeastern Sicily), were studied by Pavia (1999). It contains a fossil association dated of about 500 ka BP (Bada et al., 1991), and composed of a large list of Mediterranean taxa. The degree of isolation during the period represented by this assemblage seems to have been quite reduced.
Among the five Pleistocene faunal complexes described in Sicilian localities, one of them, Elephas mnaidriensis Faunal Complex, from the Middle Pleistocene, has yielded an important avian assemblage (Pavia, 2001). Three outcrops contribute to the avian collection: K 22, Acquedolci and Contrada Fusco –the former studied by Cassoli & Tagliacozzo (1996)-. The assemblage of birds shows the typical traits of the Mediterranean insular avifaunas (sensu Alcover et al., 1992) but revealing a reduction of the degree of isolation (Pavia, 2001). The Goosander (Mergus merganser) seems to be a northern irruptive species.
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Torre in Pietra locality is in the province of Rome and was studied by Cossoli (1978). The layer m, with a pre-würmian lithic industry, has a Mediterranean assemblage of birds, with a high proportion of water species.
In the north of Spain, in the area today included in the Eurosiberian region, Valdegoba cave, whose sediments deposited during the Mousterian period (Díez et al., 1989), offers an avian association with Mediterranean as well as Eurosiberian characteristic species. Three northern irruptive species are recorded. Two more reduced assemblages, but with similar features to Valdegoba are the Italian localities of Radice valley (Biduttu et al., 1967) and Carnello (Segre et al., 1984), both situated near Sora village and with some Mousterian tools collections.
The Upper Pleistocene deposits from Cova Nova locality (Capdepera, Mallorca island) have yielded a large sample of a Mediterranean insular avifauna (Florit & Alcover, 1987; McMinn & Alcover, 1992). The occurrence of the Bullfinch (Pyrrhula pyrrhula) in the Balearic islands has to be considered as a case of irruption of northern species. Other typical Mediterranean assemblage was collected at the locality of Es Pouàs (Florit et al., 1989). This karstic cavity, where there are no remains of irruptive taxa, is situated near Santa Agnès de Corona (Eivissa island).
 |
| Table 2 – Upper Pleistocene and Holocene localities from the Mediterranean region used in this work. 1.- Chronology (x 103 aBP). 2.- Epochs. 3.- Lithic intervals. 4.- Lithic types. 5.- Oxygen Isotopic Stadials δ 18O. [Ampliación de la tabla] |
The fossil remains from Bacho Kiro cave were studied by Bocheński (1982). This locality from the Stara Planina region (Bulgaria), show an assemblage characteristic of temperate to mild conditions. The presence of Lagopus mutus could lead to assume some relatively cool temperatures, although this species seems to have occurred in forest zones in Late Pleistocene (Bocheński, 1974).
Cau d’en Borrás is a cavity on the east coast of Spain, near Oropessa town as well as the current sea line, in spite of which the only species recorded connected with the sea is Haliaeetus albicilla. The fossil assemblage is characteristic of open Mediterranean habitats. It is important to point out that the closeness of the sea do not seem to have any consequence in the avian fossil record at the locality.
Gibraltar has a big number of karstic cavities, a lot of them with an valuable Pleistocene record. Gorham’s cave is situated close to the south end of this small peninsula. The fossil remains from this site were studied by Eastham (1968). Two avian assemblages are the most rich: one from layers B and D, attributed to Würm III (ca. 29 ka BP) an another from layers K and U, attributed to Würm I. The former is an unbalanced sample of the corresponding local avifauna. Most of the taxa are marine species, raptors and rock inhabitants. The assemblage from layers K and M is also of Mediterranean type, but records the presence of some northern irruptive birds.
Devil’s Tower shelter was situated –it was dismantled some time ago- near the isthmus of Gibraltar. Its faunal remains were identified by Bate (1928). The avian association is very similar to the ones mentioned for Gorham’s cave. Not far from here, between both localities and about 250-300 masl, there is Ibex cave, whose fossil association, studied by Cooper (2000) represents also a Mediterranean ornithofauna. The dates of Ibex cave (Rhodes et al., 2000) support a Late Pleistocene age for this cavity but later than Gorham.
With the exception of Pinguinus impennis and northern irruptive species, the taxa recorded in the Gibraltarian localities are the same ones that they are observed currently at this migratory pass (Finlayson, 1992).
Gegant cave is situated on the coast of Barcelone, in the Garraf massif. The assemblage of the layer II indicates, as in other cases, open areas under climatic conditions typical for the Mediterranean region.
Arbreda cave is near the Serinyà village, in a zone of the north of the Iberian peninsula with intense processes of travertine formation and karstification (Garcia, 1995). The avian finds of this outcrop have been studied by Vilette (1983) and Garcia (1995, 1997). The largest fossil assemblages correspond to the stratigraphic units: Mousterian, Aurignacian (dated to 25830 ± 400 aBP) and Gravettian (dated to 20130 ± 220 aBP) layers by Garcia (1995) and Aurignacian layers by Vilette (1983). All the assemblages are typical of the current Mediterranean region. In the Gravettian association, Garcia (1995) attributed one of the finds to Lagopus sp., but this fact does not determine the meaning of the whole avian association. It is important the appearance of Pinicola enucleator in the Upper Paleolithic layers of Arbreda II (Vilette, 1983).
Romaní site is a rockshelter built in a travertine formation and located just in Capellades village, north-east Iberia. The stratigraphic unit II has been dated between 40-44 ka BP (Bischoff et al., 1988). It contains a small sample of the corresponding ornithofauna. A noticeable feature is the occurrence of Pinicola enucleator.
In the Languedoc region (south of France), the Mousterian layers from Hortus cave have provided a collection of fossil birds studied by Mourer-Chauviré (1972, 1975a). It is a typical Mediterranean assemblage with no record of characteristic Eurosiberian species. Ten kilometres from Hortus locality it is Salpêtre cave (Mourer-Chauviré, 1975a), likewise containing Mousterian and Upper Paleolithic layers. But the avian assemblage from the Mousterian layers from the latter show the occurrence of Mediterranean and Eurosiberian species, fact already seen in other localities from this area. The same features can be observed in layers 1a and F1 (transition Dryas 3 – Preboreal) from Salpêtre cave, studied by Vilette et al. (1983).
Coscia localities are situated in the north end of Corse island. Two fossil avian deposits are distinguished (Bonifay et al., 1998): Coscia (rock shelter South), from the early Würm, and Coscia cave, ca. 60 ka BP. Both avian assemblages are typical of western Mediterranean islands. In the shelter South record there is evidence of irruption of northern species.
Cassoli (in Bulgarelli, 1972) identified the fossil birds from Torre Nave cave. This site is situated on the southern coast of the Italian peninsula, in Cosenza. Unexpectedly, there are not any marine species in the avian assemblage. Nor northern irruptive species are.
Near the village of Remoulins, not far from Nîmes (southern France), is Salpêtrière shelter. The stratigraphic complex CG5-SLC4 (corresponding to layer 14a) has yielded the richest avian association of this site. It has been attributed to the early Aurignacian and dated of 28180 ± 1000 aBP (Vilette, 1983). The presence of aquatic species is explained by the closeness of the Gordon river, but the record of low dispersive Eurosiberian species as well as northern irruptive birds, together with the absence of typical Mediterranean taxa points to Eurosiberian paleoenvironmental conditions.
Fumane rock shelter is located in the region of Veneto, north Italy, close to the Alps range. Therefore, it is outside of the Mediterranean region. The avian remains were studied by Cassoli and Tagliacozzo (Bartolomei et al., 1992; Cassoli & Tagliacozzo, 1994). The Mousterian collection is very scarce, but the Aurignacian one constitutes a relatively good sample of the local avifauna. It is characteristic of the Eurosiberian region with presence of some irruptive species from north Europe.
In the west end of the Mediterranean region, Figueira Brava cave (Mourer-Chauviré & Antunes, 1991, 2000) is situated south of Lisbonne, on the coast of the Setubal peninsula. The composition of its Mousterian assemblage is very influenced by the closeness of the site to the sea. The avian local fauna was of Mediterranean type with a high proportion of aquatic taxa, two of them might be considered as northern irruptive species.
Boev (1994) studied the avian remains from Temnata cave, near Karlukovo village, and situated in the middle west of Bulgaria. The largest avian assemblage come from the layer 3d, dated of 28900 ± 1100 aBP. It is not a very rich layer in fossil birds, but the corresponding association of taxa would fit better with a Mediterranean avifauna. On the other hand, the layer 3a, dated of 13600 ± 200 aBP, contained an avian assemblage characteristic of Eurosiberian conditions. None of these assemblages record the occurrence of northern irruptive species.
Šandalja I and Šandalja II caves are situated on the northern coast of Croatia. Their avian remains have been the subject of some works by Malez (fide Tyrberg, 1998). The layer d of the former has been attributed to the Gravettian lithic period (Würm 3) whereas the layer E of the latter to the Aurignacian (with a date of 23540 ± 180 aBP). Both fossil assemblages are characteristic of woodland habitats, although the presence of Lagopus in Šandalja I could be understood as an indicator of a cooler phase, in spite of not having record of any northern irruptive taxa in a so large variety of aquatic habitat species. The same author (Malez fide Tyrberg, 1998) has studied the birds from Vindija cave, situated in the north of the country. Its largest assemblage is dated of ca. 26970 aBP (Würm 3) and corresponds to the layer E + F, which is attributed to the Aurignacian. Thus, the above mentioned assemblages from these three Croatian sites are chronologically very close one to the other. Moreover, the paleoenvironmental features implicated by these local associations are practically the same.
The locality of Ohalo 2, in the Galilee region (Israel), has a rich record from the Early Epipaleolithic (Simmons & Nadel fide Tyrberg, 1998), dated of about 19400 BP. The assemblage corresponds to a typical eastern Mediterranean avifauna, but there are six northern irruptive species recorded.
The avian remains from Bois-des-Brousses shelter were studied by Vilette (1983). This locality is close to the Hérault river, 25 km from Montpellier. The layer 2B (attributed to the Solutrean) contains a no-Mediterranean fossil assemblage and the uncertain presence (cf. Gallinago media) of a northern irruptive species.
In the middle of the Iberian peninsula, the rock shelter named Jarama II has yielded an avian assemblage attributed to the Early Magdalenian phase (Jordá, 1993; Adán et al., 1995) which is clearly of no-Mediterranean type, although there is no record of irruptive species from north Europe.
Castiglione 3 PL is an Upper Pleistocene karstic infilling rich in avian remains in the above mentioned Castiglione 3 complex. Sediments from PL fissure are dated of about 15 ka BP (Salotti et al., 2000). The corresponding avian assemblage is, as for CG fissure, also typical for Mediterranean islands, with the exception of the presence of Alectoris.
Palidoro quarry is situated near Rome. It yielded a reduced avian association, which was attributed to the Upper Paleolithic period (Cassoli, 1977). The taxonomical composition of this association is of Mediterranean type.
Romains cave at Pierre-Châtel, near Virignin village (French Alps), is not today –and probably was never- in the Mediterranean region. Its Magdalenian avian association (Desbrosse & Mourer-Chauviré, 1973; Mourer-Chauviré, 1975a) is consequently characteristic of a woodland habitat under a fresh climate. The high percentage of aquatic species is explained by the nearness of the Rhone river to the cave. The Snowy owl appears in the record of this cavity.
Dimitrijević et al. (2000) have studied the Epigravettian rock shelter of Trebački. This locality is near Berane village, in Yugoslavia. The avian assemblage is characteristic of the Eurosiberian region. There is also the record of the Pine grosbeak (Pinicola enucleator).
The fossil birds from Nerja locality –a cave situated near the homonymous village, in the south of Spain, approximately 1 km from the current sea line (Jordá, 1986)- have been the subject of several works. The most informative assemblages correspond respectively to: Epipaleolithic (Boessneck and Driesch, 1980; Hernández Carrasquilla, 1995; Tyrberg & Hernández Carrasquilla, 1995), Magdalenian (Eastham, 1986) and Upper Paleolithic layers (Hernández Carrasquilla, 1995). The acquatic species constitute the largest group in every assemblage, being the small passeriforms absent or not identified.
Kozarnika cave is located near the town of Bologradchik, north-west Bulgaria. The avian finds were collected in the cultural layers III and IV, which have been attributed to the Gravettian (26 to 19 ka BP) (Boev, 2001). The fossil material was accumulated into the cavity by nocturnal raptors, most probably by the Eagle owl (Bubo bubo) (Boev, 2001). The association of birds accounts with a large representation of Eurosiberian species. It implies woodland or forest habitats with no Mediterranean conditions. The Snowy owl appears as an irruptive immigrant.
Arene Candide is one of the richest Pleistocene avian localities of Europe. The fossil birds from this cave, situated in the Ligurian coast of Italy, were studied by Cassoli (1980). Sediments from the Upper Paleolithic, with a thickness about 3.8 m, show an avian assemblage with twelve northern irruptive species, as well as typical low dispersive Eurosiberian and Mediterranean taxa. This fact may be the result of a mixture of layers –or avifaunas- or, on the contrary, to represent the reality during the Upper Pleistocene in this area.
The site of Mas-d’Azil consists of a large cave situated in the French Pyrenees, 30 km from Foix town. The avian finds, studied by Vilette (1983), come from the cavity named “galerie rive droite”. All the fossils were collected in the Magdalenian layers (13400 ± 1000 and 13200 ± 110 aBP) (Vilette, 1983). As it can be expected by the geographical situation of the locality, the avian taxa imply no-Mediterranean climate. The Snowy owl is recorded together with the Eagle owl. The same author (Vilette, 1983) studied the birds from Cingle Vermell, a shelter located near Vilanova de Sau village, northern Iberia. The avian assemblage of the Tardiglacial layers (dated to 11620 ± 140 aBP) is characteristic of Mediterranean conditions.
Razhishkata cave situated near Lakatnik station, Sofia district (Bulgaria), has a fossil record from the end of the Pleistocene and probably of the transition to the Holocene (Boev, 2000b). The avian assemblage contains low dispersive species linked to a woodland habitat under Eurosiberian conditions.
The richest avian association of the final Upper Pleistocene was recovered from Romanelli cave (south of the Italian peninsula), with two series of dates: 11930 ± 520 and 9050 ± 100 aBP (Cassoli & Tagliacozzo, 1997) and also 10640 ± 100 and 9980 ± 100 aBP, all of them for the formation Terre brune (upper part of the stratigraphical profile) (Tagliacozzo & Gala, 2000). A large amount of water species, with clear evidences of human exploitation of anseriforms (Tagliacozzo & Gala, 2002), and raptors are recorded here, and some irruptive taxa among them.
Cendres cave is in the eastern coast of Spain, near Teulada village. It has yielded an unbalanced sample of the avian paleornithocaenoses, identified by Villaverde et al. (1997), from the end of Magdalenian. The authors point out the possible presence of the irruptive Branta bernicla.
Hayonim cave, situated in the Galilee region of Israel, has some very rich collections of fossil birds related to Natufian period. The layer B, dated of 10.7 to 12.4 ka BP and studied by Tchernov (fide Tyrberg, 1998) as well as the sample studied by Pichon (fide Tyrberg, 1998) provide and exceptional insight to the local avifauna in this period. It is characteristic of the eastern part of the Mediterranean region and it is worth while to mention that we do not find any northern irruptive species in this so large fossil record, unlike we have seen in Ohalo 2, other Galilean locality.
Fontbrégoua locality is a large cavity situated near Salernes village, about 60 km from Toulon, France. The most important avian collection come from the Epipaleolithic layer (50 to 53), with an infradate of 8400 ± 110 (layer 54) and 7600 ± 100 aBP (layers 50 and 51) (Vilette, 1983). The very rich fossil assemblage of birds are characteristic of a zone under Mediterranean conditions, with a predominant woodland habitat. Eurosiberian indicator species are not recorded.
Mourer-Chauviré (1999) studied the Early Holocene fossil bird collection from Akrotiri locality. In spite of its modern age, these avian remains are the oldest known from Cyprus. The small assemblage represents an insular Mediterranean ornithofauna with no occurrence of northern species.
Among the localities from Crete studied by Weesie (1988), Liko is the richest one by far. Liko cave is near Likotenaria village, on the north-west coast of Crete, in a cliff, a few metres above sea level. The fossil assemblage is a wide sample of an eastern Mediterranean avifauna, with insular endemisms, species of eastern distribution and one or two northern irruptive taxa (Aegolius funereus and cf. Pyrrhula pyrrhula). Two other cavities situated not far from here, Gumbes B and Gumbes C, situated also on the north coast of the island, offer two more reduced assemblages with no irruptive species.
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The cooling episodes in the northern hemisphere recorded until now have been noted before in high latitudes than in low ones. Thus, the temperatures fall in Greenland and Iceland around 250 and 150 years, respectively, before than in Europe (Dansgaard et al., 1975; Burton, 1995). It is probable that short-term climatic deteriorations were noted only in high latitudes, having no consequences in the climatic conditions in middle latitudes. Although we may assume that some of such events should lead to movements of birds southwards up to refugia in the MR.
The table 3 shows that the first record of irruptive species from the north of Eurasia in the MR occurs near 1 Ma (Victoria and Dursunlu), in approximate coincidence with the Galerian mammal pulse. From this time up to the end of the Middle Pleistocene, at least, the inner part of the northern Iberian peninsula had tempered conditions of Eurosiberian type (Valdegoba, Elefante, Dolina and Galería). On the contrary, the eastern part held a Mediterranean climate during the all of the Pleistocene, with the exception of occasional enlargements of the Eurosiberian area near the Pyrenees (Gravettian layers from Arbreda). Therefore, the climate in Iberia was in general more fresh and considerable moister than today.
| Localities with irruptive Eurosiberian species | Localities with no irruptive Eurosiberian species | ||
| Presence of grouses - Pygmy - Tengmalm’s owls’ group | No presence of grouses - Pygmy - Tengmalm’s owls’ group | Presence of grouses - Pygmy - Tengmalm’s owls’ group | No presence of grouses - Pygmy - Tengmalm’s owls’ group |
| Arbreda (Gravet.) | Ambrona-Torralba | Bacho Kiro | Akrotiri |
| Arene Candide | Arbreda (Aurign.) | Elefante | Arbreda (Mouster.) |
| Bois-des-Brousses | Arbreda II | Jarama II | Áridos 1 |
| Carnello | Coscia (South) | Lazaret (bottom) | Ca na Reia |
| La Fage (CO) | Coscia (cave) | Radice | Casablanca 1 |
| Fontéchevade | Cova Nova | Razhishkata | Castiglione 3 (CG) |
| Fumane | Dolina | Salpêtre | Castiglione 3 (PL) |
| Kozarnika | Dursunlu | Šandalja I (d) | Cau d’en Borrás |
| Lazaret (loc. VIII) | Em FC Sicily | Šandalja II (E) | Cendres |
| Lazaret (C III) | Figueira Brava | Temnata (3a) | Cingle Vermell |
| Mas-d’Azil | Galería (TG 10B) | Varshets | Devil’s Tower |
| Orgnac 3 (i) | Galería (TG 10A) | Vindija | Es Pouàs |
| Romains | Gorham (K-M) | Fontbrégoua | |
| Salpêtrière | Higueruelas | Galería (TG 11) | |
| Trebački | Huéscar 1 | Gegant | |
| Valdegoba | Ibex | Gorham (B-D) | |
| Liko | Gumbes B | ||
| Nerja (Epipaleolit.) | Gumbes C | ||
| Nerja (Upper Pal.) | Hayonim | ||
| Ohalo 2 | Hortus | ||
| Pedrera de S’Ònix | Montoussé 5 | ||
| Quartaccio | Nerja (Magdalen.) | ||
| Romanelli | Palidoro | ||
| Romaní (II) | Quibas | ||
| Spinagallo | Temnata (3d) | ||
| Victoria | Torre Nave | ||
| Torre in Pietra (d) | |||
| Ubeidiya (II-23) | |||
| Table 3 – List of localities arranged by the presence of irruptive Eurosiberian species and the group of grouses, Pygmy owl and Tengmalm’s owl (see Appendix 2). Localities from islands and Middle East are underlined. | |||
The irruption of northern species in the MR is constantly recorded in the uppermost layers of the Lower Pleistocene (Huéscar 1 and layer 6 from Dolina) and during the Middle Pleistocene, with the exception of CG sediments from Castiglione 3 (Corsica) and Áridos 1 (in the middle of Iberia). The MR seems to have disappeared during this time from the south of France, and to be restricted to south half of the western peninsulas and the islands. We lack data for this period from the east of the MR.
The Mousterian period spread out the Eemian and a large part of the Würmian phases (OIS 3-5). Thus, there follow a succession of shifts in the climatic conditions in the northern hemisphere. The respective positions of some of the localities from this span of time on the chronological table 2 are particularly uncertain. Some northern irruptions are clearly recorded (Valdegoba, layers K and M from Gorham, and Carnello) and it seems also that the paleornithocaenoses from the middle of Italy were quite different than those from the south.
During the Upper Paleolithic the Eurosiberian area extended to inner Iberia (Jarama II), southern France (Bois-des-Brousses and Salpêtrière), northern Italy (Arene Candide and Fumane), Adriatic coast (Šandalja I and Šandalja II) and northern of Balkan peninsula (layer 3a from Temnata, Trebački, Kozarnika and Razhishkata).
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The Mediterranean extension during the Quaternary reached out more to the western part than to the east, as it happens at present. It seems that the Mediterranean conditions never reached the territory of current Bulgaria, although the fossil locations of that area, are situated at lower latitudes than the locations from the south of France and many from Italy and Iberia.
Some northern irruptive species, but group of grouses and Pygmy and Tengmalm’s owls, appear in islands (Coscia, Cova Nova, Sicily Faunal Complex, Liko, Pedrera de S’Ònix) and in the Middle East (Ohalo 2). The reason for their absence it is not the geographical barrier of the Mediterranean sea (at least for owls), neither the reduced time span of cool phases, but the unsuitable climatic conditions for these species.
We have data enough to assume that the geographical boundaries of avian species have changed several times during the Quaternary. Moreover, we may accept that some of these changes were not caused directly by climatic circumstances. As we can observe today, increasing individuals of some northern species lead to quick dispersal movements southwards. It is likely that icecap retreats created suitable conditions for demographic explosions, and some of such events could increase the size of populations to magnitudes unmatched by historic observations.
Some of the fossil locations in the south of France and northern Italy (Arene Candide, Bois-des-Brousses, La Fage, Fontéchevade, Lazaret, Mas-d’Azil, Orgnac 3, Romains and Salpêtrière) point to several displacements of characteristic Eurosiberian low dispersive taxa to the MR during the cool phases, following the shifts of climatic and environmental conditions southwards (Table 3). The large record of the genus Loxia (a group of birds specialized in feeding on seed-cones) in southern France and northern Italy localities (Tyrberg, 1991b) (Table 1), implies the existence of at least some phases, particularly during the Upper Pleistocene, with climatic conditions moist enough to permit the spreading of conifer (most probably of pine) forests. The Mediterranean islands, as well as Iberia and the south of the Italian peninsula, kept out this phenomenon and maintained a rather steady paleoecological condition.
From our point of view, there is no fossil record of irruptive species in the MR during upper Pliocene and early lower Pleistocene. During this span of time, cold pulses only became apparent in the arrival of wintering species into the MR. This fact is unknown until this date, and is firstly observed at Higueruelas locality, which may be considered as the first known avian fossil site of the Quaternary in the Iberian peninsula. From the Middle Pleistocene onwards, the succession of stadials and interstadials cause in Europe two patterns of avian distribution: a) north and central Europe were severely affected by climatic changes and extensions of the ice cap, which probably led to habitat destructions and latitudinal displacements following distributions in east-west belts (Made, 1992), and (b) the MR –at times, only a part of its current extension- supported during the whole of this time a Mediterranean ornithofauna.
In Quaternary fossil localities from the MR we did not find any ornithocaenoses with the same taxonomic composition as those existing at present time in north Eurasia. A number of outcrops that encompass the fossil record of a great part of the Pleistocene points to the the irruption of species from north Europe into Mediterranean avifaunas. This fact does not imply that the climatic conditions in the Mediterranean region were similar to those in the north of the continent. In the North face of the Mediterranean sea tempered conditions stayed, more or less constant, throughout all the Pleistocene. In the south of Europe, not only the Mediterranean region acted like a refuge from the end of the Lower Pleistocene onwards for species that in the interstages occupy more northern territories, but also did in other zones next to the south of Europe. During the coldest periods MR diminished, although south halves of Italy and Iberia, as well as the eastern coast of the latter, always were dominated by Mediterranean faunas. Some species currently characteristic of the MR shared the same habitats in a part of north Iberia, south France, north Italy and most of the Balkan peninsula with species today typical of the Eurosiberian region of Europe.
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Consultar el índice de este documento
Lists of fossil taxa from Mediterranean localities of the Quaternary. The localities are arranged in alphabetical order. Irruptive species for each locality, following the criterium of area of irruption (Appendix 2), are in bold. The group of Pygmy-Tengmalm's owlgrouses (see "areas of irruption" in Appendix 2) is underlined. Nowadays survivors in the east part of the region are in grey fond when recorded in the west area.
| Akrotiri
(Early Holocene) Mourer-Chauviré (1999) |
Podiceps nigricollis Puffinus puffinus Phalacrocorax aristotelis Anser anser s. fabalis Anser sp. Anas crecca cf. Circus sp. Rallus aquaticus Otis tarda Columba livia s. oenas Asio flammeus Athene noctua Turdus iliacus s. philomelos Corvus corone s. frugilegus small Passeriformes |
| Ambrona & Torralba
Sánchez Marco (1990, 1999c) |
Anser anser Tadorna ferruginea Anas strepera Anas cf. acuta Mergus serrator M. merganser Porphyrio porphyrio Fulica cf. atra Vanellus vanellus |
| Arbreda
(Mousterian layers) Garcia (1995) |
Anser s. Branta Anas platyrhynchos Anas sp. Accipiter gentilis Falco naumanni Falco tinnunculus Falco vespertinus Falco subbuteo Alectoris graeca Alectoris rufa Perdix perdix Coturnix coturnix Porzana porzana Calidris canutus Chlidonias nigra Columba livia C. oenas Athene noctua cf. Caprimulgus sp. Coracias garrulus Picus viridis Turdus viscivorus Pyrrhocorax pyrrhocorax P. graculus Corvus monedula C. corone small Passeriformes |
| Arbreda
(Aurignacian layers) Garcia (1995) |
Anser s. Branta Anas platyrhynchos Aquila cf. chrysaetos Falco naumanni F. tinnunculus F. vespertinus F. columbarius F. subbuteo Alectoris graeca A. rufa A. barbara Perdix perdix Coturnix coturnix Rallus aquaticus Porzana porzana Grus grus Tetrax tetrax cf. Charadrius vociferus Gallinago gallinago G. media Larus cf. melanocephalus Columba livia s. oenas C. oenas C. palumbus Otus scops Bubo bubo Athene noctua Turdus viscivorus Garrulus glandarius Pica pica Pyrrhocorax pyrrhocorax P. graculus Corvus monedula small Passeriformes |
| Arbreda
(layers 23-28, Aurignacian) Vilette (1983) |
Podiceps auritus s. nigricollis Branta bernicla Anas crecca Anas platyrhynchos Anas querquedula cf. Falco subbuteo Alectoris graeca Perdix perdix Coturnix coturnix Eudromias morinellus cf. Limosa limosa Columba oenas Athene noctua Asio flammeus cf. Galerida cristata Pyrrhocorax pyrrhocorax P. graculus Corvus corax |
| Arbreda
(Gravettian layers) Garcia (1995) |
Anas sp. Anas platyrhynchos Falco naumanni F. tinnunculus F. vespertinus F. subbuteo Lagopus sp. Alectoris graeca s. rufa A. barbara Perdix perdix Coturnix coturnix Porzana porzana Otis tarda Tetrax tetrax Burhinus oedicnemus Gallinago media Columba livia s. oenas Strix aluco Pica pica Pyrrhocorax pyrrhocorax P. graculus Corvus monedula C. frugilegus s. corone small Passeriformes |
| Arbreda II
(Upper Paleolithic layers) Vilette (1983) |
Falco tinnunculus Alectoris graeca Coturnix coturnix Asio flammeus Turdus torquatus cf. T. merula Pinicola enucleator Pyrrhocorax pyrrhocorax P. graculus |
| Arene Candide
(Upper Paleolithic layers) Cassoli (1980) |
Podiceps auritus Puffinus puffinus Calonectris diomedea Phalacrocorax aristotelis Cygnus cygnus Branta bernicla Anas platyrhynchos A. crecca A. penelope Melanitta nigra Bucephala clangula Mergus albellus M. merganser M. serrator Accipiter gentilis Buteo lagopus Aquila chrysaetos Circus cyaneus C. pygargus C. aeruginosus Pandion haliaetus cf. Falco rusticolus F. eleonorae F. vespertinus F. tinnunculus Tetrao tetrix Lagopus lagopus L. mutus Alectoris graeca Perdix perdix Coturnix coturnix Rallus aquaticus Crex crex Tringa totanus T. glareola Gallinago sp. Calidris canutus Pluvialis squatarola Eudromias morinellus C. alexandrinus Stercorarius pomarinus Sterna paradisaea Uria aalge Alca torda Fratercula arctica Columba livia C. oenas C. palumbus cf. Cuculus canorus Nyctea scandiaca Surnia ulula Athene noctua Strix aluco cf. S. aluco Asio otus A. flammeus Aegolius funereus Apus apus Picus canus Melanocorypha sp. Eremophila alpestris Lullula arborea Ptyonoprogne rupestris Anthus trivialis Cinclus cinclus Prunella collaris Zoothera dauma Turdus viscivorus T. pilaris T. torquatus Plectrophenax nivalis Fringilla coelebs Carduelis chloris C. carduelis C. flammea Acanthis cannabina Pinicola enucleator Loxia pytyopsittacus L. curvirostra L. leucoptera Pyrrhula pyrrhula Coccothraustes coccothraustes Montifringilla nivalis Petronia petronia Sturnus vulgaris Pica pica Nucifraga caryocatactes Pyrrhocorax pyrrhocorax P. graculus Corvus monedula C. corax |
| Áridos 1
Mourer-Chauviré (1980) |
Anas platyrhynchos A. crecca A. clypeata Accipiter nisus Alectoris graeca - rufa Perdix palaeoperdix Porzana porzana Columba oenas C. palumbus Strix aluco Upupa epops Picus viridis Dendrocopos major Galerida cristata Hirundo rustica Turdus pilaris T. iliacus Parus cristatus Coccothraustes coccothraustes Corvus monedula |
| Bacho Kiro
Bocheński (1982) |
Anas platyrhynchos Aquila chrysaetos Circus aeruginosus Lagopus mutus Alectoris graeca Perdix perdix Coturnix coturnix Gallus gallus Rallus aquaticus Porzana porzana Gallinula chloropus cf. Bubo bubo cf. Ptyonoprogne rupestris Delichon urbica cf. Lullula arborea Alauda arvensis cf. Anthus campestris Loxia curvirostra cf. Turdus philomelos Pyrrhocorax pyrrhocorax P. graculus Corvus monedula C. corax |
| Bois-des-Brousses
(Solutrean layers) Vilette 1983) |
Anas platyrhynchos Aythya ferina Falco tinnunculus Lagopus lagopus Lagopus sp. Perdix perdix cf. Gallinago media Scolopax rusticola cf. Calidris temminckii Philomachus pugnax Columba sp. Otus scops Ptyonoprogne rupestris Cinclus cinclus Pyrrhocorax graculus cf. Corvus corone |
| Ca na Reia
Alcover (1989) |
Puffinus nestori cf. Buteo sp. Palaeocryptonyx sp. Columba livia s. oenas Athene cf. veta small Passeriformes |
| Carnello
Segre et al. (1984) |
Cygnus cygnus C. columbianus Anser fabalis Anas platyrhynchos A acuta A penelope Tetrao tetrix Alectoris graeca Strix aluco |
| Casablanca 1
Sánchez Marco (1995b, 1999d) |
Geronticus eremita Gypaetus barbatus Tyto balearica Pterocles orientalis Pyrrhocorax s. Corvus monedula Corvus corone |
| Castiglione 3
(fissure CG) Salotti et al. (1997), Mourer-Chauviré et al. (1997) |
Aquila chrysaetos Buteo cf. buteo B. rufinus Accipiter nisus A. gentilis Porzana porzana Columba livia Bubo insularis Otus sp. Athene angelis Tyto balearica T. alba Pyrrhocorax graculus small Passeriformes |
| Castiglione 3
(fissure PL) Salotti et al. (2000) |
Gyps melitensis Aquila n. sp. Bubo insularis Athene angelis Gypaetus bar batus Falco tinnunculus Alectoris sp. Columba livia s. oenas Tyto alba Lullula arborea Muscicapa striata Turdus spp. Loxia curvirostra Emberiza sp. Pica pica Pyrrhocorax pyrrhoco rax P. graculus Corvus monedula |
| Cau d’en Borrás
Sánchez Marco (unpublished) |
Tadorna tadorna Anas penelope Haliaeetus albicilla cf. Aegipyus monachus Aquila chrysaetos Falco naumanni F. tinnunculus Alectoris rufa Columba livia s. oenas Bubo bubo Athene noctua Apus melba Anthus spinoletta Oenanthe oenanthe Turdus philomelos T. cf. viscivorus Pica pica Pyrrhocorax pyrrhocorax P. graculus Corvus monedula C. corone |
| Cendres
(Upper Magdalenian) Villaverde et al. (1997) |
Anser s. Branta cf. Branta bernicla Aquila chrysaetos Accipiter nisus Falco tinnunculus F. naumanni Alectoris rufa Coturnix coturnix Calidris ferruginea Haematopus ostralegus Columba livia s. oenas Bubo bubo Athene noctua Apus apus Lullula arborea Turdus sp. cf. Turdus viscivorus cf. T. merula Sylvia sp. Pica pica Pyrrhocorax pyrrhocorax P. graculus Corvus monedula C. corone |
| Cingle Vermell
(Tardiglacial layers) Vilette (1983) |
Anser cf. anser Falco tinnunculus Alectoris barbara Perdix perdix Coturnix coturnix Columba livia Apus apus A. melba Melanocorypha calandra Monticola saxatilis Turdus torquatus T. iliacus T. viscivorus Pyrrhocorax pyrrhocorax Corvus monedula |
| Coscia
(shelter South) Bonifay et al. (1998) |
Calonectris diomedea Anser erythropus Anas cf. platyrhynchos A. crecca A. cf. penelope Anatidae indet. Gyps melitensis Buteo buteo B. lagopus Accipiter nisus A. gentilis Milvus sp. Haliaeetus albicilla Circus cf. cyaneus Falco peregrinus F. subbuteo F. naumanni F. tinnunculus Falco sp. Coturnix coturnix Porzana porzana P. pusilla P. parva Crex crex Gallinula chloropus Tetrax tetrax Gallinago gallinago G. media Scolopax rusticola Tringa erythropus Fratercula arctica Columba livia C. palumbus Bubo insularis Asio otus Otus scops Caprimulgus europaeus Apus melba Merops apiaster Coracias garrulus Dendrocopos major cf. Galerida cristata Hirundo cf. rustica Anthus cf. spinoletta Motacilla sp. Lanius collurio L. cf. meridionalis L. minor Saxicola sp. Luscinia cf. megarhynchos Turdus spp. Sylvia atricapilla Emberiza citrinella Fringilla sp. Coccothraustes coccothraustes Sturnus sp. Oriolus oriolus Pyrrhocorax pyrrhocorax Corvus corone s. frugilegus Corvus corax |
| Coscia
(cave) Bonifay et al. (1998) |
Tachybaptus ruficollis Anas platyrhynchos A. platyrhynchos s. acuta A. crecca Anas spp. Gyps melitensis Accipiter nisus Haliaeetus albicilla Falco biarmicus F. naumanni F. tinnunculus Coturnix coturnix Crex crex Tetrax tetrax cf. Pluvialis squatarola Gallinago gallinago Scolopax rusticola Columba cf. livia C. palumbus Bubo insularis Apus melba Coracias garrulus Alauda arvensis Turdus sp. Coccothraustes coccothraustes Prunella collaris Sturnus sp. Pyrrhocorax pyrrhocorax Corvus monedula C. corone small Passeriformes |
| Cova Nova
(Upper Pleistocene) Florit & Alcover (1987) |
Phalacrocorax aristotelis Anas crecca Aquila chrysaetos Falco eleonorae F. tinnunculus Scolopax rusticola Columba livia Apus melba Upupa epops Melanocorypha calandra Hirundo rupestris Lanius minor L. excubitor Prunella collaris Turdus iliacus s. philomelos T. merula T. viscivorus Emberiza cia s. hortulana Loxia curvirostra Pyrrhula pyrrhula Pyrrhocorax pyrrhocorax P. graculus Corvus corone |
| Cova Nova (addenda)
(Upper Pleistocene) McMinn & Alcover (1992) |
Calonectris diomedea Accipiter nissus Falco cf. naumanni Alca torda Tyto alba Prunella modularis Erithacus rubecula Montifringilla nivalis Fringilla sp. |
| Devil’s Tower
Bate (1928) |
Calonectris diomedea Puffinus puffinus Phalacrocorax (?) carbo P. aristotelis Melanitta fusca Mergus cf. serrator Haliaeetus albicilla Gyps fulvus Hieraaetus fasciatus Hieraaetus pennatus Falco (?) naumanni F. eleonorae F. tinnunculus F. subbuteo F. peregrinus Alectoris (?) barbara Larus fuscus Sterna (?) sandvicensis Pinguinus impennis Uria aalge Columba livia C. oenas C. palumbus Apus melba Picus viridis Hirundo rustica Turdus cf. merula T. viscivorus Turdus sp. Fringilla coelebs Passer sp. Pyrrhocorax pyrrhocorax P. graculus |
| Dolina
(layer TD6) Sánchez Marco (1999b and unpublished) |
Anas crecca Melanitta fusca Falco naumanni Perdix palaeoperdix Coturnix coturnix Porzana pusilla Porzana sp. Limosa limosa Eudromias morinellus Scolopax rusticola Columba livia s. oenas Melanocorypha calandra Calandrella cf. brachydactyla Galerida cristata Lullula arborea Alauda arvensis Eremophila alpestris Hirundo rustica Ptyonoprogne rupestris Motacilla flava Anthus pratensis Cinclus cinclus Prunella collaris P. modularis Turdus merula Turdus iliacus Turdus philomelos Sylvia hortensis Muscicapa striata Emberiza citrinella Fringilla coelebs Sturnus sp. Corvus antecorax |
| Dursunlu
Louchart et al. (1998) |
Tachybaptus ruficollis Podiceps cristatus P. nigricollis Phalacrocorax cf. carbo Bubulcus ibis Ardeola ralloides Nycticorax nycticorax Ardeidae indet. Platalea leucorodia Cygnus sp. Anser anser A. cf. erythropus Tadorna ferruginea Tadorna sp. Anas platyrhynchos A. penelolpe A. crecca A. acuta A. querquedula A. aff. clypeata Marmaronetta angustirostris Aythya ferina A. nyroca A. fuligula Aythya sp. Somateria sp. Bucephala sp. Mergus albellus M. merganser Oxyura leucocephala Accipitridae indet. Perdix palaeoperdix Rallus aquaticus Crex crex Porzana porzana Gallinula chloropus Fulica atra Rallidae indet. Tetrax tetrax Otididae indet. Gallinago gallinago Limosa limosa Tringa nebularia Scolopacidae indet. Larus canus Larus sp. Turdus cf. pilaris Fringillidae indet. |
| Elefante
Rosas et al. (2001), Sánchez Marco (unpublished) |
Anas sp. Haliaeetus albicilla Falco cf. tinnunculus Lagopus mutus Perdix palaeoperdix Coturnix coturnix Vanellus vanellus Lymnocryptes minimus Columba livia s. oenas Phoenicurus ochruros Turdus spp. Acanthis flammea Carduelis carduelis C. chloris Pyrrhocorax pyrrhocorax Corvus corax s. frugilegus C. antecorax |
| Elephas mnaidriensis
Faunal Complex of Sicily (K 22, Acquedolci and Contrada Fusco) Pavia (2001) |
Tachybaptus ruficollis Podice ps cristatus P. auritus Phalacrocorax carbo Pelecanus crispus Ixobrychus minutus Botaurus stellaris Egretta garzetta Ardea cinerea Plegadis falcinellus Cygnus falconeri Anser sp. Branta sp. Tadorna tadorna Anas crecca s. querquedula A. platyrhynchos A. clypeata Anas sp. Aythya sp. Mergus merganser Oxyura leucocephala Buteo buteo Gyps melitensis Accipiter gentilis A. nisus Aquila sp. Pandion haliaetus Falco columbarius Coturnix coturnix Fulica atra Grus grus G. cf. melitensis Tetrax tetrax Otis tarda Limosa limosa s. lap ponica Numenius phaeopus Scolopax rusticola Tringa sp. Pterocles orientalis Columba livia s. oenas Bubo bubo Strix aluco Athene noctua Apus melba Coracias garrulus Anthus sp. Erithacus rubecula Turdus sp. Sylvia sp. Sturnus unicolor s. vulgaris Corvus corone |
| Es Pouàs
(Upper Pleistocene) Florit et al. (1989) |
Colonectris diomedea Puffinus puffinus Phalacrocorax aristotelis Anser sp. Falco eleonorae F. tinnunculus Coturnix coturnix Grus primigenia Otis tarda Columba livia Otus scops Asio flammeus Strigidae n. sp. Apus apus Alauda arvensis Hirundo sp. Lanius minor Turdus spp. Pyrrhocorax pyrrhocorax Corvus antecorax |
| Es Pouàs (addendum)
(Upper Pleistocene) Alcover & McMinn (1992) |
Haliaeetus albicilla |
| La Fage
(layer CO) Mourer-Chauviré (1975a) |
Anas platyrhynchos A. crecca Aegypius monachus Aquila chrysaetos Buteo buteo Accipiter nisus Circus macrourus Falco subbuteo F. columbarius F. naumanni F. tinnunculus Lagopus lagopus L. mutus Tetrao urogallus T. tetrix Alectoris graeca Perdix palaeoperdix Coturnix coturnix Rallus aquaticus Porzana porzana Crex crex Otis tarda O. tetrax Vanellus vanellus Pluvialis apricaria P. squatarola Eudromias morinellus Limnocryptes minimus Scolopax rusticola Tringa stagnatilis Calidris alpina Phalaropus fulicarius Glareola pratincola ? Alle alle Columba livia C. oenas C. palumbus Cuculus canorus Nyctea scandiaca Asio flammeus Aegolius funereus ? Picus viridis Dendrocopos major D. minor Calandrella brachydactyla Galerida cristata Lullula arborea Alauda arvensis Ptyonoprogne rupestris Hirundo rustica H. daurica Anthus campestris A. trivialis A. spinoletta Motacilla flava Cinclus cinclus Saxicola rubetra Monticola saxatilis Luscinia luscinia Turdus pilaris T. merula T. viscivorus Emberiza calandra E. citrinella Plectrophenax nivalis Acanthis cannabina Carduelis flammea Serinus citrinella ? Petronia petronia Montifringilla nivalis Pica pica Pyrrhocorax pyrrhocorax P. graculu Corvus monedula Corvus antecorax |
| Férrassie
Mourer-Chauviré (1984) |
Anas crecca Aegypius monachus Aquila chrysaetos Falco tinnunculus Lagopus mutus Lagopus sp. Tetrao tetrix Perdix perdix Coturnix coturnix Pluvialis squatarola Eudromias morinellus Gallinago media Tringa hypoleucos Calidris canutus C. alpina Callandrella brachydactyla Melanocorypha calandra Galerida cristata Lullula arborea Ptyonoprogne rupestris Delichon urbica Cinclus cinclus Turdus pilaris T. merula Emberiza citrinella Carduelis flammea Petronia petronia Pyrrhocorax graculus Corvus corax |
| Figueira Brava
Mourer-Chauviré & Antunes (1991, 2000) |
Gavia stellata Podiceps nigricollis Puffinus holeae Sula bassana Anas platyrhynchos Melanitta nigra M. fusca Clangula hyemalis Aquila chrysaetos Hieraaetus fasciatus Accipiter nisus Milvus migrans Falco cf. tinnunculus Alectoris rufa Grus primigenia Scolopax rusticola Numenius phaeopus Calidris canutus Larus fuscus Pinguinus impennis Columba palumbus Bubo bubo Athene noctua Pyrrhocorax pyrrhocorax |
| Fontbrégoua
(Epipaleolithic) Vilette (1983) |
Accipiter nisus Alectoris cf. rufa Coturnix coturnix Rallus aquaticus Porzana porzana cf. Gallinago gallinago Columba oenas C. palumbus Cuculus canorus Otus scops Strix aluco Caprimulgus europaeus Apus cf. pallidus Merops apiaster Coracias garrulus Upupa epops Picus viridis Dendrocopos major D. medius Galerida cristata Ptyonoprogne rupestris Hirundo rustica H.daurica Anthus pratensis Motacilla flava M. alba Lanius collurio L. senator Prunella modularis Saxicola rubetra Oenanthe hispanica Monticola solitarius Phoenicorus phoenicorus Erithacus rubecula Turdus pilaris T. merula T. philomelos T. viscivorus Acrocephalus palustris Hippolais polyglota Sylvia risoria S. hortensis S. atricapilla S. cantillans S. melanocephala Phylloscopus collybita P. bonelli Regulus sp. Ficedula hypoleuca Parus caeruleus P. major cf. Sitta europaea Emberiza citrinella E. hortulana Fringilla coelebs Acanthis cannabina Carduelis spinus Serinus serinus Pyrrhula pyrrhula Coccothraustes coccothraustes Garrulus glandarius Pyrrhocorax pyrrhocorax P. graculus |
| Fontéchevade
Mourer-Chauviré (1975a) |
Anas platyrhynchos Anas crecca Anas penelope Mergus merganser Mergus merganser s. serrator Buteo buteo Buteo cf. lagopus Lagopus cf. mutus Tetrao tetrix Tetrao urogallus Alectoris barbara Perdix perdix Coturnix coturnix Gallus gallus (?) Crex crex Scolopax rusticola Picus canus Ptyonoprogne rupestris Anthus cf. spinoletta Lanius excubitor Prunella collaris Phoenicurus phoenicurus Erithacus rubecula Turdus sp. Carduelis carduelis Acanthis cannabina Loxia curvirostra Pyrrhula pyrrhula Sturnus vulgaris s. unicolor Sturnus roseus Oriolus oriolus Pica pica >Corvus corone |
| Fumane
Aurignacian layers) Bartolomei et al. (1992), Cassoli & Tagliacozzo (1994) |
Anas platyrhynchos A. querquedula Buteo lagopus Aquila chrysaetos Circus pygargus Falco vespertinus F. tinnunculus F. subbuteo F. columbarius Tetrao tetrix Lagopus mutus Perdix perdix Coturnix coturnix Rallus aquaticus Crex crex Tringa hypoleuca T. glareola Scolopax rusticola Vanellus vanellus Columba oenas Nyctea scandiaca Strix aluco Asio otus Dendrocopos leucotos Eremophila alpestris Lullula arborea Ptyonoprogne rupestris Turdus viscivorus T. pilaris Achantis cannabina Loxia pytyopsittacus L. curvirostra Pyrrhula pyrrhula Montifringilla nivalis Pica pica Garrulus glandarius Nucifraga caryocatactes Pyrrhocorax graculus Corvus monedula |
| Galería
(layer TG10B) Sánchez Marco (1999a) |
Falco naumanni F. tinnunculus Perdix perdix Coturnix coturnix Rallus aquaticus Himantopus himantopus Glareola pratincola Eudromias morinellus Tringa erythropus T. totanus Actitis hypoleucos Gallinago gallinago G. media Calidris alpina Columba livia s. oenas C. palumbus Eremophila alpestris Galerida cristata Lullula arborea Alauda arvensis Prunella collaris Oenanthe oenanthe E. citrinella Pyrrhocorax pyrrhocorax P. graculus Corvus monedula C. antecorax |
| Galería
(layer TG10A) Sánchez Marco (1999a and unpublished) |
Anas platyrhynchos Aegypius monachus Falco naumanni F. tinnunculus Perdix perdix Coturnix coturnix Tetrax tetrax Eudromias morinellus Pluvialis apricaria Actitis hypoleucos Calidris alba Gallinago gallinago G. media Columba livia s. oenas Cuculus canorus Athene noctua Alauda arvensis Oenanthe oenanthe Ficedula hypoleuca Carduelis chloris Pinicola enucleator Pyrrhocorax pyrrhocorax P. graculus Corvus antecorax |
| Galería
(layer TG11) Sánchez Marco (1987a, 1987b, 1995a) |
Anas crecca A. platyrhynchos A. cf. querquedula Falco tinnunculus Perdix palaeoperdix Coturnix coturnix Porzana cf. pusilla Otis tarda Vanellus vanellus Pluvialis apricaria Limosa limosa Tringa erythropus T. totanus Gallinago media G. gallinago Calidris alpina Sterna albifrons Columba livia s. oenas Galerida cristata Lullula arborea Alauda arvensis Turdus merula T. pilaris s. viscivorus T. viscivorus Emberiza citrinella Fringilla coelebs Sturnus sp. Pica pica Pyrrhocorax pyrrhocorax P. graculus Corvus monedula C. antecorax |
| Gegant
(layer II) Sánchez Marco (unpublished) |
Puffinus puffinus Accipiter nisus Alectoris rufa Coturnix coturnix Columba livia s. oenas Tyto alba Athene noctua Strix aluco Anthus spinoletta Turdus viscivorus Emberiza calandra Fringilla coelebs Carduelis chloris Coccothraustes coccothraustes Pyrrhocorax pyrrhocorax P. graculus Corvus monedula Corvus corone |
| Gorham
(layers B and D) Eastham (1968) |
Puffinus yelkouan Phalacrocorax aristotelis Milvus milvus Haliaeetus albicilla Gyps fulvus Falco tinnunculus F. peregrinus Alectoris rufa Charadrius cf. dubius Himantopus ? himantopus Larus fuscus L. argentatus Fratercula arctica Columba livia Bubo bubo Hirundo rustica Fringilla coelebs Sturnus sp. Pyrrhocorax pyrrhocorax P. graculus Corvus monedula C. corone |
| Gorham
(layers K and M) Eastham (1968) |
Gavia stellata Phalacrocorax aristotelis Ardea purpurea ? Tadorna ferruginea Netta rufina Aythya nyroca A. fuligula Clangula hyemalis Melanitta fusca Milvus milvus Gyps fulvus Hieraaetus fasciatus Falco tinnunculus Alectoris rufa Anthropoides virgo ? Fulica atra Haematopus ostralegus Larus ridibundus L. fuscus Pinguinus impennis Alle alle Columba livia Nyctea scandiaca Strix cf. aluco Pyrrhocorax pyrrhocorax P. graculus Corvus monedula C. corone C. corax |
| Gumbes B
Weesie (1988) |
Falco tinnunculus Fulica atra Columba livia C. livia s. oenas C. palumbus Tyto alba Athene cretensis Asio flammeus Turdus sp. Garrulus glandarius Corvus monedula C. corax |
| Gumbes C
Weesie (1988) |
Columba oenas C. livia s. oenas Otus scops Athene cretensis Asio flammeus Dendrocopos leucotos Garrulus glandarius Corvus monedula |
| Hayonim
(Natufian layers) Tchernov (fide Tyrberg, 1998) |
Anser sp. Anas platyrhynchos Aquila pomarina Milvus migrans Buteo buteo Falco tinnunculus F. peregrinus Accipiter nisus A. cf. gentilis Gyps fulvus Alectoris chukar Coturnix coturnix Phasianus colchicus Crex crex Gallinula chloropus Columba livia C. palumbus Streptopelia sp. Athene noctua Otus scops Tyto alba Asio sp. Apus affinis Upupa epops Alcedo atthis Alauda arvensis Melanocorypha calandra Galerida cristata Motacilla cinerea Hirundo daurica H. rustica Hirundo sp. Miliaria calandra Regulus sp. Turdus merula T. philomelos Monticola sp. Oenanthe spp. Phoenicurus sp. Saxicola torquata Erithacus rubecula Luscinia megarhynchos L. svecica Luscinia sp. Sylvia hortensis S. melanocephala Sylvia sp. Phylloscopus sp. Nectarinia osea Lanius senator Passer domesticus Petronia petronia Carduelis carduelis C. spinus C. chloris Coccothraustes coccothraustes Loxia curvirostra Serinus serinus Emberiza caesia Garrulus glandarius Sturnus vulgaris Sturnus sp. Pica pica Pyrrhocorax pyrrhocorax Corvus monedula |
| Hayonim
(Natufian layers) Pichon (fide Tyrberg, 1998) |
Anser anser A. albifrons Tadorna tadorna Anas crecca A. platyrhynchos Aythya nyroca A. fuligula Pandion haliaetus Neophron percnopterus Gyps fulvus Aegypius monachus Accipiter nisus A. gentilis Buteo buteo B. rufinus Buteo sp. Aquila pomarina A. chrysaetos Falco naumanni F. tinnunculus F. subbuteo Perdix perdix Alectoris chukar Coturnix coturnix Phasianus cf. colchicus Grus grus Rallus aquaticus Crex crex Porzana porzana Fulica atra Tetrax tetrax Otis tarda Chlamydotis undulata Vanellus vanellus Pterocles alchata Pterocles sp. Columba livia C. palumbus Streptopelia senegalensis Streptopelia sp. Tyto alba Asio otus Otus scops Athene noctua Strix aluco Apus affinis Alcedo atthis Merops apiaster Coracias garrulus Upupa epops Melanocorypha calandra Alauda arvensis Hirundo rustica H. daurica Motacilla cinerea M. alba Lanius senator L. nubicus Troglodytes troglodytes Nectarinia osea Saxicola torquata Oenanthe oenanthe O. hispanica Monticola solitarius Phoenicurus ochruros Erithacus rubecula Luscinia svecica L. megarhynchos Turdus iliacus T. philomelos T. merula Turdus sp. Prinia gracilis Sylvia hortensis Sylvia borin Sylvia atricapilla S. melanocephala Phylloscopus collybita P. sibilatrix Phylloscopus sp. Regulus regulus Muscicapa striata Remiz pendulinus Parus major Oriolus oriolus Garrulus glandarius Miliaria calandra Emberiza caesia Fringilla coelebs Serinus pusillus Carduelis carduelis C. chloris C. spinus Loxia curvirostra Passer domesticus P. hispaniolensis Passer sp. Petronia petronia Sturnus vulgaris Sturnus sp. Pica pica Corvus monedula C. corone |
| Higueruelas
Sánchez Marco (unpublished) |
Podiceps auritus Podiceps nigricollis Ardea cinerea Nycticorax nycticorax Ixobrychus minutus Plegadis falcinellus Cygnus cygnus Anser sp. Tadorna sp. Marmaronetta angustirostris Aythya sp. Aythya marila Mergus albellus Oxyura leucocephala Palaeocryptonyx sp. Crex crex Actitis hypoleucos Pterocles alchata Columba sp. Anthus pratensis Turdus sp. Turdus iliacus s. philomelos Emberiza citrinella Passer cf. montanus Corvus monedula C. antecorax |
| Hortus
Mourer-Chauviré (1972) |
Gyps fulvus Falco peregrinus F. tinnunculus Falco sp. Alectoris graeca A. barbara Perdix perdix Coturnix coturnix Scolopax rusticola Columba livia C. palumbus Athene noctua Asio accipitrinus Apus melba Picus viridis ? Hirundo daurica Riparia riparia Ptyonoprogne rupestris Motacilla alba Anthus trivialis Muscicapa hypoleuca Turdus viscivorus Emberiza citrinella Coccothraustes coccothraustes Petronia petronia Sturnus vulgaris Nucyfraga caryocatactes Pyrrhocorax pyrrhocorax P. graculus Corvus monedula C. corone |
| Huéscar 1
Sánchez Marco (1989, unpublished) |
Tachybaptus ruficollis Tadorna sp. Anas crecca A. strepera s. acuta A. querquedula A. clypeata cf. Netta rufina Aythya nyroca Aythya sp. Melanitta nigra Mergus serrator cf. Perdix sp. Coturnix coturnix Bubo bubo |
| Ibex
Cooper (2000) |
Puffinus cf. puffinus Sula bassana Branta cf. bernicla Haliaeetus albicilla Aquila cf. chrysaetos Falco naumanni F. tinnunculus F. peregrinus Alectoris cf. rufa Gallus gallus Larus cf. cachinnans Pinguinus impennis Columba livia s. oenas C. palumbus cf. Bubo bubo Apus melba Alaudidae indet. cf. Anthus sp. Prunella cf. modularis P. cf. collaris cf. Monticola solitarius Turdus sp. Fringillidae indet. Coccothraustes s. Emberiza Pyrrhocorax pyrrhocorax Corvus cf. corone C. corax |
| Jarama II
Adan et al. (1995), Sánchez Marco (unpublished) |
Anas platyrhynchos Netta rufina Aquila chrysaetos Buteo buteo Falco tinnunculus Lagopus mutus Alectoris rufa Perdix perdix Tringidae indet. Scolopax rusticola Columba livia s. oenas Columba palumbus Strix aluco Athene noctua Alaudidae indet. Emberiza citrinella Turdus iliacus s. philomelos Pyrrhocorax graculus Corvus monedula |
| Kozarnika
Boev (2001) |
Anas crecca Falco tinnunculus F. vespertinus Tetrao tetrix T. urogallus Lagopus lagopus Lagopus sp. Bonasa bonasia Perdix palaeoperdix P. perdix Coturnix coturnix Alectoris graeca s. chukar Crex crex Gallinula chloropus Porzana cf. Parva cf. Charadrius sp. Tringa totanus T. stagnatilis Tringa sp. Apus apus Nyctea scandiaca Aegolius funereus? Athene noctua Eremophila alpestris Hirundo daurica Ptyonoprogne rupestris Riparia riparia Anthus trivialis Lanius collurio cf. Erithacus sp. Turdus merula T. viscivorus Monticola saxatilis Carduelis cannabina Carduelis carduelis Coccothraustes coccothraustes cf. Pyrrhula pyrrhula Fringilla coelebs Loxia curvirostra Pyrrhocorax cf. pyrrhocorax P. graculus Garrulus glandarius Corvus monedula C. corone Corvus sp. |
| Lazaret
(Locus VIII) Mourer-Chauviré (1975a) |
Nycticorax nycticorax Anas platyrhynchos Aegypius monachus Gypaetus barbatus Accipiter nisus A. gentilis Aquila chrysaetos Buteo buteo Circus cf. cyaneus Falco peregrinus F. tinnunculus F. subbuteo Tetrao tetrix Alectoris graeca A. barbara Perdix palaeoperdix Coturnix coturnix Rallus aquaticus Porzana porzana Crex crex Eudromias morinellus Gallinago gallinago Columba livia C. palumbus Nyctea scandiaca Bubo bubo Athene noctua Asio flammeus Otus scops Aegolius funereus Strix aluco Caprimulgus ruficollis Apus apus A. melba Coracias garrulus Dendrocopos major D. leucotos Picoides tridactylus Jynx torquilla Lullula arborea Alauda arvensis Riparia riparia Ptyonoprogne rupestris Lanius senator Troglodytes troglodytes Prunella collaris Saxicola rubetra Turdus merula T. pilaris T. iliacus T. viscivorus Sylvia communis Aegithalos caudatus Emberiza citrinella Fringilla coelebs Serinus serinus? Coccothraustes coccothraustes Passer montanus Petronia petronia Montifringilla nivalis Sturnus vulgaris Garrulus glandarius Pica pica Pyrrhocorax pyrrhocorax P. graculus Corvus monedula C. antecorax |
| Lazaret
(Bottom) Mourer-Chauviré (1975a) |
Accipiter nisus A. gentilis Aquila chrysaetos Alectoris graeca A. barbara Perdix palaeoperdix Coturnix coturnix Porzana cf. parva Scolopax rusticola Numenius arquata Larus fuscus Columba livia C. palumbus Bubo bubo Athene noctua Otus scops Aegolius funereus Strix aluco Apus apus Picus viridis Dendrocopos major Galerida cristara Hirundo rustica Anthus spinoletta Troglodytes troglodytes Prunella modularis Turdus merula T. pilaris T. iliacus T. viscivorus Fringilla coelebs Carduelis flammea Coccothraustes coccothraustes Passer domesticus Petronia petronia Montifringilla nivalis Garrulus glandarius Pica pica Pyrrhocorax pyrrhocorax P. graculus Corvus monedula C. corone C. cf. corax |
| Lazaret
(layer C III) Vilette (1993) |
Anas cf. acuta cf. A. clypeata Aythya cf. marila Bucephala clangula Aegypius monachus Gypaetus barbatus cf. Buteo buteo Aquila chrysaetos cf. Hieraaetus fasciatus Haliaeetus albicilla Circus cf. cyaneus C. macrourus Falco subbuteo F. eleonorae F. cf. columbarius F. cf. vespertinus F. tinnunculus Tetrao tetrix Alectoris graeca Perdix paleoperdix Coturnix coturnix cf. Rallus aquaticus Crex crex Tringa stagnatilis cf. Pluvialis squatarola cf. Charadrius dubius cf. Eudromias morinellus Scolopax rusticola Larus cf. canus Sterna cf. hirundo Chidonias hybrida Uria lomvia s. aalge Fratercula arctica Burhinus oedicnemus Columba livia C. cf. oenas C. palumbus Cuculus canorus Nyctea scandiaca Bubo bubo Athene noctua Strix cf. aluco Otus scops Asio otus A. flammeus Aegolius funereus Apus apus A. melba A. cf. pallidus cf. Dendrocopos major cf. D. medius cf. D. minor Galerida cristata Lullula arborea Alauda arvensis Anthus pratensis Motacilla alba Cinclus cinclus Oenanthe hispanica Phoenicurus phoenicurus Erithacus rubecula Turdus viscivorus T. pilaris T. merula T. iliacus Emberiza hortulana Plectrophenax nivalis Fringilla coelebs Pinicola enucleator Loxia pytyopsittacus Carpodacus erythrinus Pyrrhula pyrrhula Coccothraustes coccothraustes Montifringilla nivalis Petronia petronia Sturnus vulgaris Garrulus glandarius Pica pica Nucyfraga cariocatactes Pyrrhocorax pyrrhocorax P. graculus Corvus monedula C. corone C. corax |
| Liko
Weesie (1988) |
Phalacrocorax aristotelis cf. Branta ruficollis Anas penelope A. querquedula Haliaeetus albicilla Gypaetos barbatus Gyps fulvus G. melitensis Aegypius monachus Accipiter gentilis A. cf. nisus Buteo buteo Aquila chrysaetos A. c. simurgh Falco tinnunculus F. subbuteo F. eleonorae F. peregrinus Coturnix coturnix Porzana porzana Gallinulla chloropus Glareola sp. Calidris canutus Scolopax rusticola Columba livia C. oenas C. palumbus Tyto alba Otus scops Ketupa zeylonensis Athene cretensis Asio flammeus Aegolius funereus Caprimulgus cf. europaeus Apus cf. apus A. melba Dendrocopos leucotos cf. Calandrella brachydactyla cf. Hirundo daurica Prunella collaris cf. Erithacus rubecula Oenanthe cf. hispanica Monticola cf. solitarius cf. Zoothera dauma cf. Turdus iliacus Turdus spp. cf. Muscicapa striata Fringilla sp. Carduelis chloris cf. Pyrrhula pyrrhula Coccothraustes coccothraustes cf. Emberiza calandra Emberiza spp. Sturnus sp. Garrulus glandarius Pyrrhocorax pyrrhocorax P. graculus Corvus monedula Corvus sp. |
| Malagrotta
Cassoli et al. (1982) |
Anser brachyrhynchus Branta bernicla Anas platyrhynchos A. acuta A. penelope A. querquedula Calidris temminckii Rallus aquaticus Perdix perdix |
| Mas-d’Azil
(Magdalenian layers) Vilette (1983) |
Anas platyrhynchos A. crecca Aythya ferina Aegypius monachus Falco tinnunculus Lagopus lagopus L. mutus Perdix perdix Larus canus Columba oenas Columba sp. Nyctea scandiaca Bubo bubo cf. Lanius excubitor cf. Turdus viscivorus Montifringilla nivalis Sturnus vulgaris Pyrrhocorax graculus Corvus corax |
| Montoussé 5
Clot et al. (1976) |
Palaeocryptonyx sp. cf. Dendrocopos major Oenanthe sp. Turdus cf. merula T. cf. viscivorus Ficedula sp. cf. Garrulus glandarius Corvus cf. monedula C. pliocaenus |
| Nerja
(Magdalenian layers) Eastham (1986b) |
Puffinus griseus Sula bassana Anser sp. Anas platyrhynchos A. crecca Aythya nyroca Circaetus gallicus Falco tinnunculus Alectoris rufa Larus fuscus Uria aalge Pinguinus impennis Columba livia Monicola solitarius Lanius excubitor Coccothraustes coccothraustes Pyrrhocorax pyrrhocorax P. graculus C. corone |
| Nerja
(Epipaleolithic layers) Boesneck & Driesch (1980) |
? Gavia stellata Calonectris diomedea Puffinus gravis P. puffinus Sula bassana ? Anser albifrons Tadorna tadorna Anas platyrhynchos Anas sp. ? Aythya ferina Melanitta nigra Milvus milvus Gypaetus barbatus Accipiter gentilis Buteo buteo Aquila heliaca Alectoris rufa Grus grus Fulica atra ? Larus canus L. argentatus L. fuscus L. marinus Pinguinus impennis Alca torda Uria aalge Columba livia Tyto alba Bubo bubo Athene noctua Sturnus sp. Pyrrhocorax pyrrhocorax Corvus corone C. corax |
| Nerja
(Upper Paleolithic layers) Hernández Carrasquilla 1995) |
Gavia stellata Calonectris diomedea Puffinus yelkouan P. aff. griseus Sula bassana Phalacrocorax carbo P. aristotelis Branta bernicla Tadorna ferruginea T. tadorna s. ferruginea Anas clypeata Aythya nyroca A. ferina s. fuligula Milvus milvus Buteo buteo Alectoris rufa Fulica atra Grus grus Larus marinus L. canus Uria aalge Alca torda Pinguinus impennis Columba livia s. oenas Bubo bubo Alauda arvensis Hirundo rustica Turdus sp. Pyrrhocorax pyrrhocorax Corvus corax |
| Nerja
(Epipaleolithic layers) Hernández Carrasquilla (1995)
|
Sula bassana Phalacrocorax aristotelis Ardea cinerea Anas platyrhynchos Stercorarius skua Larus cachinnans s. argentatus s. fuscus Uria aalge Columba livia s. oena |
| Nerja
(Epipaleolithic layers) Tyrberg & Hernández Carrasquilla (1995) |
Catharacta skua |
| Ohalo 2
Simmons & Nadel (1998) |
Podiceps auritus P. cristatus P. grisegena P. nigricollis Tachybaptus ruficollis Phalacrocorax aristotelis P. pygmaeus Ardea cinerea A. purpurea Ardeola ralloides Egretta garzetta Platalea leucorodia Plegadis falcinellus Cygnus cygnus C. columbianus Anser anser A. albifrons A. fabalis Tadorna tadorna Alopochen aegyptiacus Anas acuta A. capensis A. clypeata A. crecca A. penelope A. platyrhynchos A. querquedula A. strepera Aythya fuligula A. marila Netta rufina Mergus merganser M. serrator Melanitta fusca Bucephala clangula Haliaeetus albicilla Milvus migrans Circus aeruginosus C. cyaneus Accipiter nisus A. gentilis Melierax metabates Buteo rufinus B. buteo Aquila rapax Falco tinnunculus F. columbarius F. biarmicus F. cherrug Alectoris chukar Ammoperdix heyi Coturnix coturnix Fulica atra Porphyrio porphyrio Tetrax tetrax Otis tarda Himantopus himantopus Recurvirostra avosetta Vanellus vanellus Numenius phaeopus N. arquata Arenaria interpres Larus minutus L. sabini L. argentatus Bubo bubo Strix aluco s. butleri Corvus monedula C. corone C. frugilegus |
| Orgnac 3
(layer i) Mourer-Chauviré (1975a) |
Anas platyrhynchos Aythya nyroca A. marila Accipiter nisus Falco naumanni F. tinnunculus Lagopus sp. Tetrao urogallus T. tetrix Alectoris graeca A. barbara Perdix palaeoperdix Coturnix coturnix Crex crex Pluvialis squatarola Scolopax rusticola Tringa cf. stagnatilis Columba livia C. palumbus Cuculus canorus Nyctea scandiaca Asio flammeus Otus scops Aegolius funereus Athene noctua Strix aluco Caprimulgus europaeus Apus melba Coracias garrulus Upupa epops Picus viridis Dendrocopos major D. medius Jynx torquilla Melanocorypha calandra Galerida cristata Lullula arborea Alauda arvensis Ptyonoprogne rupestris Motacilla alba Bombycilla garrulus Troglodytes troglodytes Saxicola rubetra Turdus merula T. iliacus T. viscivorus Acrocephalus arundinaceus ? Sylvia nisoria S. atricapilla S. melanocephala Ficedula hypoleuca Parus cristatus Emberiza citrinella ? E. hortulana ? Carduelis chloris Serinus citrinella Pyrrhula pyrrhula Coccothraustes coccothraustes Garrulus glandarius Pica pica Corvus corone C. antecorax |
| Palidoro
Cassoli (1977)
|
Anser fabalis Falco tinnunculus Perdix perdix Coturnix coturnix Columba livia Athene noctua Melanocorypha calandra Alauda arvensis Pyrrhocorax pyrrhocorax P. graculus Corvus corone |
| Pedrera de S’Ònix
Mourer-Chauviré et al. (1977) |
Aegypius monachus Scolopax rusticola Troglodytes troglodytes ? Turdus iliacus Aegithalos caudatus ? Parus cristatus ? Fringilla coelebs Carduelis carduelis Corvus pliocaenus |
| Pedrera de S’Ònix
Mourer-Chauviré in Alcover et al. (1981) |
Cygnus cf. cygnus Bucephala cf. clangula Coturnix cf. coturnix cf. Porzana porzana Otus cf. scops Tyto balearica Dendrocopos cf. major Melanocorypha cf. calandra cf. Lullula arborea Sylvia cf. atricapilla Parus cf. ater Prunella cf. modularis Erithacus cf. rubecula Turdus cf. merula T. cf. iliacus s. philomelos Fringilla cf. coelebs Carduelis cf. carduelis Coccothraustes cf. coccothraustes Pica pica Pyrrhocorax cf. pyrrhocorax Corvus pliocaenus |
| Pedrera de S’Ònix
Seguí (2001) |
Pica mourerae |
| Romains
Desbrosse & Mourer-Chauviré (1973) |
Podiceps auritus Cygnus olor C. cygnus Anas platyrhynchos A. crecca A. strepera A. acuta A. querquedula A. clypeata Aythya fuligula Melanitta nigra Clangula hyemalis Gypaetus barbatus Aquila chrysaetos Buteo buteo Falco rusticolus F. tinnunculus Lagopus lagopus L. mutus Tetrao tetrix Perdix perdix Grus grus ? Gallinago gallinago Scolopax rusticola Numenius arquata N. tenuirostris Tringa totanus Calidris alba Larus argentatus L. canus Sterna sandvicensis Columba livia Cuculus canorus Nyctea scandiaca Asio otus A. flammeus Strix aluco Apus apus A. melba Dryocopus martius Dendrocopos medius Lullula arborea Hirundo daurica Delichon urbica Prunella collaris Turdus cf. torquatus T. merula T. iliacus Acrocephalus paludicola Sylvia atricapilla Ficedula hypoleuca Emberiza hortulana Pyrrhula pyrrhula Coccothraustes coccothraustes Montifringilla nivalis Garrulus glandarius Pica pica Pyrrhocorax graculus Corvus corone C. corax |
| Quartaccio
Bedetti (2001) |
Podiceps grisegena Ardea purpurea Anser sp. Branta ruficollis Anas platyrhinchos Anas clypeata Somateria molissima Mergus serrator Fulica atra Gallinago sp. Sturnus vulgaris Pyrrhula pyrrhula |
| Quibas
Montoya et al. (1999, 2001), Sánchez Marco (unpublished) |
Geronticus eremita Gypaetus barbatus Coturnix coturnix Columba livia s. oenas Cuculus canorus Athene noctua Delichon urbica Prunella modularis Anthus pratensis Phoenicurus ochruros Saxicola torquata S. rubetra Oenanthe hispanica Parus major Carduelis chloris C. carduelis C. spinus Serinus serinus |
| Radice
Bidduttu et al. (1967) |
Tetrao urogallus T. tetrix Perdix perdix Alectoris graeca Strix aluco Pyrrhocorax pyrrhocorax P. graculus |
| Razhishkata
Boev (2000b) |
Anser sp. Anas sp. Tetrao tetrix Bonasa bonasia Perdix palaeoperdix P. perdix Coturnix coturnix Crex crex Tringa cf. stagnatilis Athene noctua Asio otus Apus melba Melanocorypha sp. Anthus cf. trivialis Anthus sp. Parus major Sylvia sp. Ptyonoprogne rupestris Fringilla montifringilla Loxia curvirostra Coccothraustes coccothraustes Carduelis chloris Carduelinae indet. Petronia petronia cf. Garrulus glandarius Pyrrhocorax pyrrhocorax P. graculus Corvus monedula C. corone s. frugilegus Corvus sp. |
| Romanelli
Cassoli & Tagliacozzo (1997) |
Podiceps auritus P. nigricollis Gavia arctica G. stellata Puffinus puffinus Pelecanus crispus Phalacrocorax carbo P. aristotelis Ardea cinerea Nycticorax nycticorax Cygnus cygnus Anser anser A. erythropus A. albifrons A. fabalis A. brachyrhynchus A. caerulescens Branta bernicla B. leucopsis B. ruficollis Tadorna tadorna Anas platyrhynchos A. crecca A. strepera A. penelope A. acuta A. querquedula A. clypeata Netta rufina Aythya ferina A. nyroca A. fuligula Melanitta fusca Bucephala clangula Clangula hyemalis Oxyura leucocephala Mergus albellus M. serrator Accipiter gentilis Buteo lagopus Aquila chrysaetos A. heliaca Haliaeetus albicilla Aegypius monachus Gyps fulvus Circus cyaneus C. macrourus C. aeruginosus Falco peregrinus F. eleonorae F. subbuteo F. vespertinus F. tinnunculus Alectoris graeca A. rufa Coturnix coturnix Crex crex Gallinula chloropus Fulica atra Grus grus G. cf. leucogeranus Anthropoides virgo Otis tetrax O. tarda Numenius arquata N. phaeopus Limosa limosa Scolopax rusticola Philomachus pugnax Vanellus vanellus Pluvialis squatarola P. apricaria Eudromias morinellus Burhinus oedicnemus Larus marinus L. argentatus Rissa tridactyla Pinguinus impennis Columba livia C. oenas Streptopelia turtur Pterocles alchata P. orientalis Tyto alba Otus scops Bubo bubo Nyctea scandiaca Athene noctua Asio otus A. flammeus Apus apus Galerida cristata Ptyonoprogne rupestris Hirundo rustica Zoothera dauma Turdus viscivorus T. iliacus T. pilaris Loxia curvirostra Pyrrhula pyrrhula Sturnus vulgaris Pica pica Nucifraga caryocatactes Pyrrhocorax pyrrhocorax P. graculus Corvus monedula C. frugilegus C. corone C. corax |
| Romaní
(unit II) Sánchez Marco (unpublished) |
Anas platyrhynchos A. acuta Buteo buteo B. rufinus Alectoris rufa Perdix perdix Columba livia s. oenas Athene noctua Delichon urbica Hirundo rustica Turdus sp. Pinicola enucleator Pica pica Pyrrhocorax pyrrhocorax |
| Salpêtre
(Mousterian layers) Mourer-Chauviré (1975a) |
Accipiter nisus Falco columbarius F. naumanni F. tinnunculus Lagopus lagopus L. mutus Lyrurus tetrix Alectoris graeca Perdix perdix Coturnix coturnix Eudromias cf. morinellus Scolopax rusticola Columba livia Cuculus canorus Asio flammeus Otus scops Athene noctua Dendrocopos major D. medius Jynx torquilla? Galerida cristata Hirundo daurica Saxicola rubetra Turdus merul T. viscivorus Emberiza citrinella Serinus citrinella Coccothraustes coccothraustes Montifringilla nivalis Garrulus glandarius Pica pica Pyrrhocorax pyrrhocorax P. graculus Corvus corax |
| Salpêtre
(Dryas 3 / Preboreal layers) Vilette et al. (1983) |
Aythya fuligula Falco subbuteo F. naumanni F. tinnunculus Lagopus sp. Alectoris graeca s. rufa Perdix perdix Coturnix coturnix Scolopax rusticola Tringa sp. Columba cf. livia Cuculus canorus Otus scops Athene noctua Glaucidium passerinum Merops apiaster Upupa epops Dendrocopos medius Jynx torquilla Ptyonoprogne rupestris Hirundo rustica Lanius senator Troglodytes troglodytes Erithacus rubecula Turdus pilaris T. merula T. iliacus s. philomelos T. viscivorus Acrocephalus arundinaceus Acrocephalus sp. Sylvia atricapilla Phylloscopus sp. Ficedula hypoleuca Parus cristatus P. ater Emberiza citrinella Fringilla coelebs Coccothraustes coccothraustes Passer domesticus Pica pica Nucifraga caryocatactes Pyrrhocorax pyrrhocorax P. graculus |
| Salpêtrière
(Aurignacian layers) Vilette (1983) |
Anas platyrhynchos A. clypeata Aythya fuligula cf. Falco peregrinus F. subbuteo Lagopus lagopus Lagopus sp. cf. Lyrurux tetrix Perdix perdix Coturnix coturnix Pluvialis apricaria Charadrius hiaticula Eudromias morinellus Lymnocryptes minimus Numenius arquata Columba oenas C. livia Asio otus Galerida cristata Lullula arborea Hirundo rustica Cinclus cinclus cf. Turdus merula T. viscivorus Loxia pytyopsittacus Petronia petronia Pletrophenax nivalis Pyrrhocorax graculus Corvus corone |
| Šandalja I
(layer d) Malez (several publications, fide Tyrberg, 1998) |
Tachybaptus ruficollis Anser fabalis Anas penelope A. acuta A. crecca Aythya ferina Aegypius monachus Accipiter nisus Falco vespertinus F. tinnunculus Lagopus lagopus L. mutus Tetrao tetrix Bonasa bonasia Coturnix coturnix Phasianus colchicus Rallus aquaticus Porzana sp. Fulica atra Tetrax tetrax Scolopax rusticola Limosa limosa Larus argentatus L. minutus Sterna hirundo Columba livia Asio otus A. flammeus Dendrocopos minor Galerida cristata Hirundo rustica H. cf. Daurica Delichon urbica Motacilla flava Bombycilla garrulus Erithacus rubecula Sitta europaea Emberiza citrinella Fringilla coelebs Coccothraustes coccothraustes Pyrrhula pyrrhula Sturnus vulgaris Lanius collurio Pica pica Pyrrhocorax graculus Corvus monedula C. corone C. corax |
| Šandalja II
(layer E) Malez (several publications, fide Tyrberg, 1998) |
Podiceps auritus Falco subbuteo F. vespertinus F. tinnunculus Tetrao tetrix T. urogallus Coturnix coturnix Grus grus Rallus aquaticus Otis tarda Tetrax tetrax Vanellus vanellus Scolopax rusticola Larus ridibundus Columba livia Tyto alba Bubo bubo Asio flammeus Dendrocopos medius Dendrocopos minor Galerida cristata Motacilla alba Garrulus glandarius Pica pica Pyrrhocorax pyrrhocorax P. graculus Corvus monedula |
| Spinagallo
Pavia (1999) |
Geronticus eremita Anser erythropus Branta sp. Anas penelope A. querquedula Marmaronetta angustirostris Accipiter gentilis A. nisus Falco tinnunculus F. columbarius F. subbuteo F. eleonorae Coturnix coturnix Rallus aquaticus Grus sp. Recurvirostra avosetta Scolopax rusticola Larus minutus L. ridibundus Columba livia C. livia s. oenas C. palumbus Streptopelia turtur Cuculus canorus Tyto sp. Otus scops cf. Surnia ulula Athene sp. Asio otus Caprimulgus cf. europaeus Apus apus s. pallidus A. melba Picus viridis Dendrocopos leucotos Calandrella brachydactyla Lullula arborea Hirundo sp. Anthus sp. Prunella modularis Erithacus rubecula Oenanthe cf. hispanica Monticola solitarius Turdus spp. Sylvia sp. Phylloscopus sibilatrix s. collybita Lanius senator Sturnus sp. Petronia petronia Fringilla coelebs s. montifringilla Serinus sp. Carduelis chloris Carduelis sp. Pyrrhula pyrrhula Coccothraustes coccothraustes Emberiza spp. Pica pica Pyrrhocorax graculus Corvidae indet. Passeriformes indet. |
| Temnata
(layer 3d) Boev (1994) |
Aquila pomarina Aegypius monachus Circus cyaneus Falco tinnunculus Falco sp. Alectoris graeca Perdix perdix Streptopelia sp. Columba oenas Bubo bubo Alauda cf. arvensis Nucifraga caryocatactes Pica pica Corvus monedula |
| Temnata
(layer 3a) Boev (1994) |
Falco tinnunculus Falco cf. subbuteo Tetrao sp. Perdix perdix Coturnix coturnix Strix aluco Asio otus cf. Aegolius sp. Apus apus Dendrocopos minor Turdus viscivorus Pyrrhula pyrrhula Sturnus vulgaris Pyrrhocorax graculus Corvus monedula Corvus sp. |
| Torre Nave
Cassoli in Bulgarelli (1972) |
Ixobrychus minutus Anas platyrhynchos Falco subbuteo F. tinnunculus Alectoris graeca Perdix perdix Coturnix coturnix Otis tetrax Crex crex Columba livia C. palumbus Pterocles orientalis Athene noctua Strix aluco Apus melba Picus viridis Dendrocopos medius Hirundo rustica Turdus pilaris T. viscivorus T. philomelos T. merula Carduelis chloris Garrulus glandarius Pyrrhocorax pyrrhocorax P. graculus Corvus monedula C. frugilegus C. corax |
| Torre in Pietra
(layer d) Cassoli (1978) |
Phalacrocorax carbo Botaurus stellaris Ixobrychus minutus Cygnus cygnus Anser fabalis A. albifrons Anas platyrhynchos A. penelope A. acuta Aythya nyroca Pandion haliaeetus Alectoris graeca Perdix perdix Crex crex Alauda arvensis Turdus pilaris T. iliacus T. merula Nucifraga caryocatactes |
| Trebački
(Epigravettian) Dimitrijević et al. (2000) |
Falco sp. Lagopus lagopus Tetrao tetrix Crex crex Gallinago gallinago Apus apus Galerida cristata Alauda arvensis Lullula arborea Hirundo rustica Delichon urbica Lanius minor Turdus viscivorus T. merula T. pilaris T. philomelos T. cf. torquatus T. cf. iliacus Luscinia luscinia Sturnus vulgaris Fringilla montifringilla F. cf. coelebs Carduelis chloris Pyrrhula pyrrhula Coccothraustes coccothraustes Pinicola enucleator Emberiza citrinella< Passer domesticus Passeriformes indet. |
| Ubeidiya
(layer II-23) Tchernov (several publications, fide Tyrberg, 1998) |
Podiceps cristatus P. auritus Phalacrocorax africanus P. carbo Anhinga rufa Tadorna tadorna Anas crecca A. acuta A. penelope Aythya sp. Milvus pygmaeus Falco subbuteo Falco sp. Aquila sp. Alectoris baryosefi Francolinus sp. Rallus aquaticus Porzana spp. Crex crex Fulica stekelesi Porphyrio porphyrio Columba livia Strix butleri Asio cf. capensis Ketupa zeylonensis Melanocorypha calandra M. gracilis Alauda jordanica cf. Calandrella sp. Motacilla alba M. cf. cinerea Saxicola cf. torquata Oenanthe sp. Cercomela cf. melanura Turdus sp. Acrocephalus sp. Lanius excubitor Lanius sp. Parus sp. Petronia brevirostris Fringilla coelebs Carduelis cf. chloris Sturnus vulgaris Sturnus sp. Pica pica Corvus monedula C. corone C. corax |
| Valdegoba
Sánchez Marco (unpublished) |
Anser fabalis Anas crecca A. platyrhynchos A. platyrhynchos s. acuta A. querquedula Somateria sp. Milvus migrans Gypaetus barbatus Gyps fulvus Aegypius monachus Accipiter gentilis Buteo buteo B. lagopus Falco naumanni F. tinnunculus F. peregrinus Lagopus mutus Tetrao tetrix Alectoris graeca Coturnix coturnix Otis tetrax Columba livia s. oenas C. palumbus Caprimulgus sp. Athene noctua Alauda arvensis Hirundo rustica Delichon urbica Anthus spinoletta Prunella collaris P. modularis Oenanthe oenanthe Turdus sp. Muscicapa striata Emberiza calandra E. citrinella Acanthis cf. cannabina Pinicola enucleator Petronia petronia Sturnus sp. Pica pica Pyrrhocorax pyrrhocorax P. graculus |
| Varshets
Boev (2002) |
Anatini indet. Accipiter sp. Circaetus sp. Aquila cf. clanga Aquila sp. Hieraaetus cf. fasciatus Hieraaetus sp. Gyps sp. Buteo sp. Accipitridae indet. Falco cf. tinnunculus F. bakalovi Falco sp. Tetrao aff. partium Lagopus balcanicus cf. Perdix sp. Chauvireria balcanica Phasianus sp. Gallinula balcanica Porzana sp. Otis aff. khosatzkii Otididae indet. Actitis balcanica Charadriiformes indet. Apus baranensis Columba spp. Streptopelia sp. Athene sp. Anthus sp. Motacilla sp. Melanocorypha sp. Alauda sp. Lullula sp. Lullula arborea Galerida spp. Eremophila sp. Parus sp. Regulus bulgaricus cf. Muscicapa sp. Fringilla sp. Loxia patevi Coccothraustes simeonovi Carduelis sp. Emberiza sp. Turdus cf. merula T. cf. philomelos T. cf. iliacus Turdus sp. Erithacus sp. Sturnus sp. Pyrrhocorax cf. pyrrhocorax P. cf. graculus Pica sp. Corvus cf. monedula Corvus sp. Corvidae indet. |
| Victoria
Sánchez Marco (unpublished) |
Cygnus cf. olor Tadorna tadorna cf. Anas crecca Milvus migrans cf. Aegypius monachus Buteo cf. buteo Falco peregrinus Tetrao tetrix Alectoris graeca Himantopus himantopus Syrrhaptes s. Pterocles Columba livias. oenas C. palumbus Tyto alba Bubo bubo Athene noctua Strix aluco Picus viridis Dryocopus martius Calandrella cf. brachydactyla Lullula arborea Alauda arvensis Hirundo rustica Anthus spinoletta Turdus sp. Prunella modularis Sylvia hortensis S. cf. atricapilla Ficedula hypoleuca Emberiza cf. citrinella E. melanocephala Carduelis carduelis Sturnus sp. Pica pica Pyrrhocorax pyrrhocorax P. graculus Corvus monedula C. frugileguss. corone C. antecorax |
| Vindija
(layer E + F) Malez (several publications, fide Tyrberg, 1998) |
Ardeola sp. Anas platyrhynchos A. querquedula A. crecca Aythya fuligula A. nyroca A. ferina Mergus merganser Buteo buteo Accipiter gentilis Falco rusticolus Falco subbuteo Lagopus mutus L. lagopus Tetrao urogallus T. tetrix Alecoris graeca Phasianus colchicus Gallinula chloropus Rallus aquaticus Vanellus vanellus Recurvirostra avosetta Scolopax rusticola Gallinago gallinago Larus minutus Tyto alba Bubo bubo Nyctea scandiaca Asio otus A. flammeus Strix aluco Dendrocopos major Hirundo rustica Lanius excubitor Sylvia borin Sturnus vulgaris Nucifraga caryocatactes Garrulus glandarius Pica pica Corvus corone C. monedula Pyrrhocorax pyrrhocorax P. graculus |
Consultar el índice de este documento
Appendix 2 – Northern irruptive species in the Mediterranean region during the Quaternary. Current distributions after Cramp (1998).
| Specie | Cygnus olor |
| Current European Distribution | Resident in Europe from France east to Black sea. Wintering and resident in zones of southern Italy, Balkans and Anatolia. |
| Areas of Irruption | Irruptive species in the islands, Iberia and part of Italy. |
| Direction of Movements | Movements to south and west. |
| Specie | Cygnus cygnus |
| Current European Distribution | Winters in north and central Europe, Adriatic coast, southern Balkans, Black sea and southern Anatolia. |
| Areas of Irruption | Irruptive species in the islands, Iberia and Italy. |
| Direction of Movements | Movements to south and west. |
| Specie | Cygnus columbianus |
| Current European Distribution | Winters in coasts of British islands, from Germany west to north France, and in inland points to Azure coast. |
| Areas of Irruption | Irruptive in the whole Mediterranean region, except in Azure and Adriatic coasts. |
| Direction of Movements | Movements to south. |
| Specie | Anser brachyrhynchus |
| Current European Distribution | Winters in British islands and north Germany. |
| Areas of Irruption | Irruptive in the whole Mediterranean region. |
| Direction of Movements | Movements to south. |
| Specie | Branta bernicla |
| Current European Distribution | Winters in coasts from Denmark to west France. |
| Areas of Irruption | Irruptive in the whole Mediterranean region. |
| Direction of Movements | Movements to south. |
| Specie | Branta leucopsis |
| Current European Distribution | Winters in British islands and north Germany. |
| Areas of Irruption | Irruptive in the whole Mediterranean region. |
| Direction of Movements | Movements to south. |
| Specie | Aythya marila |
| Current European Distribution | Winters in coasts of north Europe, Adriatic and Black sea, also in points of central Europe. |
| Areas of Irruption | Irruptive in the whole Mediterranean region, except in Adriatic coasts. |
| Direction of Movements | Movements to south. |
| Specie | Melanitta fusca |
| Current European Distribution | Winters in Atlantic coasts of Europe, in north-west Mediterranean coasts and in Black sea. |
| Areas of Irruption | Irruptive in the whole Mediterranean region, except in north Adriatic coast, and from north Italy west to coasts of northern half of Iberia. |
| Direction of Movements | Movements to south. |
| Specie | Melanitta nigra |
| Current European Distribution | Winters in Atlantic and Baltic coasts from Skandinavia to Mauritania, also in coasts of Iberia, south France and north Italy. |
| Areas of Irruption | Irruptive in the islands and south Italy to east. |
| Direction of Movements | Movements to south and west. |
| Specie | Bucephala clangula |
| Current European Distribution | Winters in north Europe, inland points, Balkan peninsula, Anatolia, Azure and Adriatic coasts. |
| Areas of Irruption | Irruptive species in the islands, Iberia and Greece. |
| Direction of Movements | Movements to south and west. |
| Specie | Somateria molissima |
| Current European Distribution | Resident and wintering in Atlantic coasts of north and central Europe. |
| Areas of Irruption | Irruptive in the whole Mediterranean region. |
| Direction of Movements | Movements to south. |
| Specie | Clangua hyemalis |
| Current European Distribution | Winters in coasts of north, south France and north Italy. |
| Areas of Irruption | Irruptive in the whole Mediterranean region, except in north Italy and south France. |
| Direction of Movements | Movements to south. |
| Specie | Mergus merganser |
| Current European Distribution | Winters in north Europe, inland points, Adriatic coasts, northern Balkan peninsula and points of Anatolia. |
| Areas of Irruption | Irruptive in west Mediterranean area, islands, southern Balkanic peninsula and Anatolia. |
| Direction of Movements | Movements to south and west. |
| Specie | Mergus albellus |
| Current European Distribution | Winters in coasts of central and western Europe, inland points, Adriatic coasts, northern Balkan peninsula and points of Anatolia. |
| Areas of Irruption | Irruptive in west Mediterranean area, islands, southern Balkanic peninsula. |
| Direction of Movements | Movements to south and west. |
| Specie | Buteo lagopus |
| Current European Distribution | Winters in north, central and east Europe, in Balkan peninsula and points of northern Anatolia. |
| Areas of Irruption | Irruptive in west Mediterranean area, islands, southern Balkanic peninsula and Anatolia. |
| Direction of Movements | Movements to south and west. |
| Specie | Gallinago media |
| Current European Distribution | Breeds in North and Eastern Europe. |
| Areas of Irruption | Irruptive in the whole Mediterranean region. |
| Direction of Movements | Movements to south and west. |
| Specie | Nyctea scandiaca |
| Current European Distribution | North Eurasia. |
| Areas of Irruption | Irruptive in the whole Mediterranean region. |
| Direction of Movements | Movements to south. |
| Specie | Surnia ulula |
| Current European Distribution | Resident in northern Eurasia. |
| Areas of Irruption | Irruptive in the whole Mediterranean region. |
| Direction of Movements | Movements to south. |
| Specie | Aegolius funereus |
| Current European Distribution | Resident in mountains and in the taiga. |
| Areas of Irruption | Irruptive in the islands, Iberia, Italy and southern Anatolia. |
| Direction of Movements | Movements to south and west. |
| Specie | Eremophila alpestris |
| Current European Distribution | Resident in Balkan peninsula, Anatolia and Atlas. Wintering species in northern and eastern Europe. |
| Areas of Irruption | Irruptive in islands and western Mediterranean region. |
| Direction of Movements | Movements to south and west. |
| Specie | Carpodacus erythrinus |
| Current European Distribution | Breeding in north and east Europe, in points of central Europe and Anatolia. |
| Areas of Irruption | Irruptive in the whole Mediterranean region except in Anatolia. |
| Direction of Movements | Movements to south and west. |
| Specie | Plectrophenax nivalis |
| Current European Distribution | Winter species in north, centre and east of Europe. Also in Atlantic coasts of France and northern coasts of Iberia. |
| Areas of Irruption | Irruptive in the whole Mediterranean region. |
| Direction of Movements | Movements to south. |
| Specie | Pinicola enucleator |
| Current European Distribution | Resident and wintering species from Scandinavia to the east. |
| Areas of Irruption | Irruptive in the whole Mediterranean region. |
| Direction of Movements | Movements to south. |
| Specie | Loxia pytyopsittacus |
| Current European Distribution | Resident in Scandinavia and in the north of Eastern Europe. |
| Areas of Irruption | Irruptive in the whole Mediterranean region. |
| Direction of Movements | Movements to south. |
| Specie | Loxia leucoptera |
| Current European Distribution | Resident in the taiga belt, from Eastern Europe to the east. |
| Areas of Irruption | Irruptive in the whole Mediterranean region. |
| Direction of Movements | Movements to south. |
| Specie | Pyrrhula pyrrhula |
| Current European Distribution | Resident and wintering species in the whole of Europe, except two southern thirds of Iberia, south of Greece, south of Anatolia and in the mediterranean islands. |
| Areas of Irruption | Irruptive in the islands and southern Iberia, Greece and Anatolia. |
| Direction of Movements | Movements to south. |
| Specie | Bombycilla garrulus |
| Current European Distribution | Winters in North, centre and east of Europe. Reaches the Balkan mountains. |
| Areas of Irruption | Irruptive in the whole region, except in the north of the Balkanic peninsula. |
| Direction of Movements | Movements to south. |
| Specie | Carduelis flammea |
| Current European Distribution | Resident in north and central Europe. Winters from France to the east. |
| Areas of Irruption | Irruptive in the whole Mediterranean region, except in north Iberia, France and north Italy. |
| Direction of Movements | Movements to south. |
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Appendix 3 – Nowadays surviving species in the east part of the MR, with fossil record in the west Mediterranean area during the Quaternary. Current distributions after Cramp (1998).
| Specie | Anser erythropus |
| Current European distribution | Breeding species in north Eurasia. It winters in Balkan peninsula and coast of Black sea. |
| Highlighted in Appendix 1 when appeared in localities… | from Italy west to Iberia. |
| Specie | Haliaeetus albicilla |
| Current European distribution | Resident and wintering species in north and east Europe, in zones of the Balkans and Anatolia. |
| Highlighted in Appendix 1 when appeared in localities… | from north Italy (also Corsica and Sardinia) west to Iberia. |
| Specie | Circus macrourus |
| Current European distribution | Breeds north Black sea. Winters in east Mediterranean region. |
| Highlighted in Appendix 1 when appeared in localities… | from north Italy (also Corsica and Sardinia) west to Iberia. |
| Specie | Luscinia luscinia |
| Current European distribution | Breeds in eastern Europe to east. Vagrant in Balkans and Anatolia. |
| Highlighted in Appendix 1 when appeared in localities… | from the west of the MR. |
| Specie | Emberiza melanocephala |
| Current European distribution | Breeds in Italy, Balkan peninsula and Anatolia. |
| Highlighted in Appendix 1 when appeared in localities… | from western islands (from Corsica and Sardinia), southern France and Iberia. |
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