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140 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY FIGURE 1.—The reconstructed "modem range" of Pinguinus impennis in the East Atlantic used to determine areas of sympatry. (•=Holocene subfossil record, A=historical breeding site, x=historical nonbreeding record, ?=occurrence uncertain.) Unfortunately, despite the fact that well over 1000 sites with late Pleistocene avifaunas are known from the West Palearctic, obligate seabirds occur in only a very small proportion of these sites. The main reason for this is the eustatic lowering of sea levels during glacial periods, which means that coastal sites of glacial age are now mostly submerged. Exceptions to this rule are mostly found on steep Mediterranean coasts and in northern Europe (e.g., Norway), where the isostatic rebound of formerly glaciated areas have kept pace with the eustatic rise of the sea level. In one case (Archi, in Calabria, Italy), a glacial coastal site has been preserved through tectonic movements (Ascenzi and Segre, 1971a, 1971b). In principle, interglacial coastal sites should be accessible, but avifaunas of interglacial age are unfortunately extremely rare. The reasons for this are not well understood, but extensive erosion and weathering (near the end of interglacials?), which have destroyed most interglacial cave deposits, are presumably at least a partial explanation.
NUMBER 89 141 METHODS Only strictly marine species are considered in this study to ensure that the occurrence of the birds truly reflects marine conditions. For birds that regularly frequent freshwater or land habitats, occurrence at a site might indicate that suitable freshwater or terrestrial, rather than marine, habitat existed in the vicinity. The species used herein are all Procellariidae, all Hydrobatidae, Moms bassanus, Phalacrocorax aristotelis, Somateria mollissima, Catharacta skua, Larus audouinii, Rissa tridactyla, Pagophila eburnea, and all Alcidae (including Pinguinus). Nomenclature for species' binomials and English names of modern birds follows Sibley and Monroe (1990). With two exceptions, these species are never found inland except as rare vagrants. The exceptions are Somateria mollissima and Rissa tridactyla, which regularly fly over land on migration but rarely stop at inland sites. Xema sabini also might have been included, but it has not yet been recorded from the Pleistocene. Larus hyperboreus has not been included because it can probably not be reliably separated from Larus marinus on osteological criteria. Puffinus puffinus and P. yelkouan have been treated as one species because they were not considered separate species at the time when most of the determinations were made. Only late Pleistocene sites where at least two marine species were reported are included in this study. Data on relevant avifaunas and their dates were obtained through an extensive Site Cyprus Akrotiri Aetokremnos Italy Archi Arene Candide Buca del Bersaglieri Cala Genovesi a Levanzo Grotta dei Fanciulli Grotta Pietro Tampoia Grotta Romanelli Norway Blomvag Skjonghelleren Portugal Grotte de Fuminha Gruta de Figueira Brava Spain Cova den Jaume Orat Cova Nova Cueva de Nerja Devil's Tower Es Pouas Gorham's Cave Great Britain Bacon Hole Creag nan Uamh Paviland Cave Potter's Cave TABLE 1.—Sites with late Pleistocene seabird faunas. Mourer-Chauvire, in litt., 1996 search of the literature. This yielded a total of 22 sites and 31 faunas that fulfill these criteria (Tables 1, 2). At stratified sites, each layer has been considered as a separate fauna. Only late Pleistocene records have been considered, both because dating of older records is usually quite uncertain and because the ecology of the birds may have changed over a longer time interval. For modern distributions, the maps in Cramp (1977, 1983, 1985) were used. These are admittedly only approximate in offshore areas, but because the fossil record necessarily samples the coastal fauna, the nearshore distribution, which is better known, is more significant. It should be noted that the ranges used are the total ranges of the species in question, breeding ranges usually being considerably more restricted. It is, however, only rarely possible to determine whether a fossil is from a breeding bird or not. It should be noted that the ranges in Cramp only define the main area of distribution and that most species are found more or less regularly in small numbers well outside the indicated range. This, of course, introduces a margin of error, but the probability of a rare or vagrant species being preserved as a fossil is certainly very low. For the recently extinct Pinguinus impennis, a "modern" distribution map was compiled from literary and subfossil data (Figure 1). Faunas The faunas (Table 1) are treated in approximately chronological order below. Sources Ascenzi and Segre, 1971a, 1971b; Cassoli and Segre, 1985 Cassoli, 1980 Lambrecht, 1933; Wolf 1938 Cassoli and Tagliacozzo, 1982 Del Campana, 1946 Lambrecht, 1933; Mayaud and Schaub, 1950; Newton, 1922; Wolf, 1938 Cassoli et al., 1979 Lie, 1986; Undas, 1942 Larsen, 1984; Larsen et al., 1987 Lambrecht, 1933; Roche, 1972; Villalta, 1964 Mourer-Chauvire and Antunes, 1991; Mourer Chauvire, in litt., 1995 McMinn etal., 1993 Florit and Alcover, 1987; McMinn and Alcover, 1992 Boessneck and von den Driesch, 1980; Eastham, 1986, 1988, 1989; Hernandez, 1993, 1994, 1995, and in litt. Garrodetal., 1928; Villalta, 1964 Alcover et al., 1981, 1992; Florit etal., 1989 Eastham, 1968, 1989; Vega Toscano, 1990 (dating only) Harrison, 1977, 1987; Stringer et al., 1986 Newton, 1917; Lambrecht, 1933; Wolf, 1938; Stuart, 1983 (dating only) Bell, 1922; Bowen, 1970 (dating only) David, 1991
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140<br />
SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY<br />
FIGURE 1.—The reconstructed "modem range" of Pinguinus impennis in the East Atlantic used to determine<br />
areas of sympatry. (•=Holocene subfossil record, A=historical breeding site, x=historical nonbreeding record,<br />
?=occurrence uncertain.)<br />
Unfortunately, despite the fact that well over 1000 sites<br />
with late Pleistocene avifaunas are known from the West<br />
Palearctic, obligate seabirds occur in only a very small proportion<br />
of these sites. The main reason for this is the eustatic<br />
lowering of sea levels during glacial periods, which means<br />
that coastal sites of glacial age are now mostly submerged.<br />
Exceptions to this rule are mostly found on steep Mediterranean<br />
coasts and in northern Europe (e.g., Norway), where the<br />
isostatic rebound of formerly glaciated areas have kept pace<br />
with the eustatic rise of the sea level. In one case (Archi, in<br />
Calabria, Italy), a glacial coastal site has been preserved<br />
through tectonic movements (Ascenzi and Segre, 1971a,<br />
1971b). In principle, interglacial coastal sites should be accessible,<br />
but avifaunas of interglacial age are unfortunately extremely<br />
rare. The reasons for this are not well understood, but<br />
extensive erosion and weathering (near the end of interglacials?),<br />
which have destroyed most interglacial cave deposits,<br />
are presumably at least a partial explanation.
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