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NUMBER 89 71<br />

Mallorca, probably in prehuman levels, and in the late Pleistocene<br />

from Corsica-Sardinia (Alcover et al., 1992). Their<br />

presence is probably related to the presence of small mammals<br />

on these islands {Hypnomys and Nesiotites in the Gymnesic Islands,<br />

Nesiotites, Rhagamys, Microtus, and Prolagus in Corsica-Sardinia),<br />

whereas small mammals are absent from Eivissa.<br />

It is not possible, however, to infer in a strict sense the absence<br />

of Tytonidae on islands without small mammals. For example,<br />

in the Galapagos and Canary islands different small mammals<br />

have been recorded on some of the islands but not on all of<br />

them. In these archipelagos, fytonids have even been obtained<br />

in prehuman deposits on islands without small mammals<br />

(Steadman, 1986; Alcover, unpublished).<br />

Strigid owls that prey on vertebrates are unknown in the fossil<br />

record of Mallorca. Although Otus scops and Athene noctua<br />

do occur in the fossil record, these species feed mainly on insects.<br />

Asio flammeus and Bubo insular is are present in the fossil<br />

record on Sardinia, and Bubo insularis, Asio otus, and a<br />

new, undescribed species of Athene have been found as fossils<br />

on Corsica (Mourer-Chauvire, pers. comm., 1996). Several remains<br />

of Asio flammeus and those of a single individual of a<br />

large species of Strigidae, probably Bubo bubo, have been<br />

found in the late Pleistocene of Eivissa (Alcover et al., 1992;<br />

Sondaar et al., 1995). Asio flammeus, the more abundant strigid<br />

on Eivissa, is more diurnally active than other Palearctic species<br />

of owls of the same size, like Asio otus or Strix aluco<br />

(Cramp, 1985). This behavioral pattern may have been advantageous<br />

for preying on birds, which were the primary food<br />

available on Eivissa before human colonization. Circadian<br />

rhythms might differ between insular and continental systems<br />

(Granjon and Cheylan, 1990, 1993, gave an example of change<br />

of circadian rhythms in a mammalian species). Asio flammeus<br />

may have been more diurnal on Eivissa than in other areas of<br />

its range.<br />

The endemic genus Grallistrix lived in the Hawaiian Islands<br />

(Olson and James, 1991). This genus shows anatomical specializations<br />

for bird-catching (elongated legs and shortened<br />

wings). Its ornithophagous habits have been confirmed by fossil<br />

pellets (Olson and James, 1991). The main activity period of<br />

Grallistrix is unknown, but its ability to catch and kill birds in<br />

flight suggests that the species of this genus may have been<br />

more diurnal than other Strigidae.<br />

Hawks and falcons are widely distributed on western Mediterranean<br />

islands and have been found as fossils on all those islands<br />

studied herein. In the Hawaiian Islands this BTT is occupied<br />

by Circus dossenus, a species convergent with bird-eating<br />

hawks of the genus Accipiter (Olson and James, 1991). On<br />

Mediterranean islands with mammals, species of Buteo (with a<br />

diet of small mammals and small- and medium-sized reptiles),<br />

species of Circus and Milvus (each with a varied diet), and<br />

small species of the genus Falco (mainly herpetophagous and<br />

entomophagous) have been found (Alcover et al., 1992). This<br />

BTT is occupied by Buteo solitarius in the Hawaiian Islands<br />

(Olson and James, 1991), whereas in Eivissa it is represented<br />

by Circus cyaneus, which is scarce in the fossil record, and by<br />

small species of the more abundant Falco (Sondaar et al.,<br />

1995).<br />

Pelagic sea birds are not found on the large western Mediterranean<br />

islands with mammals, being unknown in Corsica and<br />

Sardinia (Alcover et al., 1992). On the other hand, the small island<br />

of Tavolara, near Sardinia, has yielded fossil remains of<br />

Calonectris diomedea and Puffinus yelkouan (Mayaud and<br />

Schaub, 1950). Significantly, Procellariiformes were abundant<br />

in the Pityusic Islands. Many fossil bones of Puffinus mauretanicus<br />

and scarcer remains of Calonectris diomedea and Hydrobates<br />

pelagicus have been found in Eivissa (Sondaar et al.,<br />

1995). Similarly, abundant remains of procellariforms, such as<br />

the extinct, small-sized gadfly petrel Pterodroma jugabilis (Olson<br />

and James, 1991), have been found in Oahu and Hawaii,<br />

and many other Procellariidae are commonly encountered.<br />

The BTT that includes the nonpasseriform species with a<br />

wide diet (malacophagous, insectivorous, detritivorous, and<br />

herbivorous birds) has a better representation on the Pityusic<br />

and Hawaiian islands than on the other western Mediterranean<br />

islands. Two or more flightless species of ibis belonging to the<br />

extinct genus Apteribis lived on the Hawaiian islands Molokai,<br />

Maui, and Lanai, which constituted the Pleistocene island of<br />

Maui Nui (Olson and James, 1991). The genera Otis (bustards)<br />

and Grus (cranes) can be considered to be the Palearctic ecological<br />

analogs of these species. Both are common in the Eivissan<br />

fossil record but are not as common on the other western<br />

Mediterranean islands: so far, only a few remains of one specimen<br />

of Grus have been obtained in Mallorca (Mourer-Chauvire<br />

et al., 1975), and a few remains of Otis have been found on<br />

Corsica (Mourer-Chauvire, pers. comm., 1996).<br />

Among the rails, only a few remains of Porzana have been<br />

recorded on Mallorca, and no rails have been found in the Sardinian<br />

fossil record. Four species of rails are infrequently encountered<br />

in the fossil record of Corsica (Alcover et al., 1992),<br />

whereas rails are well represented in the late Pleistocene of<br />

Eivissa as well as on the Hawaiian Islands. Endemic species<br />

have evolved on the Hawaiian Islands and on Evissa but not on<br />

the other western Mediterranean islands. An endemic undescribed<br />

form of Rallus lived on Eivissa (McMinn, in prep.),<br />

whereas the genus Porzana underwent a significant radiation in<br />

the Hawaiian Islands (Olson and James, 1991).<br />

The BTTs occupied by Passeriformes are not as easily defined<br />

as those occupied by nonpasserines. Thus, only three<br />

BTTs were considered for the Passeriformes. Some patterns,<br />

coming mainly from corvids, emerged from our analysis. On<br />

all the islands studied, corvids played an important role. Pyrrhocorax<br />

was present during the late Pleistocene on all the<br />

western Mediterranean islands. At least one small-sized species<br />

of Corvus also was present in Corsica, Sardinia, and Mallorca.<br />

Large-sized species of Corvus are scarce in the fossil record of<br />

Mallorca, Corsica, and Sardinia. In contrast, on Eivissa abundant<br />

remains of a large Corvus, initially identified as C. antecorax<br />

(Florit et al., 1986), but whose identity is now under revision<br />

(McMinn, in prep.), have been recorded. In the<br />

Pleistocene of the Hawaiian Islands three large-sized species of

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