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70 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Nonpasseriforms Passeriforms Vertebrate predators Herbivores Malacophages, insectivores, detritivores, herbivores Omnivores Bir d trophic types TABLE 3.—The main BTTs (Bird Tropic Types) on islands with terrestrial mammals (Corsica, Sardinia, Mallorca) compared with islands without terrestrial mammals (Eivissa, Hawaiian Islands). Taxa in parentheses are known only from very scarce remains. On islands with terrestrial mammals, some of the BTTs are not present because the niches that they represent are practically monopolized by mammals (see shaded boxes). Data are from the authors; Alcover et al., 1992; Olson and James, 1991; and James and Olson, 1991. Superpredators Nocturnal predators Diumal predators Scavengers Sea birds Granivores Fish-eaters large-sized small-sized specialized ornithophagous large-sized small-sized small-sized large-sized generalists Corsica and Sardinia Aquila Bubo, Asio, Tyto alba Accipiter Falco, Milvus, Buteo, Circus Haliaeetus Gypini Procellaridae spp. Cervidae Rodentia, Lagomorpha (Otis) (Rallus) (Porzana) (Gallinula) (Crex) Sturnus, Pyrrhocorax (Corvus corax) several Fringillidae, Passeridae, Emberizidae Islands with terrestrial mammals Aquila Mallorca Tyto alba/ Tyto balearica Accipiter Falco - Gypini - Bovidae Rodentia (Grus) (Porzana) (Sturnus), (Corvus), Pyrrhocorax - several Fringillidae, Passeridae, Emberizidae works (Cramp and Simmons, 1977, 1980; Cramp, 1985; Hoyo et al., 1992, 1994, 1996). Usually, each taxon was referred only to a single BTT, although a small group of species occupied more than one type (e.g., Haliaeetus on Eivissa, which is considered to have been the superpredator on the island but was simultaneously the main fish predator and the main scavenger). The absence of medium-sized herbivorous mammals on Eivissa is related to the presence of an abundant goose, probably derived from Anser erythropus. The tribe Anserini is usually underrepresented, due to taphonomic conditions, in nonanthropogenic cave sites, excepting on some islands. Because of the great number of remains that have been exhumed from speleological sites on Eivissa, the extinct goose must have been very abundant in Pityusic paleornithological communities. This species was probably the main medium-sized herbivorous vertebrate on the island and must be considered to be the key species in understanding Pityusic paleoecosystems. The occupation of the browsing- or grazing-herbivore trophic type by mammals (Bovidae and Cervidae) on all the Eastern Mediterranean islands Aquila Tyto alba, T melitensis, Athene ere tens is, Aegolius, (Ketupa) (A. flammeus) Accipiter Buteo Haliaeetus Gypini (Puffinus) Cervidae, Elephantidae, Anser/Branta Hippopotamidae Rodentia (Otis) Grus melitensis (Porzana) (Gallinula) (Fulica) (Sturnus), Corvus, Pyrrhocorax, Garrulus (Corvus corax) several Fringillidae and Emberizidae Islands without terrestrial mammals Haliaeetus Asio flammeus Accipiter Falco (Circus) Eivissa Hawaiian Islands Haliaeetus, Pandion Haliaeetus Procellaridae spp. - Otis, Grus Rallus, Crex Pyrrhocorax Corvus several Fringillidae, Passeridae, Emberizidae Haliaeetus Grallistrix Circus dossenus Buteo Haliaeetus Haliaeetus Procellaridae spp. Branta, moa-nalos - Apteribis Porzana Aidemedia Corvus viriosus, Corvus impluviatus, Corvus hawaiiensis several Drepanidini Mediterranean islands has further implications for trophic webs. Thus, the islands with ungulates have eagles of the genus Aquila as superpredators and have vultures {Aegypius, Gyps) as the chief large scavengers. On the other hand, on islands without these mammals but with abundant Anseriformes and Procellariiformes, the role of superpredators is taken by sea-eagles of the genus Haliaeetus, which also must be considered the main scavengers there. This relationship, confirmed on the Mediterranean islands, also applies to other islands in the world (Alcover and McMinn, 1992, 1994). Sea-eagles must have been very abundant in the Pityusic Islands before human colonization. In just one fossiliferous site, remains of at least nine individuals have been found, including several practically complete associated skeletons. On Corsica and Sardinia, on the other hand, very few remains of Haliaeetus have been recovered. As far as vertebrate nocturnal predators are concerned, the absence of Tytonidae in the fossil record of Eivissa, as is the case in the Hawaiian Islands, is particularly noteworthy. Tytonidae are present in the early Pleistocene and the Holocene of
NUMBER 89 71 Mallorca, probably in prehuman levels, and in the late Pleistocene from Corsica-Sardinia (Alcover et al., 1992). Their presence is probably related to the presence of small mammals on these islands {Hypnomys and Nesiotites in the Gymnesic Islands, Nesiotites, Rhagamys, Microtus, and Prolagus in Corsica-Sardinia), whereas small mammals are absent from Eivissa. It is not possible, however, to infer in a strict sense the absence of Tytonidae on islands without small mammals. For example, in the Galapagos and Canary islands different small mammals have been recorded on some of the islands but not on all of them. In these archipelagos, fytonids have even been obtained in prehuman deposits on islands without small mammals (Steadman, 1986; Alcover, unpublished). Strigid owls that prey on vertebrates are unknown in the fossil record of Mallorca. Although Otus scops and Athene noctua do occur in the fossil record, these species feed mainly on insects. Asio flammeus and Bubo insular is are present in the fossil record on Sardinia, and Bubo insularis, Asio otus, and a new, undescribed species of Athene have been found as fossils on Corsica (Mourer-Chauvire, pers. comm., 1996). Several remains of Asio flammeus and those of a single individual of a large species of Strigidae, probably Bubo bubo, have been found in the late Pleistocene of Eivissa (Alcover et al., 1992; Sondaar et al., 1995). Asio flammeus, the more abundant strigid on Eivissa, is more diurnally active than other Palearctic species of owls of the same size, like Asio otus or Strix aluco (Cramp, 1985). This behavioral pattern may have been advantageous for preying on birds, which were the primary food available on Eivissa before human colonization. Circadian rhythms might differ between insular and continental systems (Granjon and Cheylan, 1990, 1993, gave an example of change of circadian rhythms in a mammalian species). Asio flammeus may have been more diurnal on Eivissa than in other areas of its range. The endemic genus Grallistrix lived in the Hawaiian Islands (Olson and James, 1991). This genus shows anatomical specializations for bird-catching (elongated legs and shortened wings). Its ornithophagous habits have been confirmed by fossil pellets (Olson and James, 1991). The main activity period of Grallistrix is unknown, but its ability to catch and kill birds in flight suggests that the species of this genus may have been more diurnal than other Strigidae. Hawks and falcons are widely distributed on western Mediterranean islands and have been found as fossils on all those islands studied herein. In the Hawaiian Islands this BTT is occupied by Circus dossenus, a species convergent with bird-eating hawks of the genus Accipiter (Olson and James, 1991). On Mediterranean islands with mammals, species of Buteo (with a diet of small mammals and small- and medium-sized reptiles), species of Circus and Milvus (each with a varied diet), and small species of the genus Falco (mainly herpetophagous and entomophagous) have been found (Alcover et al., 1992). This BTT is occupied by Buteo solitarius in the Hawaiian Islands (Olson and James, 1991), whereas in Eivissa it is represented by Circus cyaneus, which is scarce in the fossil record, and by small species of the more abundant Falco (Sondaar et al., 1995). Pelagic sea birds are not found on the large western Mediterranean islands with mammals, being unknown in Corsica and Sardinia (Alcover et al., 1992). On the other hand, the small island of Tavolara, near Sardinia, has yielded fossil remains of Calonectris diomedea and Puffinus yelkouan (Mayaud and Schaub, 1950). Significantly, Procellariiformes were abundant in the Pityusic Islands. Many fossil bones of Puffinus mauretanicus and scarcer remains of Calonectris diomedea and Hydrobates pelagicus have been found in Eivissa (Sondaar et al., 1995). Similarly, abundant remains of procellariforms, such as the extinct, small-sized gadfly petrel Pterodroma jugabilis (Olson and James, 1991), have been found in Oahu and Hawaii, and many other Procellariidae are commonly encountered. The BTT that includes the nonpasseriform species with a wide diet (malacophagous, insectivorous, detritivorous, and herbivorous birds) has a better representation on the Pityusic and Hawaiian islands than on the other western Mediterranean islands. Two or more flightless species of ibis belonging to the extinct genus Apteribis lived on the Hawaiian islands Molokai, Maui, and Lanai, which constituted the Pleistocene island of Maui Nui (Olson and James, 1991). The genera Otis (bustards) and Grus (cranes) can be considered to be the Palearctic ecological analogs of these species. Both are common in the Eivissan fossil record but are not as common on the other western Mediterranean islands: so far, only a few remains of one specimen of Grus have been obtained in Mallorca (Mourer-Chauvire et al., 1975), and a few remains of Otis have been found on Corsica (Mourer-Chauvire, pers. comm., 1996). Among the rails, only a few remains of Porzana have been recorded on Mallorca, and no rails have been found in the Sardinian fossil record. Four species of rails are infrequently encountered in the fossil record of Corsica (Alcover et al., 1992), whereas rails are well represented in the late Pleistocene of Eivissa as well as on the Hawaiian Islands. Endemic species have evolved on the Hawaiian Islands and on Evissa but not on the other western Mediterranean islands. An endemic undescribed form of Rallus lived on Eivissa (McMinn, in prep.), whereas the genus Porzana underwent a significant radiation in the Hawaiian Islands (Olson and James, 1991). The BTTs occupied by Passeriformes are not as easily defined as those occupied by nonpasserines. Thus, only three BTTs were considered for the Passeriformes. Some patterns, coming mainly from corvids, emerged from our analysis. On all the islands studied, corvids played an important role. Pyrrhocorax was present during the late Pleistocene on all the western Mediterranean islands. At least one small-sized species of Corvus also was present in Corsica, Sardinia, and Mallorca. Large-sized species of Corvus are scarce in the fossil record of Mallorca, Corsica, and Sardinia. In contrast, on Eivissa abundant remains of a large Corvus, initially identified as C. antecorax (Florit et al., 1986), but whose identity is now under revision (McMinn, in prep.), have been recorded. In the Pleistocene of the Hawaiian Islands three large-sized species of
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SMITHSONIAN CONTRIBUTIONS TO PALEOB
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ABSTRACT Olson, Storrs L., editor.
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EARLY WATERFOWL (ANSERIFORMES) AND
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v m SMITHSONIAN CONTRIBUTIONS TO PA
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localities, all of them situated in
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Bone of Sus scrofa Linnaeus, introd
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duboisi are larger than the largest
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8 iver (1897) but afterward were tr
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10 SMITHSONIAN CONTRIBUTIONS TO PAL
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16 Cor. Hum. Uln. Cpm. Fern. Tbt. T
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Page 54 and 55: Site 13 Research Base ENTRANCE Lowe
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120 SMITHSONIAN CONTRIBUTIONS TO PA
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The Middle Pleistocene Avifauna of
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NUMBER 89 Accordi, B. 1962. La grot
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130 FIGURE 1.—Map showing locatio
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132 Cranium Mandibula Scapula Clavi
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Seabirds and Late Pleistocene Marin
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NUMBER 89 141 METHODS Only strictly
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NUMBER 89 143 FIGURE 2.—Area of s
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NUMBER 89 151 FIGURE 10.—Area of
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NUMBER 89 153 FIGURE 12.—Area of
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NUMBER 89 155 the period studied. T
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NUMBER 89 157 Walker, C.A., G.M. Wr
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164 Vl 620 M 570 £ 520 S 470f •
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166 birds, such as the two species
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170 cional Autonoma de Mexico, for
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174 ated with this specimen, see Mi
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The Fossil Record of Condors (Cicon
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NUMBER 89 179 FIGURE 2.—Geographi
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NUMBER 89 181 FIGURE 5.—Vulturida
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NUMBER 89 183 FIGURE 7.—Referred
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Two New Fossil Eagles from the Late
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NUMBER 89 187 TABLE 1.—Measuremen
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NUMBER 89 189 carpal trochlea relat
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NUMBER 89 191 FIGURE 4.—Holotypic
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NUMBER 89 193 We compared the parat
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NUMBER 89 195 FIGURE 6.—Distribut
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NUMBER 89 197 the Florida State Mus
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A New Genus of Dwarf Megapode (Gall
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NUMBER 89 201 lis hypotarsi along t
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NUMBER 89 203 The fossil is larger
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NUMBER 89 205 Clark, George A., Jr.
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A New Genus and Species of the Fami
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NUMBER 89 209 son with other known
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NUMBER 89 211 FIGURE 1.—Argornis
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NUMBER 89 213 AM AL AM AL AM AL AM
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NUMBER 89 215 caput humeri perpendi
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Selmes absurdipes, New Genus, New S
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NUMBER 89 219 FIGURE 2.—Selmes ab
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NUMBER 89 221 Costae: Deformed frag
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A Fossil Screamer (Anseriformes: An
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NUMBER 89 FIGURE 3.—Chaunoides an
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NUMBER 89 227 B C D FIGURE 6.—The
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NUMBER 89 229 FIGURE 9.—Right tib
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The Anseriform Relationships of Ana
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NUMBER 89 233 Subfamily ANATALAVINA
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NUMBER 89 235 mal was found under t
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NUMBER 89 237 tion, with retroartic
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NUMBER 89 FIGURE 7.—Sternum and p
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NUMBER 89 241 der. The bone is very
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NUMBER 89 243 Eocene records of the
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Presbyornis isoni and Other Late Pa
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NUMBER 89 255 FIGURE 1.—Referred
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NUMBER 89 257 vical vertebrae of th
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NUMBER 89 259 (Olson and Parris, 19
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274 America. Science, 214(4526): 12
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276 concerning the amphicoelous str
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278 ment of the radius that are con
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280 The left coracoid is represente
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284 Kurochkin, E.N. 1982. [New Orde
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288 Palaeontological Institute. [Sp
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290 1991). In this paper we use a "
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Implantation and Replacement of Bir
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NUMBER 89 297 saurs (Figure 1E-G).
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NUMBER 89 299 would seem unlikely a
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Humeral Rotation and Wrist Supinati
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NUMBER 89 303 apparent. It is now c
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NUMBER 89 305 FIGURE 3.—Dorsal vi
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NUMBER 89 307 FIGURE 4.—Wing of t
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NUMBER 89 309 Jura-Museums, Eichsta
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Early Evolution of Birds: Roundtabl
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338 evolved independently twice." M
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