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58 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY<br />

COMPARISONS WITH OTHER ISLANDS<br />

The 20 species of seabirds recorded as fossils from Amsterdam<br />

Island do not include Phoebetria fusca or Sterna vittata,<br />

which currently breed there, suggesting that up to 22 species<br />

had breeding populations. Species that are very rare in the fossil<br />

record, however, such as Pachyptila desolata, Puffinus carneipes,<br />

and P. griseus (each with one individual), are likely to<br />

have been nonbreeding vagrants or the result of skua kills, so a<br />

more realistic estimate is that no more than 19 species of seabirds<br />

bred on Amsterdam Island in the recent past.<br />

Amsterdam Island is similar to the Crozet and Kerguelen islands<br />

in that most breeding species are seabirds; however, with<br />

at least 34 and 30 breeding species of seabirds, respectively,<br />

these subantarctic island groups have far richer communities<br />

than Amsterdam did (Jouventin et al., 1984). Although the fossil<br />

faunas of these islands have not been studied, it is likely that<br />

neither island group has suffered species extinctions as happened<br />

on Amsterdam because some islands in each group have<br />

remained free of predators (Jouventin et al., 1984).<br />

The Mascarene islands Mauritius and Rodriguez are well<br />

known for their extinct, endemic land birds (Newton, 1888;<br />

Hachisuka, 1953; Gill, 1967), but the fossil seabirds have<br />

largely been ignored (Bourne, 1968). These islands are much<br />

farther north than Amsterdam, and various tems and boobies<br />

dominate the fauna, with a few species of petrel present.<br />

Bourne (1968) reported Wedge-tailed Shearwaters {Puffinus<br />

pacificus), a larger shearwater (possibly P. carneipes), and the<br />

Mascarene Petrel {Pterodroma aterrima) in addition to numerous<br />

remains of the White-tailed Tropic Bird {Phaethon lepturus)<br />

in a small collection of fossils from Rodriguez. The discovery<br />

of bones of Barau's Petrel {Pterodroma baraui) on<br />

Amsterdam extends the range of this species, previously<br />

known only from Reunion, and provides a faunal link with the<br />

Mascarene Islands. Pterodroma aterrima, also known only<br />

from Reunion Island, probably bred on Rodriguez (Bourne,<br />

1968), and it may also have reached Amsterdam occasionally.<br />

Studies of the fossil avifauna of St. Helena Island in the Atlantic<br />

Ocean (16°S) have revealed remains of 21 species,<br />

among which the greatest loss of species and individuals was<br />

five of the six resident petrels, three of which were endemic<br />

(Olson, 1975). Farther north in the Atlantic, the fossil avifauna<br />

of the eastern Canary Islands, which lie off the African coast<br />

between about 28°N and 29°N, is dominated by remains of two<br />

extinct species of Puffinus (Alcover and McMinn, 1995).<br />

In the tropical Pacific Ocean the fossil faunas of numerous<br />

islands have been studied, and all are rich in terrestrial species:<br />

seabirds are dominated by procellariids, but toward the equator,<br />

the species diversity of tems, boobies, tropicbirds, and frigatebirds<br />

often equals or exceeds that of procellariids (Olson<br />

and James, 1991; Steadman, 1995). On every island studied,<br />

several species of land birds and populations of seabirds were<br />

exterminated following the arrival of humans in the last few<br />

thousand years (Steadman, 1989, 1995; Olson and James,<br />

1991). Often losses, particularly of land birds, have exceeded<br />

50% of the original species diversity, and, among seabirds,<br />

some of the greatest losses have been petrels and shearwaters,<br />

especially in eastern Polynesia (Steadman, 1989).<br />

In the South Pacific the fossil faunas of the North and South<br />

islands of New Zealand have been studied extensively: Millener<br />

(1990) listed 34 species that became extinct in the late Holocene<br />

following the arrival of humans, to which Scarlett's<br />

Shearwater {Puffinus spelaeus), South Island Adzebill {Aptornis<br />

defossor), Bush Wren {Xenicus longipes), and <strong>Lo</strong>ng-billed<br />

Wren {Dendroscansor decurvirostris) should be added, for a<br />

total of 38 species. In the New Zealand region, seabirds have<br />

not suffered the same degree of loss as elsewhere because, although<br />

numerous populations of seabirds have been extirpated<br />

from the main islands, colonies on offshore islands have ensured<br />

their species survival, with the exception of Puffinus<br />

spelaeus (Holdaway and Worthy, 1994). The fossil faunas of<br />

the New Zealand subantarctic islands have not been studied.<br />

Conclusions<br />

Like most islands where numerous extinctions followed the<br />

arrival of humans and commensal mammals, Amsterdam Island<br />

had a naive avifauna (Milberg and Tyrberg, 1993) unable<br />

to cope with predation by people and rats. Its location far from<br />

other landmasses was no doubt responsible for the paucity of<br />

land-bird species, and its relatively southern position resulted<br />

in a seabird fauna composed mainly of petrels. Studies of fossil<br />

faunas from islands throughout the world, reviewed above,<br />

show that petrels are particularly susceptible to the effects of<br />

humans and to predation by introduced commensal mammals.<br />

Amsterdam Island conforms to this generality in that only 10 of<br />

at least 19 species that formerly bred there survive; in other<br />

words, 47% of the original species have been extirpated. Five<br />

of these 10 surviving species, however, are critically endangered<br />

on Amsterdam. Only three species are endemic to Amsterdam<br />

and St. Paul, and of these the land bird Anas marecula<br />

is extinct, whereas Diomedea amsterdamensis and Pachyptila<br />

macgillivrayi survive.

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