14 BIRD BONES FROM THE TO'AGA SITE: PREHISTORIC LOSS OF SEABIRDS AND MEGAPODES DAVID W. STEADMAN INTRODUCTION A S PART OF A LONG-TERM program to reconstrUct the natural distribution and diversity of bird- life in the South Pacific (figure 14.1), I have sought bird bones from archaeological sites in many different archipelagos. Until recently, the islands of Samoa have not been represented in this data base. In 1987, T. L. Hunt and P. V. Kirch conducted excavations at the To'aga site (AS-13-1), Ofu Island, American Samoa yielding a small sample of bird bones representng at least six taxa (Steadman 1990). This first glimpse of the prehistoric avifauna of Samoa showed that at least two species (a shearwa- ter and petrel) had been lost on Ofu since the first anrival of humans more than 3000 years ago. Hunt and Kirch expanded the excavations at To'aga in 1989. This paper reports the entire bird bone assemblage from both the 1987 and 1989 field seasons. The sample of bird bones from To'aga now consists of at least fifteen taxa (table 14.1) and provides a more thorough, although still far from complete, portrayal of the birdlife of ancient Samoa. The bird bones from To'aga have been cata- logued in the University of Washington Burke Museum (UWBM) Fossil Bird Collection. Com- parative skeletal or oological specimens are from UWBM, the British Museum (Natural History) (BMNH), the New York State Museum (NYSM), and the United States National Museum of Natural History, Smithsonian Institution (USNM). hi the species accounts that follow, "Unit" refers to meter square excavations (designated with Arabic numer- als). Roman numerals refer to stratigraphic layers. For details of the stratigraphy, chronology, material culture, and non-bird faunal assemblages of the To'aga site, see Hunt and Kirch (1988), Kiivh et al. (1989, 1990), and various chapters in this volume. Unless stated otherwise, the modem distribu- dons of species within American Samoa are taken from the excellent surveys conducted in 1975-1976 by Amerson et al. (1982ab) and in 1986 by Engbring and Ramsey (1989). Modem distributions for elsewhere in Polynesia are taken from Pratt et al. (1987). The prehistoric records are from Steadman (1989a) and from unpublished data from my recent research; islands preceded by an asterisk (*) repre- sent extirpated populations. SPECIES ACCOUNTS Order Procellariiformes Family Procellanidae Pwffinuspacificus (Wedge-tailed Shearwater) MATERIAL. Humerus (UWBM 1680), T9/ 500E, Unit 21, fiB. Five ulnae (UWBM 1244, 1251, 218 The To'aga Site Table 14.1 Bird Bones from the To'aga Site Number of Identified Bones Taxa Seabirds *Puffinus pacificus *Puffnus iherminieri *Puffinus griseus *Pterodroma rostrata *Pterodroma sp., size of P. heraldica (*)Procellariidae sp. *Sula sula Fregata sp. Anous stolidus Gygis alba Steminae sp. Landbirds Egretta sacra Numenius tahitiensis (M) Gallus gallus (I) *Megapodius sp. Gallicolumba stairii TOTAL Total Seabirds (species/bones) Total Landbirds (species/bones) Total Native Landbirds, without IM (species/bones) % of Bones from Extirpated Species (without IM) 11 2 15 6 2 9 1 2 1 1 1 1 1 16 2 3 15n4 10/51 5/23 4/7 85% I=Introduced species M=Migrant Species * Extirpated on Ofu (*) represents extirpated species, but not necessarily different from those already listed. 1256, 1678, 1679), Unit 9, IIB; Unit 5, IIC; Unit 14, llIa,4; 19/500E, Unit 21, IIB (2 bones). Radius (UWBM 1664), T9/500E, Unit 20, IIIB. Carpometacarpus (UWBM 1630), T3I200W. Unit 27, fI1A. Tibiotarsus (UWBM 1642), T5/100E, Unit 28, HIB. Two pedal phalanges (UWBM 1246, 1248), Unit 4, IIB; Unit 1, IIB. REMARKS. This tropical shearwater is rarely noted at sea today in American Samoa It may nest on Pola Islet (off Tutuila) and on Ta'u, but this has not been confirmed. There are no previous rcords from Ofu. It is widespread in Polynesia today, although nesting islands are few. Other Polynesian archaeological records of PwJnuspac~lcus are from *Ua Huka and *Tahuata (Marquesas), *HuAhine (Society Islands), *Lifuka and * Eua (Tonga), and Tikopia and Anuta (Solomon Islands). Puojinus lherminieri (Audubon's Shearwater) MATERIAL. 2 ulnae (UWBM 1651, 1671), T9/500E, Unit 23, IIB; T9/500E, Unit 21, IIB. REMARKS. Within American Samoa, Puffinus iherminueri nests only on Tau, where at least 200 birds breed in the cloud forest, and on Tutila (colony size unknown). It is uncommon at sea and has not been recorded previously on Ofu. Like P. pacificus, P. lherminieri nests today on relatively few islands scattered across Polynesia. Other Polynesian archaeological records of P. Iherminieri are from *Nuku Hiva, Ua Pou, *Ua Huka, *Hiva Ok, and *Tahuata (Marquesas), *Huahine (Society Prehistoric Loss of Seabirds and Megapodes 219 I I N o 0C C C,5 J . 4. *0 0 m 0~~~ z~~~~~~~ o w -J ~~~~~~~~~~-J *. * X 0 C)( 00J (4 t ze C s* * 4 0 -0 . a Z z~~~~~ I-.a ~~~~~~~~ .00 : Y 0 - J u0 0 Z ;'= | *oa uot~~~~~~~~~~; J i Z '0 w 0I a -J 4 - 4 - z~~~~~ 0. - z z w Z 9~~~~~~~~~c *r. 220 The Tolaga Site Islands), *Mangaia (Cook Islands), Eua (Tonga), and Tikopia and *Anuta (Solomon Islands). Puffinus griseus (Sooty Shearwater) MATERIAL. Sternum (UWBM 1681), T9/ 500E, Unit 21, IIB. Coracoid (UWBM 1645), T5/ 100E, Unit 29, RIB. Scapula (UWBM 1245), Unit 9, IIB. Three humeri (UWBM 1641, 1643, 1659), T5/100E, Unit 29, 1; T5/100E, Unit 28, IIB; Unit 20, RB. Three ulnae (UWBM 1240, 1241, 1647), Unit 1 1, I-8 (2 bones); T5/100E, Unit 30,1 H. Carpometacarpus (UWBM 1259), Unit 7, hA. Manus digit (UWBM 1260), Unit 7, HA. Pelvis (UWBM 1674), T9/500E, Unit 21, IIB. Three tibiotarsi (UWBM 1252, 1253, 1652), Unit 6, hA (2 bones); T9/500E, Unit 23, IRA. REMARKS. As reported by Steadman (1990), these bones are larger than those of any species of shearwater that resides today in tropical Polynesia. They agree in all osteological details with bones of Puffinus griseus, a species that probably migrates through the Samoan region today (Harrison 1983:260, 420, Pratt et al. 1987:55), although there are no records from American Samoa. The Sooty Shearwater nests today only on islands off New Zealand, southern Australia, and extreme southern South America (Harrison 1983:260, 420). Most of the nesting localities of P. griseus are temperate or subantarctic, although in Australia nesting occurs at least as far north as 320 40' S on the subtropical Broughton Islands. Three possible explanations for the unexpected presence of Paufinus griseus on Ofu were proposed by Steadman (1990): (1) the bones represent migrant birds taken at sea by fishermen; (2) P. griseus once resided on Ofu and, like other procellarids, was extirpated through predation in their nesting burrows by humans and rats; (3) the bones represent an extinct, resident shearwater that differs specifically or subspecifically from modem P. griseiss but is osteologically very similar. Regard- ing the first explanation, I am aware of no ethno- graphic accounts of Polynesians capturing seabirds while fishing. The data now available, particularly that fifteen bones (nine different skeletal elements) are osteologically indistinguishable from modem skeletons of P. griseus, suggest that the second explanation may be corect, even though the oceano- graphic conditions near Ofu are wanmer than those of the wannest localities where P. griseus nests today. That this large shearwater (or an osteologi- cally indistinguishable subspecies [explanation 3 is not necessarily incompatible with explanation 2]) was once resident rather than migrant on Ofu is indicated by the presence at To'aga of two bones of P. griseus (UWBM 1645,1653) that, based upon porosity of the external surface, are from volant juveniles unlikely to have dispersed far from their place of birth. The fonner residency of P. griseus on Ofu is supported furtxr by the abun e of its bones at To'aga (15 of 74, or 20% of identifiable bones). Sooty Shearwaters are represented as well among the few bird bones from a Lapita site on *Niuatoputapu (Tonga), southwest of American Samoa (Steadman 1990). Pterodroma rostrata (Tahiti Petrel) MATERIAL. Two mandibles (UWBM 1250, 1682), Unit 4, RB; T9/500E, Unit 21, IIB. Scapula (UWBM 1684), 19/500E, Unit 21, IIB. Humenrs (UWBM 1657), T9/500E, Unit 20, IIB. Ulna (UWBM 1242), Unit 5, HA. Tarsometatarsus (UWBM 1247), Unit 4, IIB. REMARKS. This large petrel nests today at a few widely scattered localities in te Marquesas Islands, Tahiti, New Caledonia, and American Samoa. Its presence on the latter is based upon a colony of about 500 individuals discovered in 1976 in the montarn cloud forest of Ta'u, a single nesting bird discovered on Tutuila in 1986, and a few birds heard on Olosega in 1986. An onshore record in 1972 from Taveuni (Clunie et al. 1978) suggests th Pterodroma rostrata may nest in Fiji as well. Elsewhere in Polynesia, P. rostrata has been idend- fied from archaeological sites on *Ua Huka and Tahuata (Marquesas), *Huahine (Society Islands), *Aitutaki (Cook Islands), *Lifuka and *Eua (Tonga), Tikopia (Solomon Islands), and New Caledonia. Pterodroma sp. (unknown petrel, size of P. heraldica) MATERIAL Radius (UWBM 1670), T9/500E, Unit 21, RB. Femur (UWBM 1676), Unit 21, RIB. REMARKS. These specimens, although not adequate for species-level identification, represent a species of Pterodroma in the approximate size rae Prehistoric Loss of Seabirds and Megapodes 221 of P. heraldica (Herald Petrel, which often is considered conspecific with P. arminjonana) or perhaps the slightly larger P. exlerna (Juan Fernandez Petrel), both of which are significantly smaller than P. rostrata. In American Samoa, P. eterna is known ordy from uncommon sightings at sea, while P. heraldica nests on TaVu. Regardless of species-level identification, UWBM 1670 and 1676 indicate that a species of petrel other than P. rostrata probably once nested on Ofu. Procelladiidae sp. (unknown petrel/shearwater) MATERIAL. Coracoid (UWBM 1633), T5/ 100E, Unit 16,11. Five ulnae (UWBM 1243,1258, 1634, 1662, 1669), Unit 9, IIB; Unit 5, IIB; T5/ 100E, Unit 16, I; T9/500E, Unit 20, IIIB; T9/500E, Unit 21, fIB. Radius (UWBM 1249), Unit 4, IIB. Two carpometacarpi (UWBM 1239, 1677), Unit 9, UB; T9/500E, Unit 21, IIB. REMARKS. These fragmentary bones cannot be identified beyond the family level. They do not represent a taxon separate from those already identified. Order Pelecaniformes Family Sulidae Sula sula (Red-footed Booby) MATERIAL. Radius (UWBM 1656), T9/500E, Unit 20, IIIC. REMARKS. Sula sula nested on Ofu as recently as 1975-76 (only twenty-five birds), but the small colony no longer existed in 1986. This booby still nested in 1986 on Tutuila and Rose islands. It is widespread in tropical oceans. Other Polynesian archaeological records of S. sula are from *Henderson (Pitcaim Group); *Ua Huka, *Hiva Oa, and *Tahuata (Marquesas); *Huahine (Society Islands); *Aitutaki (Cook Islands); Niuatoputapu (Tonga); and *Tikopia and *Anuta (Solomon Islands). Family Fregatidae Fregata sp. (unknown frigatebird) MATERIAL. Humerus (UWBM 1254), Unit 14, lIa-4. Ulna (UWBM 1675), T9/500E, Unit 21, BB. REMARKS. These specimens cannot be distinguished from those of Fregata minor or F. ariel, both of which visit but do not nest on Ofu today (total of thirteen birds counted in 1975-76; still recorded as a visitor in 1986). The bones of F. minor males and F. ariel females are similar in size and difficult to distinguish from each other (Steadman et al. 1990). Both species of Fregata occur fairly commonly in Polynesian archaeological sites. Order Charadriifonues Family Laridae Anous stolidus (Brown Noddy) MATERIAL. Humerus (UWBM 1668),T9/ 500E, TP 21, IIB. REMARKS. This tern is common and wide- spread today in American Samoa as well as most of Polynesia. The Ofu population was ca. 500 in 1975- 76 and was not accurately estimated in 1986. Other Polynesian archaeological records of Anous stolidus are from Henderson (Pitcairn Group); Ua Pou, Ua Huka, and Tahuata (Marquesas); Huahine (Society Islands); Mangaia and Aitutaki (Cook Islands); Niuatoputapu, Lifuka, and 'Eua (Tonga); 'Upolu (Western Samoa); Tikopia and Anuta (Solomon Islands); and Mussau (Papua New Guinea). Gygis alba (Common Fairy-Tern) MATERIAL. Humerus (UWBM 1631), T5/ 100E, Unit 16,I. REMARKS. This distinctive tern is common and widespread today in most of Polynesia including American Samoa. The Ofu population was ca. 100 in 1975-76, and at least 500 in 1986. Other Polynesian archaeological records of Gygis alba are from Henderson Island (Pitcairn Group), Ua Huka and Tahuata (Marquesas), Huahine (Society Islands), Mangaia (Cook Islands), and Niuatoputapu and 'Eua (Tonga). Sterninae sp. (unknown tern) MATERIAL. Ulna (UWBM 1257), Unit 14, llb-5. REMARKS. This eroded, fragmentary speci- men represents a tern smaller than either of the above species. UWBM 1257 is approximately te 222 The To'aga Site size of the ulna in Sterna swnatrana orAnous minus, both of which occur today in American Samoa, although only the latter nests on Ofu (just ten birds in 1975-76, and perhaps about the same in 1986). Order Clconiifonmes Family Ardeidae Egretta sacra (Pacific Reef-Heron) MATERIAL. Coracoid (UWBM 1655), T9/ 500E, Unit 22, 1. REMARKS. Egreta sacra resides throughout American Samoa and most of Polynesia today. It is uncommon on Ofu, where only dark-phase birds have been recorded. Other archaeological records are from Nuku Hiva and Ua Huka (Marquesas), Huahine (Society Islands), Mangaia and Aitutaki (Cook Islands), and 'Eua (Tonga). Order Charadriiformes Family Scolopacidae Nwnenius tahidensis (Bristle-thighed Curlew) MATERIAL. Corcoid (UWBM 1635, 1636; originally believed to be two separate bones), T5/ 100E, Unit 15, IIID. REMARKS. This widespread but rather uncommon migrant shorebird has not been recorded previously from Ofu, although undoubtedly it still occurs there occasionally. American Samoan records of Nwnenius tahlitensis are confined to Tutuila, Ta'u, Swains, Rose, and Olosega. Other Polynesian archaeological records for N. tahidensis are from Henderson Island (Pitcaim Group), Ua Huka (Marquesas), Huahine (Society Islands), Mangaia and Aitutaki (Cook Islands), and Tikopia (Solomon Islands). During the autumn wing molt, thirteen of twenty-nine individuals of N. tahtidensis captured on Laysan (Leeward Hawaiian Islands) were flightless (Marks et al. 1990). This adaptation would seem to be viable only in a predator-free setting. It may have led to reductions in the distribution and abundance of N. ahitiensis following the human colonization of Polynesia. Order Galliformes Family Megapodiidae Megapodius sp. (Unknown Megapode) MATERIAL. Ulna (UWBM 1637), T5/100E, Unit 15, HID. Femur (UWBM 1649), Unit 30, HID. REMARKS. These two fragmentary specimens (fig. 14.2) are most similar quantitatively and qualitatively to modem specimens of Megapodius freycinet, a widespread species that now reaches its eastern limit in Vanuatu. UWBM 1649 is indistin- guishable qualitatively from the femur in modem M. freycinet, while UWBM 1637 differs from the ulna of M. freycinet in having a slightly deeper sulcus radialis and in lacking a diagonal trough on the cranial surface of tuberculum carpale. Like the archaeological specimens of Megapodiusfreycinet from *Tikopia (Solomon Islands; Steadman et al. 1990), the specimens of Megapodius from To'aga are at the extreme small end of the size range of M.freycinet (table 14.2). Among living subspecies of M.freycinet, there is no indication that individual body size decreases eastward in Oceania. For example, the tarsus (in skins) of the Vanuatu population is not smaller han that from Australia (table 14.3). The bones from To'aga are larger than in M. pritchardi (confined to Niuafo'ou, Tonga) and M. laperouse (found orny in Palau and Mariana Islands), but much smaller than in two extinct species recently described from late Holocene archaeological and paleontological sites- M. molistructor of New Caledonia and Lifuka and M. alimentum of Lifuka and 'Eua (Balouet and Olson 1989; Steadman 1989b, pers. obs.). Megapodius pritchardi is the only species of megapode that survives east of Vanuatu. Both To'aga specimens of Megapodius are from one of the site's deepest and oldest strata, Layer IIID in units 15/29/30. This suggests that megapodes may have been lost from Ofu not long after prehis- toric colonization of the island. These bones provide the first unequivocal, well-documented record of a megapode from Samoa. There is, however, compli- cated historical evidence that a megapode, described as M. stain by Gray (1861), may have existed in tbe mid-i 800s on Upolu or Savai'i (Western Samoa). Gray (1861) also described M. burnabyi from Prehistoric Loss of Seabirds and Megapodes 223 Comparison of megapode bones: ulna in ventral aspect (A, B) and femur in lateral aspect (C, D). A, C. Megapodius cf.freycinet, archaeological specimens (UWBM 1637, 1649), To'aga site, Ofu, American Samoa. B, D. M. freycinet, modem specimen, USNM 557015, male, Halmahera, Northern Moluccas. Scale = 20 mm. Ha'apai (Tonga). The true identities of M. stairi and Mf. burnabyi are uncertain because only a single egg -ver was collected of each, and these two eggs cannot be assigned unequivocally to M. pritchardi, M.freycinet, or any other species of megapode (Steadman 1991). One or both of these eggs may represent the same species as the bones from Ofu, which also are at the very lower limit of the size range of M. freycinet and larger than, or at the uppermost size limit of, M. pritchardi. The original, handwritten data slip with the holotypical egg of Megapodius burnabyi in the British Museum notes that this egg was "called the chief s egg' as they are only allowed to eat them." Such a chiefly tabu might suggest rarity of the bird and a knowledge that overexploitation, which probably was the cause of the rarity, would eventu- ally lead to extinction. Alternatively, being called the "chief's egg" could suggest that megapode eggs were prestigious trade items brought to Ha'apai from another island. An extensive exchange network operated among Fiji, Tonga, and Samoa in late prehistoric and early historic times (Kirch 1984:238- 42; 1988:257-60). The Samoan voyages recorded by Stair (1895) included Fiji and Tonga as well as much of East Polynesia. Bennett (1862:247) noted that the nesting grounds of M. pritchardi on Niuafo'ou were "under the protection of the king or chief, and by his permission only can the birds or eggs be procured." Even if megapodes of the Fiji/Tonga/Samoa region were confined by the nineteenth century to Niuafo'ou, they would have been known to Tongans in Ha'apai, as well as to Samoans. Unless additional evidence comes forth, neither C% A B Figure 14.2 224 The To'aga Site Table 14.2 Measurements (in mm) of the Femur and Ulna of Megapodius, with Mean, Range, and Sample Size Specimen Locality A B C D E Megapodious sp. UWBM 1637 (U) Megapodius sp. UWBM 1649 (U) M. f cf. eremita Lab #: 66, 68 (U) M. f freycinet USNM 556995-557002 557016, 557017 (M) M. f freycinet USNM 557006-557008 557010-557013, 557019-557022 (F) M. f gilberti USNM 226175, 226176 (F, M) M. f cumingi BM(NHl) 1862.2.10.2 (U) M. f nicobariensis USNM 19686, 19700 (M) M. pritchardi tJSNM 319633, 319634 (IJ) Samoa: Ofu: To'aga Site Samoa: Ofu: To'aga Site New Ireland N. Moluccas: Halmahera N. Moluccas: Halmahera Sulawesi Philippines: island unknown Nicobar Islands Tonga: Niuafo'ou M. laperouse USNM unnumbered (U) M. alimenturm UWBM 2100 (IJ) N. Mariana Islands: Rota: Payapai Cave Tonga: 'Eua: 'Anatu (Ground-Dove Cave) 3.4 2.5 1 1 - 6.0 4.4 1 1 F = Female, M = Male, U = Sex unknown Column Headings: A. Femur: width at deepest proximo-lateral muscle scar B. Femur: depth at deepest proximo-lateral muscle scar C. Ulna: minimum width of shaft D. Ulna: minimum depth of shaft E. Ulna: width of distal end 6.5 1 7.7 1 4.4 1 3.3 1 8.4 1 6.8 1 4.9 1 7.3 1 8.2 7.6-9.1 10 8.3 7.7-8.8 10 7.4 7.3-7.6 2 5.0 4.7-5.3 10 4.9 4.6-5.4 10 4.3 4.3 2 3.8 3.3-4.1 10 3.7 3.5-4.1 10 3.2 3.2-3.3 2 7.4 7.0-7.9 10 7.3 7.0-7.8 10 6.5 6.5 2 8.2 7.5-8.7 10 8.2 7.6-8.8 10 7.2 6.9-7.5 2 8.1 1 8.3 8.0-8.6 2 5.6 5.4-5.7 2 8.0 1 5.2 4.1 1 1 7.4 1 8.3 8.2-8.4 2 6.0 5.9-6.1 2 5.0 4.9-5.0 2 3.4 3.3-3.5 2 3.8 3.6-4.0 2 2.8 2.8-2.9 2 7.8 7.6-7.9 2 5.2 5.0-5.3 2 5.4 1 Prehistoric Loss of Seabirds and Megapodes 225 Table 14.3 Tarsal Length (in mm) from Skins of Selected Subspecies of Megapodius freycinet, with Mean, Range, and Sample Size. Based upon Specimens from BM(NH). Subspecies Tarsal Length M. f layardi Vanuatu: Santo, Vate (F, 5M) M. f eremita Solomon Is.: San Cristobal, Guadacanal, Bouganville (3F, 2M, tI M. f eremita Papua New Guinea: New Britain (3F, 2M, 3U) M. f yorki Australia: Queensland (2F, 4M, 4tJ) 73.8 (71-76), 6 72.2 (67-77), 6 67.1 (62-71), 8 71.6 (70-74), 10 F = female, M = male, U = sex unknown Megapodius stairi orM. burnabyi should be re- garded as certain records of indigenous populations of megapodes in nineteenth century Samoa or Ha'apai, although this cannot be ruled out. The survival of M. pritchardi on Niuafo'ou has been due to chiefly control of exploiting eggs and birds at the conspicuous nest mounds, as first described by Bennett (1862). The people of Niuafo'ou must have realized that conserving megapodes, which probably occurred nowhere else in the region, would help to maintain their share of commerce in the Samoa- Tonga-Fiji trade network. Family Phasianidae Gallus gallus (Chicken) MATERIAL. Sternum (UWBM 1261), Unit 11, 1. Coracoid (UWBM 1654), T9/500E, Unit 23, IIIB. Scapula (UWBM 1663), T9/500E, Unit 20, HIB. Ulna (UWBM 1255), Unit 14, Illa-4. Radius (UJWBM 1660), T9/500E, Unit 20, IhB. Two pelves (UWBM 1665, 1686, 1688; the last two originally believed to be separate bones), T9/500E, Unit 20, RIB; T9/500E, Unit 21, IIB. Two femora (UWBM 1666, 1667), T9/500E, Unit 20, hIB. Four tibiotarsi (IJWBM 1632, 1644, 1658, 1661), T5/100E, Unit 16, 1; T5/1OOE, Unit 29, IIIB; T9/500E, Unit 20, IIB; T9/500E, Unit 20, IhB. Two tarsometatarsi (UWBM 1646, 1683), Unit 29, rIB; Unit 21, II. REMARKS. Feral and/or domestic populations of Gallus gallus occur neatly throughout Polynesia, including all inhabited Samoan islands. All chickens recorded on Ofu in 1986 were near human habitation and not from deep within forests. This non-native species has been found thugh virtually all of Polynesia in archaeological sites of any age, except that it is absent from all sites on Henderson Island (Schubel and Steadman 1989). Chicken bones occur throughout the To'aga sequence, but are especially well represented in the Layer IIIB occupation in Units 20/23 along Transect 9: This indicates that G. gallus was a commonly eaten bird during the Ancestral Polynesian phase, ca. 2500 yr B.P. 226 The To'aga Site Order Columbiformes Family Columbidae Gallicolwnba stairii (Shy Ground-Dove) MATERIAL. Humerus (UWBM 1638), T5/ 100E, Unit 15, 111D. Ulna (UWBM 1658), Unit 30, IIID. Tarsometatarsus (UWBM 1690), T5/100E, Unit 29, IIID. REMARKS. This species occurs only in very old deposits at To'aga, primarily Layer IIID in Units 29/30. Gallicolwnba stairii is extremely rare on Ofu today, with a very roughly estimated 100 birds surviving in 1975-76. Only two or three ground- doves were seen on Ofu during the 1986 surveys; no population estimate was made. Within American Samoa, only perhaps on Olosega does another small population of G. stairii survive. Similar declines or losses of populations of G. stairii have occurred in Tonga, where the only other archaeological record of G. stairii is from *'Eua. DISCUSSION Virtually all of the bird bones from To'aga are broken, often with both articular ends missing. Most of these breaks are not fresh, although often it is difficult to distinguish whether human or sedimen- tary processes have caused the breakage. A few bones are rounded, suggesting some post-mortem sedimentary transport. The breakage and rounding might indicate that the calcareous sands at To'aga represent a somewhat higher energy deposit than the calcareous sands at certain other Polynesian archaeo- logical sites, such as Hane (Ua Huka, Marquesas) or Tongoleleka (Lifuka, Tonga). Among the shearwa- ters and petrels, however, a systematic butchering technique is suggested by the fairly consistent pattern of both ends of the humerus, ulna, and tibiotarsus being broken off. Two of the chicken bones had been chewed by rats. None of the bird bones seems to have been modified into recogniz- able artifacts. Among indigenous, resident species recorded from the To'aga site, five of ten seabirds and one of three landbirds are extirpated on Ofu (table 14.1). At least two of the surviving species (Sterninae sp., Gallicolwnba stairit) exist today on Ofu only in very small, threatened populations. Should these species be lost from Ofu, the proportion of bones of extir- pated species at To'aga would increase from 85% (table 14.1) to 93%. The majority of bird bones from To'aga (46 of 74, or 62%) are of at least five species of petrels or shearwaters (table 14.1), none of which nests on OfN today (Amerson et al. 1982a:90). Only two of hse species (Audubon's Shearwater and Tahiti Petrel) are known certainly to nest today anywhere in American Samoa. As seems to be case throughout Polynesia (Steadman 1989a, Dye and Steadman 1990), the island of Ofu had a diverse and probably abundant seabird fauna when humans first arrived. In the case of Ofu, not a single species of procellariid has survived the three millennia of human occupa- tion. Compared to avian assemblages from other Polynesian archaeological sites, the dominance of procellariids at To'aga would characterize a fairly early site, i.e., one that dates to within the first thousand years of human occupation (Dye and Steadman 1990). When compared to sites that seem to represent the initial human occupation of an island, however, such as the Hane site (Ua Huka, Marquesas; Steadman 1989a) orTongoleleka site (Lifuka, Tonga; Steadman 1989b), the To'aga site's lower percentage and diversity of bones from native landbirds and higher percentage of chicken bones would suggest that this site may not represent the first 500 years of human occupation of Ofu. This suggestion is compatible with the radiocarbon chronology at To'aga which indicates occupation of the site from about 2800 to 1900 yr B.P. (see Kirch, chapter 6). Although the bird bones from To'aga provide much new data on the prehistoric distribution of Samoan birds, a bone sample about an order of magnitude larger would be necessary to provide a fairly complete picture of the past birdlife of Ofu. That six of the fifteen taxa of birds from To'aga are represented by only a single bone indicates that more species await discovery if a larger bone sample were available. The 1987 sample of twenty-three bones yielded six taxa, which increased to fifteen taxa with the addition of fifty-one bones from the 1989 excavations. The point of diminishing returns is difficult to determine, however, from comparison with assemblages from other sites. For example, thirty-five species of birds are represented in a sample of ca. 350 bones from the Fa'ahia site (Huahine, Society Islands; Steadman and Pahlavan Prehistoric Loss of Seabirds and Megapodes 227 1992), while thirty-nine species of birds ae repre- sented in a sample of ca. 12,000 bones from the Hane site (Ua Huka, Marquesas) (Steadman 1989a, pers. obs.). Twelve of the species from Hane are known from either one or two bones. Another gauge of the incompleteness of the To'aga avifauna is the low percentage of species in the modem avifauna of Ofu tha are represented at the site. In this case, only two of sixteen possible species of resident landbirds (12.5%) ar represented. The To'aga site has given us an intriguing introduction to Samoa's prehistoric birdlife. Our understanding of the relationship between native birds and the first human inhabitants of Samoa undoubtedly will improve as more early archaeologi- cal sites are discovered and carefully excavated. ACKNOWLEDGEMENTS This research was supported in part by National Science Foundation grant BSR-8607535. I than T. L. Hunt and P. V. Kirch for inviting me to study the bird bones from To'aga. L. Nagaoka and D. S. Pahlavan assisted in curation. S. L. Olson and M. P. Walters kindly shared information on megapodes. For comments on the manuscript, I thank T. L. Hunt, P. V. Kirch, N. G. Miller, S. L. Olson, and M. P. Walters. 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