A Chronostratigraphic Analysis of Landbird Extinction on Tahuata, Marquesas Islands

Journal of Archaeological Science (1996) 23, 81–94

A Chronostratigraphic Analysis of Landbird Extinction onTahuata, Marquesas Islands

David W. Steadman*

Biological Survey, New York State Museum, Albany, New York 12230, U.S.A.

Barry Rolett

Department of Anthropology, University of Hawaii, Honolulu, Hawaii 96822, U.S.A.

(Received 23 July 1994, revised manuscript accepted 16 December 1995)

The Hanamiai archaeological site (Tahuata, Marquesas Islands) has yielded a stratified assemblage of bird bonesassociated with prehistoric Polynesian artefacts, cultural features, and other faunal materials spanning an 800-yearperiod beginning at c. 1025–1100. The majority of the c. 650 identifiable bird bones are from seabirds, most speciesof which are extirpated on Tahuata. Indigenous landbirds (rails, pigeons, doves, parrots, kingfishers, warblers) arerepresented by 70 bones of 10 species, among which two rails, a pigeon, a dove, and three parrots no longer occur onTahuata. The stratigraphic distribution of bird bones suggests that exploitation of indigenous species was most intenseearly in the Hanamiai cultural sequence (Phase I; c. 1025–1300). By c. 1450 or before, the seven extirpated speciesof landbirds (and probably other species not recorded in the small bone sample) either had been eliminated or hadbecome uncommon enough to elude archaeological sampling. A similar chronostratigraphic pattern of prehistoriclandbird extinction has been documented for Mangaia, Cook Islands. While details of timing and taxonomy vary fromisland to island, an overall trend of early prehistoric depletion of birds is the rule across East Polynesia and all ofOceania. ? 1996 Academic Press Limited

Keywords: AVIAN ARCHAEOFAUNAS, PREHISTORIC SUBSISTENCE, PREHISTORIC EXTINCTIONS,POLYNESIA.

Introduction

A rchaeological evidence provides numerousrecords of Pacific island avian extinctions,indicating an inextricable link between wide-

spread faunal losses and prehistoric human coloniz-ation (Olson & James, 1982; James et al., 1987;Steadman, 1989a). As first determined for theMarquesas Islands by Kirch (1973) and now deducedfrom zooarchaeological assemblages in a number ofPolynesian island groups (Dye & Steadman, 1990;Steadman & Kirch, 1990), human subsistence on ter-restrial vertebrates involved relatively heavy exploita-tion of indigenous birds during the early stages ofisland cultural sequences. Many populations or speciesof indigenous birds do not seem to have survived theearly period of exploitation. After the depletion ofmost native birds, domesticates and commensals in-cluding chickens, pigs, dogs, and rats became moreimportant in the diet (Steadman & Kirch, 1990; Rolett,1992; Steadman, Vargas & Cristino, 1994). Never-

theless, only on Mangaia (Cook Islands) havearchaeologically-derived data elucidated the precisestratigraphic, chronological, environmental, and cul-tural contexts of human-induced declines in native birdpopulations (Steadman & Kirch, 1990; Kirch, Flenley& Steadman, 1991; Kirch et al., 1992, in press; Kirch &Ellison, 1994; Ellison, 1994).Thus, although the occurrence of prehistoric avian

extinctions is now well established, the exact timing ofthese events, and the processes responsible for them,are only beginning to be documented. An improvedunderstanding of the environmental and cultural fac-tors at play, and the dynamic aspect of their relation-ship, requires detailed records of specific episodes ofextinction. Such data are best derived from strati-graphically controlled excavations of archaeologicalsites with well preserved bones from a continuoussequence of precisely dated deposits.This paper focuses on the landbird losses associated

with the prehistoric human occupation of Tahuatain the Marquesas Islands. Our study is based onintegrated zoological and archaeological analyses ofbones excavated from the Hanamiai Dune, a coastalsite with deeply stratified deposits representing a

*Present address: Florida Museum of Natural History, University ofFlorida, Gainesville, FL 32611, U.S.A.

810305-4403/96/010081+14 $12.00/0 ? 1996 Academic Press Limited

800-year cultural sequence (Rolett, 1992, in press). Thecontinuous series of occupations, from as early as 1025 until the 19th century, provides a well con-trolled chronological context for interpreting the ex-tinction of various rails, pigeons, doves, and parrots.We will present the evidence of a changing landbirdcommunity within the framework of local human-induced changes in an island ecosystem.

The Hanamiai SiteSite settingThe Marquesas Islands are a somewhat isolatedgroup of 14 eroded volcanic islands situated along theeastern margin of Polynesia (Figures 1 & 2). Thearchipelago lies between latitudes 7)50* and 10)35*S,about 5500 km west of the coast of South America.The nearest islands are coral atolls of the Tuamotus,to the south and west. The closest high islands areTahiti and other islands in the Societies, lying1400 km to the southwest.The Marquesan landscape is characterized by steeply

sloping valleys separated by knife-edged ridges de-scending from mountainous interiors. The sharp ridgesand the absence of a coastal plan combine to restrictbiotic and cultural interchange between separatevalleys. In addition to this relatively high degree ofintra-island isolation of habitable valleys, another keyenvironmental factor in the Marquesas is the un-even distribution of rainfall. Orographic precipitationassociated with prevailing tradewinds produces anaverage annual rainfall on windward coasts that isabout 40% more than that of nearby leeward areas(Cauchard & Inchauspe, 1978).Human activities have substantially transformed the

native vegetation since initial Polynesian discovery and

colonization of the Marquesas, which probably oc-curred about 2000 years ago (Rolett, 1993). At present,introduced species and secondary vegetation dominateall areas except the rain forest zone, generally found onmountain ridges and summits above 900 m elevation(Decker, 1970; Hallé, 1978). As is the case elsewhere inPolynesia (Kirch, 1983), much of this vegetationalchange dates to prehistory and results from (1) theintentional Polynesian introduction of a wide range ofspecies including fruit trees and other economicallyimportant plants and (2) deforestation, especially inassociation with agricultural activities and periodicburning.Tahuata lies in the southern half of the Marquesas,

only 3 km from the much larger (336 km2) island ofHiva Oa. At 53 km2, Tahuata is one of the smallestinhabited islands in the archipelago. The HanamiaiDune archaeological site, named after the valleyin which it lies, is on the leeward (western) coastof Tahuata. Hanamiai and the adjacent valley ofVaitahu are part of a large amphitheatre-shaped for-mation, bounded in back by a 1000 m high section ofthe central mountain chain and on the sides by ridgesthat descend abruptly from this chain to the rockyshoreline. While situated on a leeward coast, thisvalley of about 5 km2 lies close enough beneathTahuata’s central mountain ridge to benefit fromorographic showers and thus receives more rainthan is typical of leeward areas. Lush anthropogenicforests dominated by coconut palms and fruit trees(mango, breadfruit, citrus) blanket the bottoms andlower slopes of the Hanamiai and Vaitahu valleys,with patches of a much less altered wet tropical forestat high elevations near the cloud zone. The presentpopulation of Vaitahu and Hanamiai is only about350 persons, although the number of inhabitants

Philippines

Palau

MarianaIslands

CarolineIslands

Australia

SulaweslNewGuinea

HalmaheraNewIreland

SolomonIslandsNew

Britain

Vanuatu

New Caledonia

Fiji

Tonga

TuvaluTokelau

GilbertIslands

MarshallIslands

Hawaii

Line Islands

Samoa

NiueCookIslands

Tubuai

SocietyIslands

MarquesasIslands

TuamotuArchipelago

PitcairnGroup Easter

Sala-y-Gomez

Clipperton

Revillagigedo

Mexico

140° 160° 180° 160° 140° 120° 100°

30°

20°

10°

0°

10°

20°

30°

Figure 1. The tropical Pacific, showing the location of the Marquesas Islands.

82 D. W. Steadman and B. Rolett

during late prehistory probably reached into thethousands.

Stratigraphy and Chronology

The Hanamiai Dune archaeological site was excavatedin 1984 and 1985 by B. Rolett and field assistantsin cooperation with the Départment Archéologie duCentre Polynésien des Sciences Humaines, Tahiti(Rolett, in press). The main focus of investigation, a21 m2 area located some 30 m from the coast, revealeda clearly defined series of stratified cultural depositsextending to depths of about 1·6 m below the sur-face. The stratigraphic sequence, consisting of eightprincipal deposits (Layers A–H; Figure 3), recordscontinuous human occupation of the site from about 1025–1100 until the mid 19th century. Layers A–H,each judged to represent a separate depositional event,were distinguished on the basis of physical propertiessuch as colour and sediment texture, with attention aswell to differences in cultural content. Only trowelswere used for digging and all of the excavateddeposits were sieved through screens of 1/8 in. mesh.Within stratigraphic layers, excavation proceeded by 5or 10 cm arbitrary levels. The artefacts and faunal

remains for each level were bagged separately for each1 m2 unit. Artefacts found in situ, and cultural featuressuch as hearths, postholes, and stone pavements, wereplotted to scale.The earliest archaeological deposits in the Hanamiai

sequence represent an occupation located near whatwas probably the top of the dune, at an elevation of

Hatutu

Eiao

Marquesas Islands

Ua Huka

Nuku Hiva

Ua Pou

Hiva Oa

Tahuata

Fatu Hiva

Motane

0 10 20 30 40 50 km

140° W

10° S

Fatu Huku

Figure 2. The Marquesas Islands.

L11 M11 N11A and B

Sterile sand mixed with alluvium

GH mixed with H H 14C sample AA2820* 890±80 BP

Pavement stone

C

F

G

E D

E

50 cmBasaltCoral

Fire-cracked basaltBurnt surface

Hanamialcultural

chronology

Phase V – (AD 1800–1850) –

Phase IV(AD 1450–1800)

Phase III(AD 1300–1450)

Phase II

– (AD 1025–1300) –

Phase I

Figure 3. Stratigraphic profile, Hanamiai Dune site, Tahuata,Marquesas Islands. Northwest face of areal excavation.

Landbird Extinction on Marquesas Islands 83

1·7 m above the present high tide line. Prior to humansettlement, a small meandering stream had traversedthe dune in this area, leaving a 12–15 m wide channeland deposits of volcanic alluvium. The area of initialsettlement extended to the edge of the stream channel,creating two contemporaneous but geologically dis-tinct cultural deposits—Layer GH (mixed calcareousand volcanic sands) and Layer H (sands with largeamounts of stream-deposited alluvium).Layers GH and H represent Phase I of the Hanamiai

cultural sequence, a period dated by three samples ofwood charcoal, two securely associated with in situburnt surfaces and the third recovered in isolation.These samples yielded radiocarbon dates ranging from890&80 to 790&80 years (Table 1). At 2ó (95%probability), calibrated ranges of two samples from thedeepest levels of Layers GH and H (AA2820, Beta-15567) are within the interval from 1025–1300.Sample AA2819, from the upper levels of Layer H,yielded an overlapping but somewhat later age range of 1162–1398. Because the stratigraphically super-posed Layer G is dated to the same age range, theactual time period represented by Layers GH and H isprobably less than 300 years, and it likely falls into theearly or middle portions of the interval 1025–1300.Although a single posthole represents the only possiblearchitectural evidence for Phase I, other remainsincluding hearths, pearl shell and whale bone tool

manufacturing areas, and a wide range of utilitarianartefacts, such as adzes and fishhooks, clearly identifythe site as a habitation area during this initial stage ofoccupation (Rolett, in press).Layer G, a mixed calcareous and volcanic sand

deposit representing Phase II of the Hanamiai se-quence, is distinguished by a massive stone pavement,probably the veranda area of a pole and thatch struc-ture marked by an associated array of postholes. Asingle wood charcoal sample from a hearth in Layer Gprovides an age estimate of 870&80 years , which isstatistically inseparable from the three dates for PhaseI (Table 1). This suggests that both Phases I and IIoccurred during a time period spanning no more than300 years, perhaps even less than 200 years. Theartefact assemblages from Layer G indicate continueduse of the site as a general habitation area, with specificevidence for animal butchering, woodworking, and themanufacture of tools including basalt adzes and pearlshell fishhooks.Phases III and IV, documented by Layers C through

F, reveal continued use of the dune as a generalhabitation area. The archaeological evidence fromboth of these phases includes low stone platforms(presumably for houses), hearths, and a wide variety ofmaterials that represent the reworking of adzes, animalbutchering, and other domestic activities. While theartefactual and faunal evidence indicates notable

Table 1. Radiocarbon chronology for the Hanamiai Dune cultural sequence, Tahuata, Marquesas Islands

Laboratorynumber Provenience Context

13C/12Ccorrected14C age cal

Calibratedage range(2ó)

AA2820 Unit M11 Isolated chunks 890&80 1218 1019–1295V3738 Layer H

257 cmbd

AA2819 Unit M11 Burnt surface 790&80 1282 1162–1398V3737 Layer H (Feature 22)

234 cmbd

Beta- Unit I10 Hearth 850&60 1245 1051–129615567 Layer GH (Feature 45)

235–240 cmbd

AA2822 Unit I10 Hearth 870&80 1225 1025–1300V3740 Layer G (Feature 44)

215–220 cmbd

AA2821 Units K9, K10 Hearth 660&80 * 1326, 1287–1436V3739 Layer F (Feature 38) 1334, 1395

179–184 cmbd

Beta- Units K9, K10 Hearth 620&90 * 1326, 1287–143615566 Layer F (Feature 38) 1334, 1395

179–184 cmbd

Beta- Unit K10 Earth oven 130&100 1896, 1644–195515565 Layer B (Feature 32) 1902, 1955

99 cmbd

All samples are wood charcoal. Calibration according to Stuiver & Pearson (1993), including adjustment forsamples from the southern hemisphere. cmbd: centimetres below datum. * weighted average of both samples fromFeature 38.


transformations in some aspects of subsistence andmaterial culture during Phases III and IV, the overallfunction of the site remained unchanged. Phase III isdated by two independently analysed samples of woodcharcoal from a large hearth in Layer F. These samplesyielded similar 14C dates of 600&80 and 620&90 years , with a 2ó range of 1287–1436 based onthe weighted average. There are no radiocarbon esti-mates for Phase IV (Layers C, D). The presence ofhistoric era artefacts in Layers A and B, and asingle date of 130&100 years , place the terminalHanamiai occupation, Phase V, in the early 19thcentury. Layers A and B represent transformation ofthe Hanamiai Dune into part of a large ceremonialground (tohua) and an associated fisherman’s precinct(mahia avi a’a), as evidenced by terracing of the siteand the construction of monumental stone platforms.

Materials and MethodsThe Hanamiai bird bones are catalogued in the collec-tions of the Vertebrate Zoology Department, BerniceP. Bishop Museum, Honolulu, Hawaii (BPBM).Identification of the bones (by DWS) is based upondirect comparison with modern skeletons from theAmerican Museum of Natural History (AMNH),Delaware Museum of Natural History (DMNH),National Museum of Natural History, SmithsonianInstitution (USNM), New York State Museum(NYSM), and Thomas Burke Memorial Museum,University of Washington (UWBM). Overall labora-tory methodology follows Rea (1986). NISP is thenumber of identified specimens and years are theradiocarbon years before present ( 1950).The brief species accounts include pertinent informa-

tion on osteology, systematics, past and present distri-bution, and current status on Tahuata. Osteologicalterminology generally follows Baumel et al. (1993).Unless stated otherwise, modern distribution andpopulation status are derived from Holyoak (1975),Thibault & Thibault (1975), duPont (1976), Holyoak &Thibault (1984), Pratt, Bruner & Berrett (1987), andSeitre & Seitre (1991, 1992). The sequence of speciesfollows Steadman (1989a). The landbird category com-prises all species that do not feed over the open ocean.The seabird bones from Hanamiai, still under study,will be the topic of a future paper. This rich assemblagerepresents more than 15 species of shearwaters, petrels,prions, storm-petrels, boobies, frigatebirds, tropic-birds, and terns, most of which no longer occur onTahuata.

Systematic AccountsOrder: GalliformesFamily: PhasianidaeGallus gallus (Chicken) (Linnaeus, 1758)

Material. Twenty-two specimens: 2 vertebrae, BPBM166562, 166719; 3 synsacra, BPBM 166521, 166524,166525; 3 sterna, BPBM 166507, 166508, 166592; cora-coid, BPBM 167110; 2 humeri, BPBM 166539, 166883;ulna, BPBM 166563; radiale, BPBM 166594; pelvis,BPBM 166523; 2 femora, BPBM 166503, 166564;fibula, BPBM 166585; 2 tarsometatarsi, BPBM 166983;167012; 3 pedal phalanges, BPBM 166453, 166595,166724.

Remarks. The chicken, like the pig, dog, and Pacificrat, was introduced prehistorically to the MarquesasIslands. Chicken bones are relatively rare in Phase I.They become more common in Phase II and remain sointo Phase IV. This may reflect an increased con-sumption of chickens as native bird populationsbecame smaller, as documented on Mangaia, CookIslands (Steadman & Kirch, 1990) and Easter Island(Steadman, Vargas & Cristino, 1994).Nevertheless, chickens are never a major component

of the Hanamiai assemblage. The absence of chickenbones in Phase V may be an artefact of small samplesize.

Order: GruiformesFamily: RallidaeGallirallus new sp. (Tahuata Rail)

Material. Twenty-four specimens: rostrum, BPBM166456; 5 mandibles, BPBM 166436, 166439, 166457,166458, 166542; coracoid, BPBM 166455; 3 femora,BPBM 166435, 166445, 166447; 5 tibiotarsi, BPBM166437, 166438, 166444, 166446, 166452; 5 tarsometa-tarsi, BPBM 166440, 166441, 166448, 166451, 166454;2 pedal phalanges, BPBM 166449, 166450; 2 ungualphalanges, BPBM 166442, 166453.

Remarks. This large flightless rail is one of four extinct,undescribed species of Gallirallus from the MarquesasIslands (Steadman, 1989a), each species being endemicto a single island. Osteological characters for Poly-nesian species of Gallirallus versus Porzana andPorphyrio are given by Steadman (1987, 1988). Repre-sented by 24 bones, Gallirallus new sp. is the mostcommon landbird at Hanamiai, with 19 of these bonesfrom Phase I.

Porphyrio paepae (Marquesas Swamp-Hen)(Steadman, 1988)

Material. Fourteen specimens: ulna, BPBM 166677;carpometacarpus, BPBM 166427; synsacrum, BPBM166432; 4 femora, BPBM 166424, 166426, 166430,166434; 3 tibiotarsi, BPBM 166425, 166428, 166433;tarsometatarsus, BPBM 166429; 3 pedal phalanges,BPBM 166431, 166442, 167117.

Remarks. This extinct species is known from archaeo-logical sites on Hiva Oa (including the holotype femur)


and Tahuata. It is the only species of Porphyrioknown unequivocally from East Polynesia. Four ofthe 14 specimens reported herein (BPBM 166442,166677, 166431, 167117) were not among thosedescribed by Steadman (1988). Except for the threebones from below Phase I (possibly pre-culturalsediment; see Table 2), the stratigraphic distributionof P. paepae at Hanamiai resembles that of Gallirallusnew sp.

Gallirallus/Porphyrio (Indeterminate Rail)

Material. Femur, BPBM 166443.

Remarks. This specimen cannot be assigned with cer-tainty to either of the two large rallids from Hanamiai.

Order: ColumbiformesFamily: ColumbidaeGallicolumba rubescens (Marquesas Ground-Dove)(Vieillot, 1818)

Material. Five specimens: coracoid, BPBM 166802; 2ulnae, BPBM 166553, 166785; 2 tibiotarsi, BPBM166783, 166801.

Remarks. Osteological characters of Gallicolumba (andother Polynesian genera of columbids) are outlined inSteadman (1992). Although recorded in archaeologicaldeposits from Nuku Hiva, Hiva Oa, Ua Huka, and

Tahuata (Steadman, 1989a, pers. obs.), modernrecords of G. rubescens are confined to Nuku Hiva,Hatutu, and Fatu Huku. This small dove, endemic tothe Marquesas, now survives only on the small, un-inhabited islands of Hatutu and Fatu Huku (Gifford,1925; Collar & Andrew, 1988: 63). All bones ofG. rubescens at Hanamiai are from Phase I.

Ptilinopus dupetithouarsii (White-capped Fruit-Dove)(Neboux, 1840)

Material. Two tarsometatarsi, BPBM 166795, 166798.

Remarks. This is the most common and widespreadcolumbid in the Marquesas today, although archaeo-logical evidence shows that all six Marquesan speciesof pigeons and doves, three of which are extinct, werewidespread in early prehistoric times (Steadman,1989a). Considered a delicacy, the White-cappedFruit-Dove commonly feeds in banyans (Ficus sp.) andwas captured traditionally by spreading bird-lime(made from breadfruit sap and coconut oil) on thelower branches of these trees (Rollin, 1929: 138). TheTahuata population of White-capped Fruit-Dove isdeclining (Seitre & Seitre, 1991), perhaps because ofpredation from the non-native Great Horned Owl(Bubo virginianus).

Ducula galeata (Nuku Hiva Pigeon) (Bonaparte, 1855)

Table 2. Bird bones from the Hanamiai Dune archaeological site, Tahuata, Marquesas

Species #

Phase

TotalI II III IV V

Gallus gallus (i) — 3 9 4 6 — 22Gallirallus new sp.‡ — 19 3 2 — — 24Porphyrio paepae‡ 3 9 1 1 — — 14Gallirallus/Porphyrio‡ — 1 — — — — 1Gallicolumba rubescens† — 5 — — — — 5Ptilinopus dupetithouarsii — 1 — 1 — — 2Ducula galeata† — 6 — — — — 6Columbidae sp. — 1 — — — — 1Vini ultramarina† 1 — — 1 3 — 5Vini vidivici‡ — 4 — — — — 4Vini sinotoi‡ — 3 — — — — 3Vini sp.‡ — 2 — — — — 2Halcyon godeffroyi — — 1 — — — 1Acrocephalus mendanae — — — — 2 — 2TotalsAll native landbirds 4 51 5 5 5 0 70Species† 1 11 0 1 3 0 16Species‡ 3 38 4 3 0 0 48Species†/‡ 4 49 4 4 3 — 64% (vs all landbirds)†/‡ 100 96 80 80 60 — 91% (vs all species of vertebrates)†/‡ — 1·5 0·16 0·16 0·16 0·07 0·39Native landbird species 2 7 3 4 2 — 10Species†/‡ 2 6 2 3 1 — 7

Numbers are NISP.(i) introduced species (not included in totals).† extirpated species.‡ extinct species.# possibly pre-cultural sediment.


Material. Six specimens: scapula, BPBM 166808;humerus, BPBM 166804; radius, BPBM 166807;pelvis, BPBM 166809; tarsometatarsus, BPBM 166805;ungual phalanx, BPBM 166806.

Remarks. Critically endangered and confined now toNuku Hiva (Collar & Andrew, 1988: 66–67; Seitre &Seitre, 1991), D. galeata is a very large pigeon that onceoccurred across much of East Polynesia (Steadman &Olson, 1985; Steadman, 1989a; Steadman & Kirch,1990; Steadman & Pahlavan, 1992). All six bones fromHanamiai are from Phase I.

Columbidae sp. (Undetermined dove)

Material. Femur, BPBM 166793.

Remarks. This fragmentary specimen is approximatelythe size of the femur in Ptilinopus, but is otherwiseundiagnostic.

Order: PsittaciformesFamily: PsittacidaeVini ultramarina (Ultramarine Lorikeet) (Kuhl, 1820)

Material. Five specimens: mandible, BPBM 166057;sternum, BPBM 166063; humerus, BPBM 166058;carpometacarpus, BPBM 172009; femur, BPBM166593.

Remarks. These bones were reported by Steadman &Zarriello (1987). Like the other two species of Vini inthe Marquesas, V. ultramarina is larger on Tahuatathan on Ua Huka. This may warrant trinomial (sub-species) recognition, pending a more thorough analysisof interisland osteological variation. V. ultramarina isknown historically from Nuku Hiva and Ua Pou,although the largest extant population lives on UaHuka, derived from a re-introduction made in the1940s (Collar & Andrew, 1988: 68–69; Seitre & Seitre,1991). Seven individuals of V. ultramarina recentlywere translocated to Fatu Hiva as well (Anonymous,1994). Three of the five bones of V. ultramarina fromHanamiai are from Phase IV, showing that this speciesprobably survived on Tahuata a century or two longerthan most of the other extirpated or extinct species.

Vini vidivici (Conquered Lorikeet) (Steadman &Zarriello, 1987)

Material. Four specimens: coracoid, BPBM 166064;humerus, BPBM 166782; 2 tarsometatarsi, BPBM166059, 166061.

Remarks. Except for the humerus, these specimenswere first reported by Steadman & Zarriello (1987). Werefer the humerus to Vini because of its more roundedcaput humeri and the more distinct distal margin ofsulcus ligamentum transversus. BPBM 166782 agrees

in size with paratype specimens of V. vidivici from UaHuka, being smaller than in V. sinotoi and larger thanin V. ultramarina. V. vidivici is an extinct species knownfrom archaeological sites on Nuku Hiva, Ua Huka,Hiva Oa, and Tahuata in the Marquesas (Steadman& Zarriello, 1987; Steadman, pers. obs.), Huahine,Society Islands (Steadman & Pahlavan, 1992), andMangaia, Cook Islands (Steadman & Kirch, 1990). AtHanamiai, all bones of V. vidivici and V. sinotoi arefrom Phase I, suggesting that both of these parrotsbecame rare or extinct relatively early in the permanenthuman occupation of the island.

Vini sinotoi (Sinoto’s Lorikeet) (Steadman & Zarriello,1987)

Material. Three specimens: rostrum, BPBM 166062; 2carpometacarpi, BPBM 166065, 166896.

Remarks. These bones were reported by Steadman &Zarriello (1987). V. sinotoi is the largest species in itsgenus. Like V. vidivici, this extinct species is known aswell from archaeological sites on Nuku Hiva, Hiva Oa,and Ua Huka (Steadman & Zarriello, 1987; Steadman,pers. obs.) and Huahine, Society Islands (Steadman &Pahlavan, 1992). Unlike V. vidivici, however, V. sinotoihas not been recorded from the Cook Islands.

Vini vidivici/sinotoi (Conquered/Sinoto’s Lorikeet)(Steadman & Zarriello, 1987)

Material. Two specimens: pedal phalanx, BPBM166060; ungual phalanx, BPBM 166066.

Remarks. These specimens are from either V. vidivici orV. sinotoi, being larger than in V. ultramarina.

Order: CoraciiformesFamily: AlcedinidaeHalcyon godeffroyi (Marquesas Kingfisher) (Finsch,1877)

Material. Humerus, BPBM 166949.

Remarks. This endangered species survives only onHiva Oa (very rare) and Tahuata (declining; Collar& Andrew, 1988: 92; Seitre & Seitre, 1991). Otherarchaeological records of H. godeffroyi are from NukuHiva and Ua Huka (Steadman, 1989a, pers. obs.).

Order: PasseriformesFamily: SylviidaeAcrocephalus mendanae (Marquesas Reed-Warbler)(Tristram, 1883)

Material. Two specimens: coracoid, BPBM 166948;humerus, BPBM 167013.

Remarks. Osteological characters of Acrocephalus ver-sus Aplonis follow Steadman (1989b). BPBM 167013


agrees in size and qualitative characters with twomodern specimens of A. mendanae from Tahuata. Thecorpus humeri is too stout for Pomarea or Myiagra.A. mendanae is a Marquesan endemic that still surviveson many islands in the group, including Tahuata.Other prehistoric records of oceanic Acrocephalus areA. kerearako from Mangaia, Cook Islands (Steadman,1985) and A. vaughani from Henderson Island, PitcairnGroup (Schubel & Steadman, 1989). Neither of theserecords represents an extirpated population. Thespecies of Acrocephalus are less dependent upon forestthan other East Polynesian passerines, and thus aremore tolerant of anthropogenic changes in terrestrialhabitats.

Discussion and ConclusionsChronological changes in subsistenceThe Hanamiai Dune archaeological site has a clearlystratified sequence of undisturbed cultural depositswith excellent preservation of vertebrate and inverte-brate faunal materials. The continuous, nearly 800 yearlong Polynesian occupation of the site began atc. 1025–1100 and extended until the mid 19th cen-tury. Our data derive from a single excavated area of21 m2, within which the horizontal variation in stratig-raphy is minimal. Quantitative analysis of the exca-vated faunal remains points to several long-term trends

in subsistence strategies. These results provide a cul-tural and chronological context for interpreting theprehistoric declines in bird populations that eventuallyled to the disappearance of species.The earliest archaeological deposits (Layers GH and

H), representing Phase I in the Hanamiai culturalsequence, yielded 503 of the 637 identifiable bones ofnative birds recovered from the site (Table 3). Thesebird bones are part of a large Phase I bone assemblage(NISP=3290) that reflects a prehistoric subsistencestrategy in which seabirds, and landbirds to a lesserextent, were part of a diversified diet dominated by fishbut also including sea turtle, marine mammals, pig,dog, and rat. In Phase I, however, the bones ofnon-native species (pig, dog, rat, human, chicken)represent only 2·4% of the entire assemblage of identi-fiable bones, a value that increases to 12·3% by PhaseII and 21·9% by Phase V (Table 3). Pigs account formuch of the steady increase through time in the dietaryimportance of domesticates, whether evaluated as per-centages or rank order (Table 4). A large number(94%) of the 447 identified pig bones date to PhasesIII–V, suggesting an expansion of animal husbandrypractices and the broader food production economybeginning around 1300 (Rolett & Chiu, 1994). Thisfinding is at variance with the recent claim by Sweeney,Graves & Hunt (1993) that stability, rather thanchange, was the predominant subsistence trend in

Table 3. The bone assemblage from Hanamiai Dune archaeological site, Tahuata, Marquesas

Taxon

Phase

TotalI II III IV V

MammalsPig (Sus scrofa) 4 22 65 251 105 447Dog (Canis familiaris) 2 26 20 50 11 109Rat (Rattus exulans) 60 269 165 506 28 1028Small whale (Odontoceti) 28 17 94 166 14 319Medium whale (Odontoceti) 10 4 0 2 0 16Human (Homo sapiens) 0 1 3 2 2 8*Unidentifiable medium mammal 59 45 108 459 106 777

BirdsNative seabirds 448 88 13 14 4 567Native landbirds 55 5 5 5 0 70Chicken (Gallus gallus) 3 9 4 6 0 22*Unidentifiable 26 10 2 4 0 42

ReptilesSea turtle (Cheloniidae sp.) 20 10 7 9 3 49

Fish 2292 2216 2001 4401 501 11,411

Other*Unidentifiable medium vertebrate 283 337 708 1666 228 3222

TotalsAll species 3290 3059 3195 7541 1002 18,087All species (without *) 2922 2667 2377 5412 668 14,046Introduced species 69 327 257 815 146 1614% introduced species 2·4 12·3 10·8 15·0 21·9 11·4

Numbers for seabirds and unidentifiable birds are approximate. Modified from Rolett (1992).Numbers are NISP.Bones listed under ‘‘#’’ in Table 2 included under Phase I herein.* These categories not included in calculating percentages in Tables 3 and 4.


Marquesan prehistory. Rather, the Hanamiai datasupport Kirch’s (1989: 159–160) general model for thedevelopment of Polynesian production systems, onethat links long term transformations in subsistence to adynamic interplay between cultural and environmentalfactors. This interplay was initiated by the humancolonization of previously uninhabited islands.Rats, whose predatory impact on ground-dwelling

and cavity-nesting birds is well known, were present inlarge numbers by Phase II. It may be no coincidencethat, during Phases I, II, and III, native seabirdsplunge from 15·3 to 3·3 to 0·5% (rank order 2, 3, and 6)and native landbirds decrease from 1·9 to 0·2% (rankorder 4, 9, and 8).Throughout the Hanamiai sequence, the dominant

trend in subsistence is a heavy reliance on fishing,reflected by the high frequency (75–84%) and rankorder (always 1) of fishbone in each of the five phases(Table 4). The lowest frequency (75%) of fishbone atHanamiai, in the Phase V assemblage, may be linked atleast partly to the change in site function from generalhabitation to a ceremonial area. Within the context ofa strong maritime orientation, there is a notable shift,however, from the diversified exploitation of bothoffshore and inshore marine environments duringPhases I–III to a more narrow focus on inshore fishingduring Phases IV and V (Rolett, in press). Dye (1990)regarded this shift in Marquesan marine exploitationas being due more to social than environmental fac-tors. In spite of the undisputed abundance of fishbonein Marquesan archaeofaunas, even higher percentagesof fishbone occur at sites in the Cook Islands(Allen, 1992; Steadman & Kirch, 1990; Kirch et al., inpress), where fringing reefs (absent in the Marquesas)facilitated exploitation of shallow water species.

Extinction of birds: process

Human predation, habitat loss, and the impacts ofintroduced predators (such as rats, pigs, and dogs),

competitors, or pathogens are generally cited as theprimary causes for the modern as well as prehistoricextinctions of island birds (Diamond, 1984; Steadman,1989a; Steadman, Greiner & Wood, 1990; Seitre &Seitre, 1991, 1992). The last three factors are a complexsets of related phenomena whose impact is difficult todocument directly through the archaeological record.Human predation and habitat loss are more amenableto archaeological and sedimentological investigation.The value of the Hanamiai data in helping to explain

these processes stems both from the inherent wellstratified nature of the site itself and the interdiscipli-nary approach used in its investigation. Finding thebones of extinct species in an archaeological contextdoes not prove, however, that people caused the extinc-tion, even when the bones are burned or show otherdistinct evidence of butchering, cooking, or consump-tion. Such evidence does argue, however, for humanhunting and consumption of the extinct species, whichwe presume, based upon modern analogy and commonsense, to have been at least one of the factors leadingtoward extinction (see also Weisler & Gargett, 1993).The direct evidence for habitat changes in prehistoric

Polynesia comes not from the bones themselves butfrom palaeobotanical and geomorphological studies,such as Kirch et al. (1991) and Ellison (1994) forMangaia (Cook Islands), Lepofsky, Harries & Kellum(1992) for Mo’orea (Society Islands), and Flenley et al.(1991) for Easter Island. These studies document pre-historic deforestation, the introduction of non-nativeplants, and the erosion of uplands and infilling oflowland sedimentary basins. Such activities are associ-ated with agricultural expansion and intensification(Kirch, 1983).Among the generally recognized causal factors for

modern and prehistoric extinctions (human predation,habitat loss, and introduced predators, competitors,and pathogens), it seems safe to assume that at least thefirst three were involved in the loss of all seven extir-pated species of landbirds on Tahuata. Each of these

Table 4. The bone assemblage from Hanamiai Dune archaeological site, Tahuata, Marquesas

Taxon

Phase

Total% (RO)

I% (RO)

II% (RO)

III% (RO)

IV% (RO)

V% (RO)

Pig 0·1 (8) 0·8 (5) 2·7 (4) 4·6 (3) 15·7 (2) 3·2 (4)Dog 0·1 (10) 1·0 (4) 0·8 (5) 0·9 (5) 1·6 (5) 0·8 (6)Rat 2·0 (3) 10·0 (2) 7·0 (2) 9·3 (2) 4·2 (3) 7·3 (2)Small whale 1·0 (5) 0·6 (6) 4·0 (3) 3·0 (4) 2·1 (4) 2·3 (5)Medium whale 0·3 (7) 0·1 (10) 0 (11) <0·1 (10) 0 (9) 0·1 (10)Human 0 (11) <0·1 (11) 0·1 (10) <0·1 (10) 0·3 (8) 0·1 (11)Native seabirds 15·3 (2) 3·3 (3) 0·5 (6) 0·3 (6) 0·6 (6) 4·0 (3)Native landbirds 1·9 (4) 0·2 (9) 0·2 (8) 0·1 (9) 0 (9) 0·5 (7)Chicken 0·1 (9) 0·3 (8) 0·2 (9) 0·1 (8) 0 (9) 0·2 (9)Sea turtle 0·7 (6) 0·4 (7) 0·3 (7) 0·2 (7) 0·4 (7) 0·3 (8)Fish 78·4 (1) 82·7 (1) 84·2 (1) 81·3 (1) 75·0 (1) 81·2 (1)Totals 2922 2667 2377 5412 668 14,046

Expressed in relative percentages (%) and rank order (RO). Based upon data in Table 3, but with the threeunidentified categories excluded.


species is palatable and also a potential source offeathers for decoration (see Linton, 1923: 432–435) andbones for tools (Steadman, in press). The two ground-dwelling rails (Gallirallus new sp., Porphyrio paepae)and the ground-dove (Gallicolumba rubescens) prob-ably were easy to snare. They, along with the pigeonand parrots, could have been taken also with slings,thrown rocks, or by raiding nests. The last activity mayhave been particularly destructive to the cavity-nestingparrots. Bird-lime, used in capturing the fruit-dovePtilinopus dupetithouarsii (Rollin, 1929: 138), probablyalso was a means for trapping other volant species(pigeons, parrots, kingfishers, passerines).All of the seven extirpated species were forest in-

habitants, and thus would have undergone populationdeclines as forests were cut and burned for agriculture.Introduced predators (pigs, dogs, rats) would havetaken ground-dwelling species (rails, ground-doves),although except for the rat, these predators probablyhad minimal impact on pigeon and parrot populations.Prehistorically, the chicken may have been a poten-

tial competitor with the native birds of Tahuata,particularly the rails. We suspect, but cannot prove,that this impact was minimal. On the other hand,chickens may have been involved in the introduction ofavian pathogens to the Marquesas. While also difficultto demonstrate, clues to the role of chickens in theprehistoric extinction of native Polynesian birds mightbe gathered through additional studies of the currentdistribution and frequency of avian pathogens, espe-cially haematozoans, across Oceania (Steadman et al.,1990).

Extinction of birds: biogeography

Results of the Hanamiai research support a number ofconcepts derived from previous investigations of pre-historic East Polynesian bird bone assemblages. First,for most or all islands, more species of native birdshave disappeared than have survived. Second, al-though both seabird and landbird populations weredepleted by human activities, nearly all of the extinctspecies are landbirds. Most taxa of Marquesan sea-birds still survive somewhere, an exception beingCostello’s race of Abbott’s Booby, Papasula abbottiicostelloi (Steadman, Pahlavan & Schubel, 1988). Third,the current ranges of most surviving species of nativelandbirds are diminished compared to their prehistoricranges. An extreme example of this is the Nuku HivaPigeon, Ducula galeata, a species that is confined todayto Nuku Hiva but for which the prehistoric rangeextended not only throughout the Marquesas but alsoto the Societies and Cook Islands, archipelagoeslocated thousands of kilometres to the west (Steadman,1989a).At least 20 species of landbirds once lived in the

Marquesas, judging by current evidence from avianarchaeofaunas on four islands (Table 5). Of thesespecies, 10 are extinct and six others have been shown

to be extirpated on one or more islands. If the prehis-toric record were complete, it might well show thatvirtually all volant species were found on all majorislands in the Marquesas, with an understanding thatinterisland variation differed among the various taxa oflandbirds. For example, within each of the threespecies of Vini, size variation might warrant subspecieslevel recognition (Steadman & Zarriello, 1987), where-as the various forms of Pomarea have been treated as ablend of subspecies and superspecies (duPont, 1975;Pratt et al., 1987). Only flightless species were likely tohave been confined to single islands.

Extinction of birds: stratigraphy, chronology,cultural context

Six of the seven extinct or extirpated species of land-birds found at Hanamiai are recorded from the Phase Ideposits, while only two of the seven are recorded fromPhase II (Table 2). The species missing from Phase I isVini ultramarina, which was recorded just below PhaseI in sediments interpreted as precultural. Presumably,therefore, V. ultramarina was present during Phase Ibut was not sampled.Two of the three species of landbirds that still live on

Tahuata are represented only in early stages of theHanamiai sequence. The two bones of Ptilinopusdupetithouarsii are from Phases I and III, and the singlebone of Halcyon godeffroyi is from Phase II. Thissuggests either a greater propensity to hunt all land-birds early in the sequence, or larger overall landbirdpopulations, or, more likely, both of these relatedfactors.There is a dramatic overall decline between Phases I

and II in the representation of bird bones. This isreflected by a sharp decrease in the concentrationindex, or density of all bird bones per cubic metre ofcultural deposit (bones m"3), from 104 for Phase I to20 for Phase II. There is an even greater, 10-folddecrease in the bones of extinct or extirpated landbirdsbetween Phases I and II. The magnitude of this de-crease is roughly the same whether evaluated on thebasis of absolute counts (NISP) or as relative abun-dance (percentage of bird bones compared with othervertebrate remains; see Tables 2 & 4). Despite themarked difference between the Phase I and II birdassemblages, the archaeological evidence indicatesneither a change in site function nor a decrease in theintensity of human occupation. In addition to artefactassemblages similar to those of Phase I, Phase IIyielded a stone house pavement with adjacent evidenceof tool manufacturing and animal butchering, clearlyidentifying the site function as that of a generalhabitation area. Comparison of the overall faunalassemblages from Phases I and II (Tables 3 & 4)further indicates that, except for increases in non-native mammals and decreases in native birds, thegeneral subsistence strategy emphasized the exploita-tion of marine resources.


Given the near contemporaneity of Phases I and II,as measured by radiocarbon, the widely different dis-tribution of bird bones in these successive occupationsof the Hanamiai Dune is striking. In evaluatingthis difference, one must consider the possibility ofdisturbances that may have resulted in large scalepost-depositional stratigraphic displacement of the ar-chaeologically recovered materials. Although a limiteddegree of movement across stratigraphic boundarieshas occurred among materials deposited near the inter-faces (Rolett, in press), the vertical distribution ofsmall specimens presumably deposited at the same time(specifically, 73 fragments of the same whale bonemandible and a group of 19 pierced cowrie shells)suggests that few if any of the bones in the Phase I andPhase II deposits have been displaced more than 30 cmfrom their original depths. These findings supportthe hypothesis that stratigraphic proveniences of theHanamiai bird bones provide accurate data forseparating the avifaunal remains into chronologicallydistinct assemblages.

The analysis of land snails reveals another impor-tant difference between Phases I and II. While thePhase I land snail assemblage consists almost entirelyof native species, that from Phase II is dominated byspecies introduced inadvertently during the Poly-nesian colonization of the Marquesas (Rolett, inpress). This nearly complete faunal turnover indicatesa substantial environmental change, as introducedspecies of land snails are typically associated withanthropogenic vegetation and tend not to occupyundisturbed habitats (Harry, 1966). The abruptnessof this inferred shift in vegetation, and the sharplydeclining abundance of bird bones between Phases Iand II, reflect an intense period of environmentaltransformation linked to human activities. In additionto the introduction and cultivation of a large numberof utilitarian plants, these activities included the in-troduction and propagation of pigs, dogs, rats, andchickens on an island without indigenous terrestrialmammals. Quantified faunal data also suggest a de-clining availability of certain easily accessible species

Table 5. Landbirds from four Marquesan islands

Nuku Hiva Hiva Oa Ua Huka Tahuata

RailsGallirallus new spp.† A A A APorzana tabuensis M — — —Porzana new sp.† — — A —Porphyrio paepae† — A — ASandpipersProsobonia sp.† — — A —Pigeons, dovesGallicolumba rubescens A,M A A AGallicolumba nui† — A A —Macropygia heana† A — A —Ptilinopus dupetithouarsii A,M M A,M A,MPtilinopus mercierii† M M A —Ducula galeata M A A AParrotsVini ultramarina A,M — A AVini vidivici† A A A AVini sinotoi† A A A ASwiftsCollocalia ocista M M M MKingfishersHalcyon godeffroyi A M A A,MPasserinesPomarea iphis — — M —Pomarea mendozae M M — Mcf. Myiagra sp. — — A —Acrocephalus mendanae M M A,M A,MTotal species (M only) 9 6 3–4 5Total species (A, M) 14 13 16–17 12NISP *19 168 2056 70Island area (km2) 337 324 78 53Island elevation (m) 1185 1190 855 1000Distance (km) to nearest island 36 3 36 3

A: archaeological record.M: modern ( 1800 or later) record, not including 20th century re-introductions. Modified from Holyoak (1975)and Steadman (1989a).* a considerable amount of unstudied bird bone exists for Nuku Hiva.† extinct species.


of coastal marine molluscs between Phases I and II(Rolett, 1992).Unfortunately, none of the other prehistoric avian

assemblages from the Marquesas (Table 5) hasadequate stratigraphic and chronometric control topermit detailed comparisons with the Hanamiai assem-blage. Larger, more uniformly collected bone samplesare needed from each of these islands to evaluaterigorously the interisland differences in the prehistoricMarquesan avifauna. The data that exist, while ex-tremely useful in demonstrating the former presence ofvarious taxa, are not comprehensive. Inadequate sam-pling could account for most of the absences in the‘‘A’’ category of Table 5. Even the very large bird bonesample from the Hane site on Ua Huka is unlikely tobe complete, being biased against small species such asswifts, kingfishers, passerines, and certain shorebirdsbecause no finer mesh than 1/4 in. was used to sieve thesediments.The chronology of landbird extinction at Hanamiai

is similar to that from the Tangatatau Rockshelter (siteMAN-44), Mangaia, Cook Islands. Based upon exca-vations conducted at MAN-44 in 1989, Steadman &Kirch (1990) reported eight species of extinct or extir-pated landbirds, seven of which were confined to stratadated at 1000–1300. When combined with bonesdiscovered during more extensive excavations in 1991,a total of 13 species of extinct or extirpated landbirdsnow are known from MAN-44, 11 of which disappearfrom the record about or before 1300 (Kirch et al.,in press).We believe that initial settlement of the Marquesas

Islands may have occurred approximately 2000 yearsago (Rolett, 1993), although we stress that the chronol-ogy of initial human colonization of the Marquesas isnot well understood and is the subject of considerabledebate (Kirch, 1986; Irwin, 1992; Spriggs & Anderson,1993; Kirch & Ellison, 1994; Anderson et al., 1994).While it seems safe to assume that the Polynesians whodiscovered and colonized the Marquesas were aware ofall islands in the group, this does not necessarily meanthat each island within the Marquesas supported apermanent human population two millennia ago. Be-cause of the topographic barriers between Marquesanvalleys, it may be useful to evaluate the colonizationprocess on a valley-by-valley as well as island-by-islandbasis. With this perspective, it could be that manyinhabitable valleys in the Marquesas, while undoubt-edly visited at least occasionally, did not sustain per-manent settlement for centuries after initial humancolonization of the archipelago (Rolett, in press). Forexample, following permanent settlement of the largestvalleys on Nuku Hiva and Hiva Oa, considerable timemay have elapsed before growing populations ex-panded into the smaller valleys on small islands, suchas Hanamiai on Tahuata.Just how Tahuata fits into this hypothetical scheme

of dispersal and colonization is uncertain, pendingnew, well-dated archaeological excavation of sites

in other valleys on Tahuata and throughout theMarquesas. Regardless, the Hanamiai Dune archaeo-logical site has sampled a prehistoric extinction eventthat occurred, as we presently understand it, primarilyfrom about 1000 to 1300. We still do not knowwhat sort of human impact, if any, had been sustainedby the birds of Tahuata before 1000.Several years ago, we might have interpreted the

presence of two flightless rails at Hanamiai as evidencethat the basal deposits represent the first arrival ofpeople on Tahuata. We now know, however, that it ispossible even for flightless species to survive for manycenturies after first human contact. For example, thefaunal sequence at site MAN-44, Mangaia, shows thatflightless rails, while becoming extinct prehistorically,survived for more than a millennium after people firstarrived on Mangaia (Steadman & Kirch, 1990; Kirch& Ellison, 1994; Kirch et al., in press). At 52 km2,Mangaia has essentially the same land area as Tahuata(53 km2). Substantial portions of both islands are toosteep or rugged for habitation or cultivation. Suchinhospitable places would have retarded the processesof anthropogenic predation and deforestation. Thesedelays added centuries to the existence of variousspecies, but did not prevent their eventual extinction.

AcknowledgementsFieldwork by BVR was funded by the National ScienceFoundation (Doctoral Dissertation ImprovementGrant BNS-8514067) and the Augusta Hazard Fundof Yale University. Laboratory research by DWSwas funded by the National Science Foundation(Systematic Biology Grant BSR-8607535). DWSthanks the Department of Anthropology, University ofCalifornia, Berkeley, for providing the opportunity towork on the manuscript during a semester in residence.Fieldwork was greatly facilitated by M. Navarro,

Director of the Départment Archéologie du CentrePolynésien des Sciences Humaine, and T. Tetahiotupa,Mayor of Tahuata. D. S. Pahlavan, S. E. Schubel,E. L. Wootan, and M. C. Zarriello assisted in curationof specimens and certain aspects of data compilation.For comments on the manuscript, we thank P. L.Bruner, H. F. James, and S. L. Olson.

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A Chronostratigraphic Analysis of Landbird Extinction on Tahuata, Marquesas Islands

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