New Species of Extinct Rails (Aves: Rallidae) from ... · 145 New Species of Extinct Rails (Aves: Rallidae) from Archaeological Sites in the Marquesas Islands, French Polynesia1 Jeremy

145

New Species of Extinct Rails (Aves: Rallidae) from Archaeological Sitesin the Marquesas Islands, French Polynesia1

Jeremy J. Kirchman2,4 and David W. Steadman3

Abstract: We examined 53 bones of rails (Rallidae), previously referred to Gal-lirallus n. spp., from archaeological sites on four islands in the Marquesas Is-lands, French Polynesia. We describe three new, extinct, flightless species ofGallirallus: G. roletti (Tahuata), G. gracilitibia (Ua Huka), and G. epulare (NukuHiva). Two bones from Hiva Oa, although probably representing another ex-tinct species of Gallirallus, are regarded as an inadequate basis for describing aspecies. At first human contact, the genus Gallirallus probably included manyscores if not hundreds of flightless species on islands from the far western Pa-cific (Okinawa, Philippines, Halmahera) eastward across most of Oceania. Ascurrently understood, the Marquesas Islands represent the eastern range limitof Gallirallus.

Bird bones from Pacific islands (Figure 1)have revealed extensive Late Holocene ex-tinctions of land birds following prehistorichuman arrival ( James and Olson 1991, Olsonand James 1991, Steadman 1995, 2006,Worthy and Holdaway 2002). Radiocarbonchronologies of stratified zooarchaeologicalassemblages indicate that, in the earlieststages of occupation, people consumed a rel-atively large number of indigenous birds(Steadman and Rolett 1996, Steadman et al.2002). In East Polynesia, from 50 to 100%of the species of land birds were extirpatedfrom most islands. Especially prevalentamong the extinct species are rails (OrderGruiformes, Family Rallidae), most of whichwere flightless species endemic to single is-

lands. These ground-nesting birds were espe-cially vulnerable to predation by people andtheir introduced rats, pigs, and dogs.

Nearly all of the known extinct (y) speciesof flightless rails have been placed in wide-spread, extant genera, including 7–10 speciesof Porzana from the Hawaiian Islands alone(Olson and James 1991); one to several spe-cies each of Gallirallus, Gallinula, Porphyrio,and Fulica from New Zealand (Worthy andHoldaway 2002); and many species of Por-zana, Gallirallus, and Porphyrio from the restof Oceania (Steadman 1995, 2006, Kirchmanand Steadman 2005, 2006). Thus flightless-ness, which became terminally maladaptiveonly after Polynesians and their commensalscolonized the Pacific, evolved independentlyin insular rails numerous times, even withinarchipelagos. Of the crudely estimated hun-dreds (Pimm et al. 1994, Livezey 2003) to500 to 1,600 (Steadman 2006) to nearly2,000 (Steadman 1995) species of flightlessrails that once existed across the Pacific, fewerthan 20 have been described (Worthy 2004).Here we describe the prehistoric bones ofrails from archaeological sites on four islandsin the Marquesas Islands.

Marquesan Archaeological Sites

Centered at ca. 9� S and 140� W, the Mar-quesas Islands lie ca. 500 km northwest of theTuamotu Archipelago and 1,400 km north-

Pacific Science (2007), vol. 61, no. 1:145–163: 2007 by University of Hawai‘i PressAll rights reserved

1 Financial support came from a McGlaughlin Fel-lowship to J.J.K. from the University of Florida Collegeof Liberal Arts and Sciences, and NSF grants EAR-9714819 and DEB-0228682 to D.W.S. Manuscript ac-cepted 14 February 2006.

2 New York State Museum, 3140 CEC, Albany, NY12230.

3 Florida Museum of Natural History, University ofFlorida, Gainesville, Florida 32611.

4 Corresponding author (phone: 518-474-1441;e-mail: [email protected]).

Figure 1. Oceania, showing location of the Marquesas Islands and location of the four archaeological sites discussed in this paper.

east of the Society Islands, which are thenearest high islands (Figure 1). The Marque-sas consist of 10 major eroded volcanic is-lands with surface areas that range from <2to 330 km2 and maximum elevations from420 to 1,252 m. Minimum interisland dis-tances among these 10 islands range from 3to 21 km. Hiva Oa and nearby Tahuata prob-ably were connected to each other duringthe last Pleistocene glacial interval, whereasNuku Hiva and Ua Huka remained as sepa-rate islands even when the sea level was 120–130 m lower than at present.

The archaeological specimens reportedhere are 53 bones that were previously re-ferred to ‘‘Gallirallus new spp.’’ or ‘‘Gallirallusundescribed spp.’’ by Steadman and Rolett(1996) and Steadman (1989, 2006). The sitesrepresent early human occupations developedin coastal calcareous sands on four islands.The Hanamiai site on Tahuata was excavatedby B. V. Rolett in 1984–1985 (Rolett 1998).The sediment excavated at Hanamiai wasscreen-washed through 1/8-inch (3.2-mm)mesh, producing 716 identifiable bird bones,among which are 70 land bird bones from 10species (Steadman and Rolett 1996). The Ha-natekua Rockshelter on Hiva Oa was exca-vated in 1967–1968 by Y. H. Sinoto and P.Bellwood (Bellwood 1972), yielding 146 landbird bones from seven species (Steadman2006). The Hane Dune site (often calledmerely the Hane site) on Ua Huka, excavatedby Y. H. Sinoto in the 1960s (Sinoto 1966,1970, 1979), is the richest Marquesan site interms of faunal remains, having yielded morethan 12,000 identifiable bird bones, including2,187 land bird bones from 17 species (Stead-man 2006). The Ha‘atuatua Dune site (oftencalled merely the Ha‘atuatua site) on NukuHiva was excavated initially by R. Suggs in1956 and 1958 (Suggs 1961) and in 1992–1994 by B. V. Rolett and E. Conte (Rolett1998). The combined excavations at Ha‘atua-tua Dune yielded 27 bones of land birds be-longing to nine species (Steadman 2006).

Based on radiocarbon (14C) dates from re-liable stratigraphic levels, the initial humanoccupation for Tahuata, Hiva Oa, and NukuHiva was ca. 1,000 yr B.P. (years before pres-ent) and ca. 1,350 yr B.P. for Ua Huka (Ro-

lett 1998). Hane Dune (Ua Huka) is theearliest well-dated site in the Marquesas. Esti-mates that people arrived in the Marquesas asearly as 2,000 yr B.P. (e.g., Sinoto 1979) arefrom single 14C samples from poorly estab-lished stratigraphic contexts or are based on14C dates with suspected laboratory errors(Rolett 1998).

materials and methods

Skeletons used for comparison with fossils arefrom the American Museum of Natural His-tory (amnh); Bernice P. Bishop Museum(bpbm); Florida Museum of Natural History,University of Florida (uf); National Museumof Natural History, Smithsonian Institution(usnm); National Museum of New Zealand(nmnz); Department de Archeologie, CentrePolynesien des Sciences Humaines, Tahiti(dapt); University of Michigan Museum ofZoology (ummz); Thomas Burke MemorialMuseum, University of Washington (uwbm);and Yale Peabody Museum (ypm). We exam-ined these modern specimens: Porzana tabu-ensis, uwbm 42501, 42528; Rallus longirostris,uf 40956, 24200; Gallirallus striatus, usnm

85892, 343214, 559919, ypm 107205; G. tor-quatus, ummz 228275, 228279, 228280, amnh

17715–17717, usnm 290445; G. owstoni, uf

39918–39921, 39256, 42968, ummz 215472,usnm 561968, 611816, 612616, 613738–614744, 614233–614235, 614771, 614772; G.australis, uf 24326, 24327, ypm 102249,110760, 110789, 110790, 110844; G. philip-pensis, uf 39854, 39855, 42902, 42935, 42934,42933, 43224, uwbm 42865, 42866, 42863,usnm 560651, 560791, 620196; G. [‘‘Nesoclo-peus’’] woodfordi, uf 39399, 39406, 39409,39547, 39556, 39574; Amaurornis olivaceus, uf

40216; A. phoenicurus, uf 24387; Porphyrioporphyrio, uf 39388, 39407; P. martinicus, uf

39927, 42418, 42419; Gallinula chloropus, uf

39927. We also examined these fossil speci-mens: yGallirallus huiatua tarsometatarsus,nmnz S37708 (holotype), ulna, nmnz S37709(paratype), femur, nmnz S37710 (paratype),tibiotarsus, nmnz S37711 (paratype); yG. rip-leyi rostrum, uf 55596, coracoid, uf 54711,usnm 402896 (paratype), humeri, uf 51402,55752, ulnae, uf 55215, 54901, femur, uf

Fossil Rails from the Marquesas Islands . Kirchman and Steadman 147

51320, tibiotarsi, uf 55283, 59574, usnm

402895 (holotype), tarsometatarsi, uf 55223,usnm 402895 (holotype); yG. storrsolsoni ros-tra, bpbm 166036 (holotype), dapt 21, hu-merus, bpbm 166022, ulnae, bpbm 166033,femur, dapt 27/105, tibiotarsi, bpbm 166023,dapt 119, tarsometatarsi, bpbm 166034, dapt

7; yG. vekamatolu rostrum, uf 52292, mandi-bles, uf 51836, 52525, coracoids, uf 52204,52966, scapulae, uf 52179, 52318, humeri,uf 52333, 52707, ulna, uf 51734, femora, uf

52058, 52518, tibiotarsi, uf 51729, 52211,tarsometatarsi, uf 52002, 52137; yPorphyriopaepae femora, bpbm 165649, 166434, tibio-tarsus, bpbm 165651; and yP. mcnabi femora,bpbm 166031 (holotype), dapt 39, dapt 53(paratypes).

Measurements were taken with electronicdigital calipers and rounded to the nearest0.1 mm. Osteological terminology followsBaumel and Witmer (1993).

comparative osteology and

systematics

Family Rallidae

The phylogeny of genera in the Rallidae isnot well resolved. As a starting point forgenus-level classification, we regard all ‘‘typ-ical long-billed rails’’ (as opposed to swamp-hens, moorhens, coots, and crakes) fromOceania as species of Gallirallus sensu lato.We agree with Worthy (2004) that the re-cently discovered fossil rail yVitirallus watlingifrom Fiji belongs in a separate genus despitehaving a long (though decurved) bill andsharing other osteological similarities withGallirallus. Our classification agrees with thatof Olson (1973) by including in Gallirallusthe species australis, philippensis, owstoni, okina-wae, ywakensis, torquatus, insignis, sylvestris,ydieffenbachii, ymodestus, striatus, and pectoralis.Among these species, only G. philippensis, G.torquatus, G. striatus, and G. pectoralis are vol-ant. Olson (1973) provisionally retained G.woodfordi and yG. poecilopterus in Nesoclopeusbut highlighted their close affinity with Gal-lirallus, within which we classify these twospecies. This treatment differs from that ofLivezey (1998, 2003), who acknowledged the

difficulty of establishing relationships in thisgroup solely on the basis of osteology butwho divided the 14 species mentioned hereamong Gallirallus, Nesoclopeus, Tricholimnas,Cabalus, and Habropteryx.

As defined herein, Gallirallus includes atleast 17 named flightless species endemic tosingle islands or islands connected during pe-riods of lowered sea level. This total includeseight extant species (G. australis of New Zea-land; G. sylvestris of Lord Howe Island; G.owstoni of Guam; G. okinawae of Okinawa;G. insignis of New Britain; G. woodfordi s.l. ofthe Solomon Islands [minimally Bougainville,Santa Isabel, and Guadalcanal]; G. rovianae ofNew Georgia, Solomon Islands; and G. cal-ayanensis of Calayan, Philippines), five histor-ically extinct species (yG. wakensis of WakeIsland; yG. lafresnayanus of New Caledonia;yG. modestus of the Chatham Islands, NewZealand; yG. poecilopterus of Viti Levu, Fiji;and yG. pacificus of Tahiti, Society Islands),and four previously named prehistoric species(yG. huiatua from noncultural Holocene cavedeposits on Niue [Steadman et al. 2000]; yG.ripleyi from cultural and noncultural late Hol-ocene sites on Mangaia, Cook Islands [Stead-man 1987]; yG. vekamatolu from preculturalstrata on ‘Eua, Kingdom of Tonga [Kirch-man and Steadman 2005]; and yG. storrsolsonifrom cultural deposits on Huahine, SocietyIslands [Kirchman and Steadman 2006]).

Genus Gallirallus Lafresnaye, 1841

We refer the 53 fossils from the MarquesasIslands to Gallirallus rather than to other gen-era of oceanic rails on the basis of the follow-ing characters. Rostrum: long, narrow, andshallow with elongate nares. Mandible: cotylalateralis narrow and concave, fossa for condy-lus medialis quadratum (the main articulationsurface in the os articulare) shallow and wide.Coracoid: acrocoracoid extends medially overthe sulcus musculo supracoracoidei such thatthe foramen triosseum is less open cranially.Scapula: facies articularis clavicularis rela-tively small and oriented at a more obtuseangle from corpus scapulae. Humerus: fossapneumotricipitalis deep and wide with prom-inent crus ventrale fossae. Ulna: thin in

148 PACIFIC SCIENCE . January 2007

cranial aspect with rectangular (rather thanrounded) margo cranialis. Pelvis: ala preaceta-bularis ilii broadly continuous with cristadorsalis of synsacrum. Femur: distal end ofcorpus femoris becomes gradually wider; con-dylus medialis subcircular in medial aspect;impresso ansae musculo iliofibularis abuts su-clus fibularis; rotolar groove broad. Tibiotar-sus: craniolateral and craniomedial margins ofcorpus tibiotarsi rounded rather than sharp;impresso ligamentum collateralis medialisdeep and wide; facies articularis femoris large;depressio epicondylaris lateralis deep; condy-lus medialis subcircular in medial aspect. Tar-sometatarsus: corpus tarsometatarsi muchwider than deep; medial sulcus hypotarsi notenclosed; fossa parahypotarsalis medialis shal-low in proximal aspect; fossa metatarsi I shortand shallow; crista plantaris mediana slopesgradually (not steeply) to hypotarsus; distalend of trochlea metatarsi tertii sloped towardmedial trochlea; cotyla medialis is rectangularin proximal aspect with flat (not rounded)dorsal margin.

yGallirallus roletti Kirchman & Steadman, n.sp.Figures 2A, 3A, 4A, 5A, 6A

holotype. Associated complete right fe-mur, bpbm 166447; right tibiotarsus, bpbm

166446; right tarsometatarsus, bpbm 166448;and pedal phalages, bpbm 166449, 166450(Figure 2A). From the Hanamiai archaeolog-ical site, Tahuata, Marquesas Islands. Col-lected by B. V. Rolett and colleagues in1984–1985.

paratypes. Distal rostrum, bpbm 166456(Figure 3A); left articular, bpbm 166458; rightarticular, bpbm 166542; distal dentaries, bpbm

166436, 166439, 166457; left coracoid (hu-meral end), bpbm 166455; left proximal fe-mur, bpbm 166435; right femur shaft, bpbm

166445; left distal tibiotarsus, bpbm 166444;left tibiotarsi shafts, bpbm 166437, 166452;right distal tibiotarsus, bpbm 166438; leftdistal tarsometatarsus, bpbm 166451; nearlycomplete right tarsometatarsus, bpbm 166440;right proximal tarsometatarsus, bpbm 166454;right tarsometatarus shaft, bpbm 166441. Allare from the same locality as the holotype.

diagnosis. A medium-sized, flightlessspecies of Gallirallus (Table 1) distinguishedfrom all examined congeners (except wheresimilarities are noted) as follows. Rostrum(Figure 3A): more robust and deep withheight to width ratio (at anterior margin ofnares) of 1.24 (a1.03 in all other species ofGallirallus); in ventral aspect, trough in ospremaxilare deep and wide, crista tomialisthin and sharp. Mandible: in caudal aspect,fossa caudalis with straight lateral and medio-ventral sides and a deep, narrow groove ontop (ventral) side; pars symphysialis long,with rami sloped steeply to form a V-shaped(rather than U-shaped) trough in cranial as-pect. Femur (Figures 2A, 4A): mediodistalmargin of neck deeply excavated below faciesarticularis acetabularis, forming a sulcus inanterior aspect; in ventral aspect, trochanterfemoris forms a prominent right angle withfacies articularis antitrochanterica; the mostproximal impressiones obturatoriae deep andlong, forming a groove parallel to the cristatrochanteris; in medial aspect, corpus femorusstout and straight, especially on leading edge(facies cranialis). Tibiotarsus (Figures 2A,5A): fossa retropatellaris narrow but deep;impresso ligamentum collateralis medialisshallow; fossa flexoria shallow; crista fibularisshort but broad, projecting farther from cor-pus tibiotarsis at distal end than in all exceptG. owstoni; tuberculum retinaculi musculo fi-bularis prominent (as in yG. storrsolsoni andG. torquatus) but not forming a tube (as in G.woodfordi); condylus medialis and condyluslateralis large relative to width and depth ofcorpus tibiotarsus. Tarsometatarsus (Figures2A, 6A): proportionately stout, although notso much as in yG. vekamatolu or yG. ripleyi;sulcus extensorius deeply concave; corpus tar-sometatarsi shallow relative to its breadth; fa-cies medialis thin in medial aspect, especiallyproximally (approaching the condition inAmaurornis olivaceus or Porphyrio spp.); sulcusproximal to the foramen vasculare distaleshort and shallow; distal trochleae large andwidely splayed.

etymology. Named in honor of BarryV. Rolett, whose outstanding research in theMarquesas Islands has been of great impor-tance to both biologists and archaeologists.


Figure 2. A, yGallirallus roletti holotype consisting of associated femur (bpbm 166447), tibiotarsus (bpbm 166446),tarsometatarsus (bpbm 166448), and two pedal phalanges (bpbm 166449, 166450), top to bottom, respectively, Tahuata,Marquesas Islands. Shown with the same elements from B, G. philippensis (uf 39855, Tutuila, Samoa), and C, G. owstoni(uf 39921, Guam, Mariana Islands). Scale bar ¼ 5 cm.

In particular, his careful excavations at theHanamiai site yielded the most extensive anduseful series of Gallirallus bones from theMarquesas Islands.

remarks. In overall size, yGallirallus ro-letti resembles G. owstoni, G. philippensis, G.striatus, and G. torquatus. It is larger than yG.ripleyi and yG. wakensis, and smaller thanG. australis, yG. vekamatolu, and G. woodfordi.The material from Hanamiai represents fourindividuals, minimally. Lacking the sternum,scapula, and elements of the forelimb, ouronly evidence that yG. roletti was flightless

comes from the relatively small size of thecoracoid (bpbm 166455) and the larger, moreopen shape and more medial position of thecotyla scapularis on the dorsal surface ofthe coracoid, which more closely resemblesthe condition in flightless rather than volantspecies of Gallirallus. This specimen is pittedover much of its surface, indicating that itmay represent a juvenile bird. Comparing thiscoracoid among those of known juveniles andadults in G. philippensis (volant) and G. owstoni(flightless) suggests that bpbm 166455 is froma bird 3 to 4 months old and that the cora-

Figure 3. Rostra of A, yGallirallus roletti (bpbm 166456, Tahuata, Marquesas Islands); B, yG. storrsolsoni (holotype,bpbm 166036, Huahine, Society Islands); C, G. philippensis (uf 39855, Tutuila, Samoa); and D, G. owstoni (uf 39921,Guam, Mariana Islands) in lateral (left) and dorsal (right) aspects. Scale bars ¼ 2 cm.


Figure 4. Femora of A, yGallirallus roletti (holotype, bpbm 166447, Tahuata, Marquesas Islands); B, cf. yGallirallus sp.(bpbm 168539, Hiva Oa, Marquesas Islands); C, yG. gracilitibia (bpbm 176974, Ua Huka, Marquesas Islands); D, yG.epulare (bpbm 181659, Nuku Hiva, Marquesas Islands); and E, G. philippensis (uf 39855, Tutuila, Samoa) in dorsal(top) and ventral (bottom) aspects. Scale bars ¼ 5 cm.

Figure 5. Tibiotarsi of A, yGallirallus roletti (holotype, bpbm 166446, Tahuata, Marquesas Islands); B, yG. storrsolsoni(bpbm 166023, Huahine, Society Islands); C, yG. epulare (bpbm 181661, Nuku Hiva, Marquesas Islands); D, yG. graci-litibia (holotype, bpbm 166013/176387, Ua Huka, Marquesas Islands); E, G. philippensis (uf 39855, Tutuila, Samoa); andF, G. owstoni (uf 39921, Guam, Mariana Islands) in dorsal (top) and ventral ( bottom) aspects. Scale bars ¼ 5 cm.

Figure 6. Tarsometatarsi of A, yGallirallus roletti (holotype, bpbm 166448, Tahuata, Marquesas Islands); B, yG. storrs-olsoni (dapt 7, Huahine, Society Islands); C, yG. epulare (bpbm 167119, Nuku Hiva, Marquesas Islands); D, G. philip-pensis (uf 39855, Tutuila, Samoa); and E, G. owstoni (uf 39921, Guam, Mariana Islands) in acrotarsial (top) and plantar(bottom) aspects. Scale bars ¼ 5 cm.

Figure 7. Humeri of A, yGallirallus storrsolsoni (bpbm 166022, Huahine, Society Islands); B, yG. epulare (bpbm 181657,Nuku Hiva, Marquesas Islands); C, yG. gracilitibia (bpbm 163130, Ua Huka, Marquesas Islands); D, G. philippensis(uf 39855, Tutuila, Samoa); and E, G. owstoni (uf 39921, Guam, Mariana Islands) in ventral (top) and dorsal (bottom)aspects. Scale bars ¼ 5 cm.

coid’s small size (relative to leg elements) isdue to flightlessness rather than to the age ofthe bird.

Steadman and Rolett (1996) referred 24specimens to ‘‘Gallirallus new sp. (TahuataRail),’’ which is named herein as G. roletti,based on 22 specimens. The discrepancy isaccounted for by an ungual phalanx ofyPorphyrio paepae (bpbm 166442) being mis-takenly listed among the specimens of ‘‘Gal-lirallus new sp.’’ in Steadman and Rolett

(1996) and by another pedal phalanx (bpbm

166453) whose identity cannot now be deter-mined.

yGallirallus gracilitibia Kirchman & Steadman,n. sp.Figures 4C, 5D, 7C

holotype. Right tibiotarsus lackingproximal end, bpbm 166013, 176387 (Figure5D; two pieces with different catalog numbers

TABLE 1

Skeletal Measurements (in mm) in yGallirallus roletti and Select Congeners, with Mean, Range, and Sample Size

Parameter

yG.roletti

U

yG.storrsolsoni

U

G.owstoni

M

G.owstoni

F

G.australis

M

G.australis

F

yG.ripleyi

U

yG.vekamatolu

U

Rostrum: heightat anteriormargin ofnares

5.21

3.73.5–3.92

3.43.2–3.85

3.33.1–3.54

5.35.1–5.43

4.74.4–5.04

2.91

4.81

Rostrum: widthat anteriormargin ofnares

4.21

4.14.0–4.22

4.44.0–4.75

4.23.8–4.54

5.55.1–5.73

5.04.5–5.54

2.81

5.91

Femur: length 54.51

49.41

56.054.0–59.2

7

52.449.2–54.611

80.678.7–82.3

3

69.966.1–73.4

4

39.81

—

Femur: distalwidth

9.81

9.21

9.58.9–9.97

8.78.4–9.0

11

16.515.9–17.6

3

14.313.2–15.0

4

6.71

10.910.7–11.0

2Femur: minimum

shaft width4.04.02

3.91

3.83.3–4.07

3.63.4–3.9

11

6.76.5–6.93

5.65.1–5.94

3.01

—

Tibiotarsus:length

79.51

68.61

79.976.1–84.6

7

75.572.9–79.811

117.4116.6–118.7

3

100.093.4–105.0

4

— —

Tibiotarsus:distal width

7.87.7–7.92

6.91

7.46.8–7.97

6.86.5–7.1

11

12.512.1–13.1

3

10.810.4–11.2

4

6.05.6–6.33

8.07.9–8.02

Tarsometatarsus:proximal width

7.91

7.41

7.67.0–8.27

7.06.7–7.3

11

12.612.2–13.4

3

11.010.7–11.7

4

5.71

8.37.8–8.72

Tarsometatarsus:distal width

8.71

— 8.07.2–8.57

7.47.2–7.8

11

14.013.6–14.3

3

12.111.2–13.0

4

6.46.3–6.53

8.58.0–9.02

Tarsometatarsus:shaft width

3.5–3.93.72

4.11

3.83.5–4.07

3.43.2–3.8

11

6.15.6–6.43

5.45.2–5.64

3.02.8–3.12

4.34.0–4.62

Tarsometatarsus:shaft depth

2.72.5–2.82

2.81

2.82.5–3.07

2.52.4–2.8

11

4.84.5–5.03

4.13.8–4.64

2.22.1–2.32

3.23.0–3.32

Note: Specimens of all available subspecies of G. australis and G. philippensis are combined, given that subspecific differences in sizeare much smaller than those between males and females. F, female; M, male; U, sex unknown; —, cannot be evaluated.


glued together). From the Hane Dunearchaeological site, Ua Huka, Marquesas Is-lands. Collected by Y. H. Sinoto and col-leagues on 11 October 1965.

paratypes. Right coracoid (humeralend), bpbm 166015; left proximal humerus,bpbm 163130 (Figure 7C ); right humerusshaft, bpbm 166014; right femur shaft, bpbm

176974 (Figure 4C ); left distal tibiotarsi,bpbm 163240, 166016, 169146, 170909; lefttibiotarsus shaft, bpbm 175384; right distal

tibiotarsi, bpbm 166008, 166012, 169145,175145; right tibiotarsi shafts, bpbm 171294,175322, 175395, 176725, 176746, 176971,176972. All are from the same locality as theholotype. A minimum of eight individuals isrepresented.

diagnosis. A small to medium-sized(Table 2), flightless species of Gallirallus thatdiffers from congeneric species as follows.Coracoid: sulcus musculo supracoracoidei rel-atively shallower and wider than in volant

TABLE 1(continued)

G.philippensis

M

G.philippensis

F

G.striatus

M

G.torquatus

M

G.torquatus

F

G.woodfordi

M

G.woodfordi

F

3.12.8–3.45

2.82.6–2.95

2.81

3.23.1–3.32

— 4.34.2–4.63

4.23.9–4.43

3.73.4–4.25

3.23.1–3.45

3.11

3.43.4–3.42

— 4.74.6–4.93

4.64.5–4.83

53.851.3–54.9

6

47.345.0–49.7

6

45.142.2–46.9

4

55.553.3–57.5

5

50.61

72.172.0–72.3

3

69.669.0–70.3

38.78.4–9.16

7.47.0–7.96

45.142.2–46.9

4

9.38.6–9.95

8.51

13.313.2–13.4

3

12.712.5–12.9

34.03.7–4.26

3.33.1–3.56

2.92.8–3.14

3.93.5–4.15

3.21

5.35.2–5.33

4.84.6–5.13

76.971.7–80.3

6

66.763.0–70.6

6

62.158.2–64.3

4

84.280.9–87.5

5

74.61

105.0100.6–108.8

3

102.899.0–105.8

36.96.5–7.26

6.15.6–6.76

5.35.1–5.64

7.26.7–7.65

6.61

10.09.9–10.13

9.59.4–9.63

7.06.7–7.26

6.15.8–6.66

5.45.3–5.54

7.47.0–7.95

6.61

10.510.2–10.7

3

10.09.9–10.13

7.16.7–7.56

6.36.0–6.96

5.65.4–5.74

7.87.3–8.45

7.01

10.810.7–11.0

3

10.610.5–10.6

33.53.1–3.66

3.02.8–3.36

2.72.5–2.94

3.63.3–3.85

3.31

4.94.8–5.03

4.64.4–4.73

2.82.6–3.06

2.52.3–2.86

2.12.0–2.24

2.72.2–2.95

2.61

4.03.9–4.23

3.83.7–4.03


species. Humerus (Figure 7C ): crista bicipita-lis small; corpus humeri thin and curved; dis-tal junction of crista pectoralis and corpushumeri abrupt, rather than gradually sloping;sulcus ligamentum transversus deep. Femur(Figure 4C ): corpus femoris gracile. Tibiotar-sus (Figure 5D): corpus tibiotarsus slenderrelative to its length; depressio epicondylarismedialis deep.

etymology. From the Latin words graci-lis (slender, slim, thin) and tibia (the shinbone,tibia); see Brown (1956:469, 791). The namegracilitibia is a feminine noun in appositionto Gallirallus. It refers to the distinctivelyslender tibiotarsus in this species, especiallycompared with that of yG. roletti.

remarks. yGallirallus gracilitibia is a smallto medium-sized species (Table 2), exceededin stoutness of skeletal elements by all congen-ers but resembling the medium-sized G.owstoni, G. philippensis, and G. torquatus inmeasurements along the long axes of leg ele-ments. Based on length-to-width ratios of ti-biotarsi, yG. gracilitibia has the thinnest shaftrelative to length of any species of Gallirallus,whether flightless or volant (Table 2). De-scribed features of the coracoid and humerusof yG. gracilitibia, as well as their size relativeto hind limb elements (see Livezey 2003,

Kirchman and Steadman 2005, 2006), indi-cate that this species was flightless.

yGallirallus epulare Kirchman & Steadman, n.sp.Figures 4D, 5C, 6C, 7B, 8B

holotype. Nearly complete left ulna,bpbm 181658 (Figure 8B). From the Ha‘atua-tua archaeological site, Nuku Hiva, Marque-sas Islands. Collected by B. V. Rolett, E.Conte, and colleagues in 1994–1995.

paratypes. Left humerus shaft, bpbm

181657 (Figure 7B); left femur shaft, bpbm

181659 (Figure 4D); left tibiotarsi shafts,bpbm 181660, 181661 (Figure 5C ), 181662;right tibiotarsus shaft, bpbm 167191; left distaltarsometatarsus, bpbm 167119 (Figure 6C ).All are from the same locality as the holotype.

diagnosis. A small species of Gallirallusdistinguished from congeneric species asfollows. Humerus and ulna very small andslender relative to leg elements, especiallycompared with those of yG. gracilitibia fromnearby Ua Huka. Ulna: straighter than in vol-ant species. Tarsometatarsus: facies dorsalisof corpus tarsometatarsi highly convex proxi-mal to trochlea metatarsi III; foramen vascu-lare distale relatively large and round.

TABLE 2

Skeletal Measurements (in mm) of yGallirallus gracilitibia and Selected Congeners with Mean, Range, and Sample Size

Parameter

yG.gracilitibia

U

yG.storrsolsoni

U

G.philippensis

M

G.philippensis

F

G.owstoni

M

G.owstoni

F

G.torquatus

M

G.torquatus

F

Humerus: shaftwidth

2.22.1–2.32

2.21

3.33.2–3.45

2.92.7–3.36

2.82.6–3.17

2.72.6–3.0

11

3.23.0–3.45

2.91

Tibiotarsus: shaftwidth

3.02.9–3.38

3.43.2–3.73

3.63.3–4.06

3.12.9–3.36

4.13.8–4.57

3.83.4–4.1

11

3.83.4–4.05

3.31

Tibiotarsus: lengthto fibular cresta

54.51

50.250.2–50.2

2

52.851.9–55.3

5

47.143.3–51.6

6

57.855.2–61.2

7

54.452.9–55.911

59.3–61.52

59.01

Tibiotarsus: distalwidth

6.46.3–6.54

6.91

6.96.5–7.26

6.15.6–6.76

7.46.8–7.97

6.86.5–7.1

11

7.26.7–7.65

6.61

Note: F, female; M, male; U, sex unknown.a Distal end of bone to distal edge of fibular crest.


etymology. From the Latin epularis(pertaining to a banquet, belonging to a ban-quet [Brown 1956:292]). The name epulare isan adjective that modifies the masculine Gal-lirallus. It refers to the archaeological contextin which the specimens of yG. epulare werefound, namely that of a kitchen midden do-minated by the bony and shelly remains offoods eaten by prehistoric Polynesians.

remarks. The humerus, femur, and ti-biotarsi lack both the proximal and distalends and therefore also lack diagnosticfeatures. The most striking feature ofyGallirallus epulare is its tiny wing elementsrelative to its leg elements; the leg elementsclosely resemble those from a small female

G. owstoni, but the humerus and ulna aremuch smaller than those of G. owstoni and in-deed more closely resemble those of yG. rip-leyi and yG. wakensis.

cf. yGallirallus sp.Figures 4B, 9A

material. Complete left scapula, bpbm

165655 (Figure 9A); shaft of right femur,bpbm 168539 (Figure 4B). From the Hanate-kua Rockshelter archaeological site, Hiva Oa,Marquesas Islands. Collected by Y. H. Sinotoand colleagues in 1965–1966.

remarks. These two bones represent alarge, probably flightless species of Gallirallus.

Figure 8. Ulnae of A, yGallirallus storrsolsoni (bpbm 166033, Huahine, Society Islands); B, yG. epulare (holotype, bpbm

181658, Nuku Hiva, Marquesas Islands); C, G. philippensis (uf 39855, Tutuila, Samoa); and D, G. owstoni (uf 39921,Guam, Mariana Islands) in dorsal aspect. Scale bar ¼ 5 cm.


We regard these bones as an inadequate basisfor describing a new species but note that thescapula differs from that in all congenericspecies in having a relatively thick area be-tween the facies articularis humeralis andfacies articularis clavicularis, in proximal as-pect, and in having a distinctive flange onventral surface of corpus scapulae near ex-tremitas caudalis. The femur lacks both theproximal and distal ends but is remarkable inhaving a relatively wide divergence of the twolinea intermuscularis caudalis.

The scapulae of the three named Marque-san species of Gallirallus and of yPorphyriopaepae (formerly of Tahuata and Hiva Oa) areunknown, thus precluding direct comparisonwith bpbm 165655. The femoral shaft fromHiva Oa (bpbm 168539 [Figure 4B]) is toolarge to belong to yP. paepae and also is largerthan that of yG. roletti, suggesting that theGallirallus species from Hiva Oa was quitelarge and that Gallirallus from the southernislands were larger than those of the twonorthern islands.

discussion

The rail bones from archaeological sites onthree Marquesan islands (Tahuata, NukuHiva, and Ua Huka) represent three newflightless species of Gallirallus, one endemicto each island. A probable fourth species,from Hiva Oa, requires more material beforea description is possible. No species of Galli-rallus exist anywhere today in East Polynesia,where all known species are flightless andextinct. They are yG. ripleyi from Mangaia,Cook Islands; yG. storrsolsoni from Huahine,Society Islands ( both known only from bonesfrom archaeological sites [Steadman 1987,Kirchman and Steadman 2006]); and yG. pa-cificus from Tahiti, Society Islands, which sur-vived to the eighteenth century (Ripley 1977,Taylor 1998).

The species from the Marquesas Islandsrepresent the northwestern limit of Galliral-lus. The very rich prehistoric record of birds(15,000þ bones from 12 archaeological sites)from Henderson Island (Pitcairn Group) has

TABLE 3

Skeletal Measurements (in mm) of yGallirallus epulare and Selected Congeners with Mean, Range, and Sample Size

Parameter

yG.epulare

U

yG.storrsolsoni

U

G.philippensis

M

G.philippensis

F

G.owstoni

M

G.owstoni

F

G.torquatus

M

G.torquatus

F

Ulna: length 25.5*1

37.51

43.941.4–44.8

6

39.235.2–43.6

6

39.136.8–41.5

7

36.634.8–38.711

44.043.0–45.3

5

40.01

Ulna: shaft width 2.01

2.71

3.02.8–3.16

2.72.6–2.86

3.02.8–3.27

2.72.6–3.0

11

3.02.8–3.15

2.81

Tibiotarsus: shaftwidth

3.41

3.43.2–3.73

3.63.3–4.06

3.12.9–3.36

4.13.8–4.57

3.83.4–4.1

11

3.83.4–4.05

3.31

Tarsometatarsus:shaft width

3.51

4.11

3.53.1–3.66

3.02.8–3.36

3.83.5–4.07

3.43.2–3.8

11

3.63.3–3.85

3.31

Tarsometatarsus:shaft depth

2.41

2.81

2.82.6–3.06

2.52.3–2.86

2.82.5–3.07

2.52.4–2.8

11

2.72.2–2.95

2.61

Tarsometatarsus:distal width

6.51

— 7.16.7–7.56

6.36.0–6.96

8.07.2–8.57

7.47.2–7.6

11

7.87.3–8.45

7.01

Note: F, female; M, male; U, sex unknown; —, cannot be evaluated.* Estimated.


not yielded evidence of Gallirallus (Steadmanand Olson 1985, Wragg 1995). Between theCook Islands (where yG. ripleyi lived) andHenderson Island lie the Austral (Tubuai) Is-lands, where the single prehistoric land birdbone known represents an undescribed,extinct species of Ptilinopus (Columbidae)(Steadman 2006). Likewise, the well-studiedfossils from dunes, lava tubes, and lakes inthe Hawaiian Islands, which have yielded7–10 flightless species of Porzana rails, lackevidence of Gallirallus (Olson and James1991).

Aside from the species of Gallirallus, ex-tinct species of Marquesan land birds known

only from prehistoric bones are the swamp-hen yPorphyrio paepae, an undescribedspecies of sandpiper (Prosobonia), two doves(yGallicolumba nui and yMacropygia heana),and two species of parrots (yVini vidivici andyV. sinotoi) (Steadman and Zarriello 1987,Steadman 1988, 1992, 2006). The extinctionof flightless rails in the Marquesas probablytook place in prehistoric times. On Tahuata,all but 1 of the 22 specimens of yGallirallusroletti were excavated from Levels G, GH, orH (Phases I and II) at Hanamiai, which rep-resent the early occupation of the site at ca.1,000–700 yr B.P. (Rolett 1998). A singlebone of yG. roletti is from Phase III (Level

Figure 9. Scapulae of A, cf. yGallirallus sp. (bpbm 165655, Hiva Oa, Marquesas Islands); B, G. philippensis (uf 39855,Tutuila, Samoa); and C, G. owstoni (uf 39921, Guam, Mariana Islands) in dorsal aspect. Scale bar ¼ 5 cm.


F), which dates to 700–550 yr B.P. This maybe the approximate time of extinction for yG.roletti, bones of which were not recovered inthe younger Levels A–D (Phases IV, V ).Similarly on Ua Huka, all of the bones ofyG. gracilitibia were from strata dated to>800 yr B.P. (Steadman 1991).

acknowledgments

For access to modern and fossil specimens,we thank Paul Sweet (amnh), James Deanand Storrs Olson (usnm), Carla Kishinami(bpbm), Alan Tennyson (nmnz), Barry Rolett(University of Hawai‘i), Tom Webber andAndrew Kratter (uf), Janet Hinshaw (ummz),Sievert Rohwer and Chris Wood (uwbm), andKristof Zyskowski (ypm). For helpful com-ments on the manuscript, we thank RichardHoldaway and an anonymous reviewer.

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New Species of Extinct Rails (Aves: Rallidae) from ... · 145 New Species of Extinct Rails (Aves: Rallidae) from Archaeological Sites in the Marquesas Islands, French Polynesia1 Jeremy