Ducula shutleri sp. nov. Hypotaenidia vavauensis sp. nov. (1 Viewer)

TREVOR H. WORTHY and DAVID V. BURLEY, 2019

Prehistoric avifaunas from the Kingdom of Tonga

Zoological Journal of the Linnean Society, 2019, XX, 1–48. With 5 figures

Free pdf: https://watermark.silverchair.com/z...SRjknchdsFOqdSXuzPmKmgZQ1ATkZujth36fV9GKl-t7A

Abstract:

Avifaunas derived from Lapita archaeological sites excavated between 2004 and 2014 from four sites in the Vava'u Group and two on Tongatapu, Kingdom of Tonga are described, revealing birds encountered by the first human arrivals. A total of 741 identifiable bones revealed 24 avian taxa, among which terrestrial birds, especially rails, pigeons and parrots, were the most abundant. At a minimum, eight taxa, or 50% of the original non-passerine land bird diversity in the sample, are globally extinct. These include two megapodes (Megapodius alimentum and a larger unnamed megapode), three pigeons (a large Caloenas sp. indet., Didunculus placopedetes and Ducula shutleri sp. nov.), two rails (Hypotaenidia vavauensis sp. nov. and an unnamed one) and the parrot Eclectus infectus. The rail H. vavauensis was restricted to Vava'u and was flightless, with reduced wings, and larger than Hypotaenidia
woodfordi of the Solomons, the largest congener hitherto found in the Pacific. The pigeon Du. shutleri was volant, but was the largest species in its genus and was widespread in the Kingdom. The evolution of Tongan avifaunas is related to varying ages (Pliocene to Pleistocene) of the island groups, where geological youth apparently precluded true giantism in the fauna.

Enjoy,

Fred

Abbreviations and definitions

Abbreviations and definitions

Institutions:

AIM, Auckland Institute and Museum, Auckland, New Zealand; AM, Australian Museum, Sydney, NSW, Australia; AMNH, American Museum of Natural History, New York, NY, USA; ANWC, Australian National Wildlife Collection, Canberra, ACT, Australia; CM, Canterbury Museum, Christchurch, New Zealand; FM, Fiji Museum, Suva, Fiji; MVZ, Museum of Vertebrate Zoology, University of California, Berkeley, CA, USA; NHMUK, Natural History Museum, London, UK [formerly British Museum (Natural History)]; NMNZ, Museum of New Zealand Te Papa Tongarewa, Wellington, New Zealand; NMV, Museum Victoria, Melbourne, Australia; NTM, Northern Territory Museum/Museum and Art Gallery of the Northern Territory, Darwin, NT, Australia; SAM, South Australian Museum, Adelaide, SA, Australia; UWBM, Burke Museum, University of Washington, Seattle,
WA, USA.

Other abbreviations

For skeletal elements:

ant, anterior; cmc, carpometacarpi; cor, coracoids; cran, cranial part of; fem, femora; fib, fibulae; frag/s, fragment(s); hum, humeri; juv, juvenile; mand, mandibles; phal, phalanges; pmx, premaxillae; quad, quadrates; rad, radii; scap, scapulae; stern, sternum; tmt, tarsometatarsi; tib, tibiotarsi. When listing material, bones are identified as left (L) or right (R) elements; L or R prefixed by ‘p’, ‘s’ or ‘d’ indicates ‘proximal’, ‘shaft’ or ‘distal’ part of the element, e.g. pR fem means the proximal part of a right femur; MNI, minimum number of individuals; NISP, number of individual specimens

SYSTEMATIC PALAEONTOLOGY: Galliformes

SYSTEMATIC PALAEONTOLOGY

Galliformes Temminck, 1820
Megapodiidae Lesson, 1831
Megapodius Gaimard, 1823
Megapodius alimentum Steadman, 1989
Material: Hopoate:NISP = 12, MNI = 2. Nukuleka: NISP = 1, MNI = 1. Ofu: NISP = 35, MNI = 4. Falevai: NISP = 7, MNI = 2. Vuna: NISP = 21, MNI = 4.

Megapodius magn. M. alimentum Steadman, 1989
Material: Hopoate: NISP = 2, MNI = 1. Falevai: NISP = 1, MNI = 1. Ofu: NISP = 5, MNI = 1. Otea: NISP = 1, MNI = 1. Vuna: NISP = 6, MNI = 1.

Megapodius pritchardii G. R. Gray, 1864, Tongan megapode
Material: Hopoate: NISP = 2, MNI = 1. Vuna: PNISP = 1, MNI = 1.

Megapodius magn. M. pritchardii G. R. Gray, 1864, Tongan megapode
Material: Ofu: NISP = 1, MNI = 1. Vuna: NISP = 1, MNI = 1.

Megapodius indet sp.
Material: Hopoate: NISP = 4, MNI = 1. Falevai: NISP = 2. Ofu: f1.05.12.07, ungual phal; NISP = 1. Vuna: NISP = 8, MNI = 2.

Megapode genus & species indet. large sp.
Material: Ofu: Of1.05.22.07, dL hum (SW = 7.85 mm, DW = 18.8 mm); NISP = 1, MNI = 1. Vuna: Pa1.04.18.10a, four frags L cor; NISP = 1, MNI = 1.

Phasianidae Horsfield, 1821
Genus Gallus Brisson, 1760
Gallus gallus (Linnaeus, 1758), red junglefowl
Material: Hopoate: NISP = 36, MNI = 3. Nukuleka: NISP = 12, MNI = 2. Falevai: NISP = 14, MNI = 2. Ofu: NISP = 14, MNI = 2. Otea: NISP = 5, MNI = 2. Vuna: NISP = 13, MNI = 5.

Cf. Gallus gallus (Linnaeus, 1758), red junglefowl
Material: Otea: Ka1.05.04.13, pt pR tmt. NISP = 1, this specimen does not add to the MNI for Gallus.

Galliform family indet
Material: Hopoate: NISP = 3.

SYSTEMATIC PALAEONTOLOGY: Columbiformes part I

Columbiformes Latham, 1790
Columbidae Illiger, 1811
Alopecoenas Sharpe, 1899
Alopecoenas stairi (G.R. Gray, 1856), friendly ground dove
Material: Hopoate: NISP = 8, MNI = 3. Nukuleka: NISP = 1, MNI = 1. Ofu: NISP = 3, MNI = 2. Vuna: NISP = 6, MNI = 2.

Columbid magn. A. stairi (G.R. Gray, 1856)
Material: Hopoate: NISP = 2, MNI = 1. Nukuleka: NISP = 3, MNI = 1.

Caloenas G.R. Gray, 1840
Caloenas sp. indet.
Material: Ofu: Of1.05.12.06, dL sR tmt; Of1.05.12.06, pL tib; Of1.05.12.07, pL rad; Of1.05.12.07, two sternal ends L cor; Of1.05.12.08, d+sL tmt; Of1.05.12.08, ant stern; Of1.05.12.08, p+sR tib; Of1.05.12.09, dL tmt; Of1.05.12.09, pL cmc; Of1.05.12.10, dR hum; Of1.05.12.10, dR ulna, pR rad; Of1.05.20.06, L ulna; Of1.05.20.07, s+dR tmt; Of1.05.20.07, pRsL cmc; Of1.05.20.09, dR ulna; Of1.05.20.10, cran pt L cor; Of1.05.21.08, pLdL rad; Of1.05.22.04, cran pt L cor; Of1.05.22.05, dR ulna; Of1.05.22.05, pL tmt; Of1.05.22.06, dL rad; Of1.05.22.07, dLsR tib; Of1.05.22.07, dL ulna; Of1.05.22.07, one sternal pt L cor; Of1.05.22.08, cran and sternal pts L cor; Of1.05.22.08, sLsR tmt; Of1.05.22.08, sR tib; Of1.05.22.08, dRsL ulna; Of1.05.22.09, pRpL tmt; Of1.05.22.09, cran and sternal pts R cor; Of1.05.22.09, R scap; Of1.05.22.09, pR rad; f1.05.22.09, pR fem; Of1.05.22.10, dL hum; NISP = 44, MNI = 4. Vuna: Pa1.04.21.09, dL cmc; NISP = 1, MNI = 1.

Didunculus Peale, 1849
Didunculus placopedetes Steadman, 2006, Tongan tooth-billed pigeon
Material: Hopoate: Hop.14.11D.12, sL tmt; NISP = 1, MNI = 1. Ofu: Of1.05.21.06, pR fem; Of1.05.22.09, Ofu, d+p L tib; NISP = 2, MNI = 1. Vuna: Pa1.04.24.07, sR tmt; NISP = 1, MNI = 1.

COLUMBIFORMES: part II

Ducula Hodgson, 1836
Ducula pacifica (J. F. Gmelin, 1789), Pacific imperial pigeon
Material: Hopoate: NISP = 22, MNI = 6. Nukuleka: NISP = 2, MNI = 1. Falevai: NISP = 2, MNI = 1. Ofu: NISP = 8, MNI = 2. Otea: NISP = 1, MNI = 1. Vuna: NISP = 20, MNI = 4.

Columbid magn. Ducula pacifica (J. F. Gmelin, 1789), Pacific imperial pigeon
Material: Hopoate: NISP = 3, MNI = 2. Nukuleka: NISP = 11, MNI = 4. Ofu: NISP = 4, MNI = 1. Otea: NISP = 2, MNI = 1. Vuna: NISP = 31, MNI = 6.

Ducula shutleri Worthy & Burley sp. nov., Shutler's fruit pigeon
http://zoobank.org/urn:lsid:zoobank.org:act: C7BC1177-FC61-4BA8-8BBC-C770C6884A20
Holotype: NMNZ S.48354, field number Of1.05.12.08, left tarsometatarsus, collected by David Burley et al. on 6 June 2005.

Paratypes: NMNZ S.48355, field number Of1.05.21.09, distal right tarsometatarsus preserved distal to metatarsal facet; NMNZ S.48356, field number Of1.05.22.04, left tarsometatarsus minus trochlea; NMNZ S.48357, field number Of1.05.22.09, distal left tarsometatarsus

Etymology: Richard Shutler Jr was an early pioneer in Oceanic archaeology, and in 1952 excavated, with Edward Winslow Gifford, at the eponymous Lapita site (Site 13) in New Caledonia (Sand & Kirch, 2002). He
was instrumental in introducing David Burley to Pacific archaeology in the early 1990s and was a part of Burley's Tongan field projects in the 1990s and early 2000s.

Type locality: Layer 8, 70–80 cm depth, Unit 12, Ofu site, Ofu Island, Vava'u Group, Tonga.

Paratypes: See Table 7 and NMNZ S.48355 (Of1.05.21.09), dR tmt, depth of trochlea metatarsi III, 5.0; NMNZ S.48356 (Of1.05.22.04), L tmt, preserved length, 41.2; NMNZ S.48357 (Of1.05.22.09), 1dL tmt,
preserved length, 36.3; depth of trochlea metatarsi III, 5.1.

Referred material: Ducula shutleri Hopoate: Hop.14.TP2.06, sL tmt; NISP = 1, MNI = 1. Falevai: Ka2.05.03.15a, L scap; Ka2.05.XX.00FE0a, sR tmt; NISP = 2, MNI = 1. Ofu: Of1.05.11.05, R scap; Of1.05.12.06, L cor, sR tmt; Of1.05.12.07, L scap, dL cmc, sL tmt; Of1.05.12.07, pR fem; Of1.05.12.08, R scap, cran and sternal pts L cor, 2sR cor, dL tib; Of1.05.12.09, dR tib, dR tmt; Of1.05.12.10, ant stern, R scap, cran and sternal pts 1R cor, sternal pts LandR cor, pL fem; Of1.05.20.06, pL fem; Of1.05.20.06, L scap; Of1.05.20.07, R tib (missing proximal end and distal condyles); Of1.05.20.07, sternal pt L, cran pt L cor; Of1.05.20.07, L fem; Of1.05.20.08, dL tib, sR tmt, pR ulna, R MII.1; Of1.05.20.09, 3pL1dR fem; Of1.05.20.09, 2L scap; Of1.05.20.10, cran pt R cor, L scap, dL tib; Of1.05.21.06, R(-d) fem; Of1.05.21.08, sL tmt; Of1.05.21.08, pR dL rad; Of1.05.21.09, R scap; Of1.05.21.10, s+dR tmt, sR tmt; Of1.05.21.10,
s+pL sL ulna; Of1.05.21.10, R scap; Of1.05.21.10, dR ulna; Of1.05.21.10, pR fem; Of1.05.21.11, cran pt R cor; Of1.05.22.04, sR cor; Of1.05.22.05, pLdL cmc; Of1.05.22.07, pR rad, dL cmc, L scap, 1sR cor, sR tib, sL tmt; Of1.05.22.08, pLdL rad, L scap; Of1.05.22.09, 2pL3dL tmt, 1R one cran pt R one sternal pt R cor; Of1.05.22.09, 2pL rad, R scap; Of1.05.22.10, two cran
pt L 1R cor, pt pL dR tmt; Of1.05.22.10, 3pL1dL fem, 2sR tib, 1dL cmc; including type material NISP = 93, MNI = 11. Vuna: Pa1.04.12.08, pL fem; Pa1.04.12.10, R scap; Pa1.04.22.04, pL fem medullary bone; Pa1.04.23.09, dR tmt; Pa1.04.24.09, pR fem; NISP = 5, MNI = 2.

Tentatively referred material: Identified as ‘Columbid lge, cf. Ducula’. Ofu: Of1.05.12.05, pLdR rad, dLdR ulna, two ulnar shaft frags; Of1.05.12.06, sL ulna, 2sR tib; Of1.05.12.08, d+sR rad, pR rad; d+sL sL sR
d+sR cmc; Of1.05.12.09, 3dL cmc, sL ulna, sL tib; Of1.05.12.10a, sR hum, sL ulna; Of1.05.14.08, sL fem; Of1.05.15.05, dR rad; Of1.05.20.06, sR fem; Of1.05.22.05, L scap, dL rad; Of1.05.22.06, 3pL two pt shaft ulnae, two pts 1L cor, pt R cmc, 2R scap, 1dR rad, 2sR1sL tib, sLsR fem, pt sR tib; Of1.05.22.09, 1dL rad, sLsR tib, pmx; Of1.05.22.10, 1dL2dR cmc; NISP = 51, does not alter Ofu MNI. Vuna: Pa1.04.12.11, sR tmt; Pa1.04.17.08b, sR cor; Pa1.04.24.09, sL tmt; NISP = 3, MNI for Vuna unchanged by these.

Columbid sp. indet. large pigeon
Material: Hopoate: NISP = 2, MNI = 1. Nukuleka: NISP = 6, MNI = 3. Falevai: NISP = 2, MNI = 1. Ofu: NISP = 2, MNI = 1. Otea: NISP = 1, MNI = 1. Vuna: NISP = 7, MNI = 1.

Phaethontiformes-Gruiformes

Phaethontiformes Sharpe, 1891
Phaethontidae Brandt, 1840
Phaethon Linnaeus, 1758
Phaethon lepturus Daudin, 1802, white-tailed tropicbird
Material: Ofu: NISP = 1, MNI = 1.

Phaethon rubricauda Boddaert, 1783, redtailed tropicbird
Material: Vuna: NISP = 1, MNI = 1.

Cuculiformes Wagler, 1830
Cuculidae Leach, 1819
Cacomantis S. Müller, 1843
Cacomantis flabelliformis (Latham, 1801), fan-tailed cuckoo
Material: Hopoate: Hop.14.TP1.10, dR hum; NISP = 1, MNI = 1.

Gruiformes Bonaparte, 1854
Rallidae Rafinesque, 1815
Hypotaenidia Reichenbach, 1853
Hypotaenidia philippensis (Linnaeus, 1766), buff-banded rail
Material: Nukuleka: NISP = 1, MNI = 1. Falevai: NISP = 1, MNI = 1. Ofu: NISP = 4, MNI = 1. Otea: NISP = 3, MNI = 1. Vuna: NISP = 1, MNI = 1.

Hypotaenidia vavauensis Worthy & Burley sp. nov., Vava'u rail
http://zoobank.org/urn:lsid:zoobank.org:act:
D83755C3-07F8-4947-9B45-CED20275BD80
Holotype: NMNZ S.48358, field number Pa1.04.23.05, right tarsometatarsus (Fig. 5A), collected by David Burley et al. on 7 June 2004.

Paratypes: NMNZ S.48359, field number Pa1.04.11.10, R tarsometatarsus lacking trochleae metatarsorum III and TIV; NMNZ S.48360, field number Pa1.04.24.06, pL tarsometatarsus (Fig. 5B); NMNZ S.48361, field number Pa1.04.17.7, near-complete reassembled R tibiotarsus, lacking cnemial crests (Fig. 5C).

Etymology: Named for Vava'u, the island group in Tonga where this species was discovered, and to which it is so far restricted. The the Latin locative suffix–ensis was added to the name.

Type locality: Layer 5, 40–50 cm depth, Unit 23, site is Vuna on Pangaimotu Island, Vava'u Group, Tonga.

Referred material: Falevai: Ka2.05.06.15, dL tmt; Ka1.05.12.15, sR cor; NISP = 2, MNI = 1. Otea: Ka1.05.01.21, dL fem; NISP = 1, MNI = 1. Vuna: Pa1.04.00.00Aa, p+d pts L tib; Pa1.04.00.12, p+sL fem; Pa1.04.02.06a, pL tib; Pa1.04.03.09, pR tib; Pa1.04.06.09, dL tib; Pa1.04.09.07, dL tib; Pa1.04.10.07, pL tib; Pa1.04.10.12, L ramus mand; Pa1.04.11.11 + Pa1.04.11.08b, d+sL tib, two pts L fem; Pa1.04.13.09, sL tib; Pa1.04.14.07, dL hum; Pa1.04.14.09, dL hum, L fib; Pa1.04.14.10b, dR fem, R cmc; Pa1.04.14.11, dL cmc; Pa1.04.15.04, pL fem; Pa1.04.15.07b, dR fem; Pa1.04.16.08, pR tib; Pa1.04.16.10, L cor; Pa1.04.17.07, sR tmt; Pa1.04.17.08b, dR ulna; Pa1.04.17.10, dL ulna; Pa1.04.18.08a, dL tib, cran pt L cor; Pa1.04.18.09, two pts R tmt; Pa1.04.18.11, dL tib; Pa1.04.19.11a, pR fem; Pa1.04.20.10, dL tib, dR fem, dR hum; Pa1.04.21.07, synsacrum; Pa1.04.21.08, ant sternum; Pa1.04.21.09, sR tmt, sR hum; Pa1.04.21.09, dR ulna; Pa1.04.22.04, sL tib, pt sR tmt; Pa1.04.22.07, sR fem; Pa1.04.22.08, R ulna; Pa1.04.22.11, pt pelvis; Pa1.04.23.06, dL tib; Pa1.04.24.10, cran pt L cor; Pa1.04.25.06, dR hum; Pa1.04.25.08, dR fem, dR tib, sR tmt; Pa1.04.25.09b, dR tib; Pa1.04.25.11, s+dL hum; Pa1.04.26.07, L fem, pR fem; Pa1.04.26.10, L(-p) hum, dL tmt; Including type material, NISP = 58, MNI = 9.

Tentatively referred material: Nukuleka: To2.07.38.05, sR hum; To2.07.26.11, L fem; NISP = 2,
MNI = 1.

Porphyrio Brisson, 1760
Porphyrio melanotus Temminck, 1820, southwest Pacific swamphen
Material: Hopoate: NISP = 1, MNI = 1. Nukuleka: NISP = 3, MNI = 1. Falevai: NISP = 1, MNI = 1. Ofu:
NISP = 1, MNI = 1. Vuna: NISP = 1, MNI = 1.




Bones of Hypotaenidia vavauensis from Vuna Island, Vava'u Group, Tonga, showing relative proportions of leg and wing elements. A, holotype right tarsometatarsus NMNZ S.48358 (Pa1.04.23.05) in dorsal (A1), plantar (A2) and medial (A3) views. B, paratype proximal left tarsometatarsus NMNZ S.48360 (Pa1.04.24.06) in proximal view. C, paratype left tibiotarsus NMNZ S.48361 (Pa1.04.17.7) in anterior view. D, left humerus Pa1.04.2b.10 in cranial (D1) and caudal (D2) views. E, left femur Pa1.04.26.07 in caudal (E1), cranial (E2) and medial (E3) views. F, right ulna Pa1.04.22.08 in ventral view. G, left coracoid missing sternal articulation, Pa1.04.16.10, in dorsal view. H, distal right femur Pa1.04.15.07b in caudal view. Scale bars: 10 mm. Abbreviations: cmfh, crista medialis flexoris hallucis longus; dRET, distal attachment for retinaculum extensorium tibiotarsi; fbl, sulcus for musculus fibularis longus; fdl, sulcus for tendon of musculus flexor digitorum longus; fhl, sulcus for tendon of musculus flexor hallucis longus; fp2, sulcus for the tendons of musculus flexor perforatus digiti 2 and musculus flexor perforans et perforatus digiti 2; ire, impressiones retinaculi extensorii; pRET, proximal attachment for retinaculum extensorium tibiotarsi; tmgl, tuberculum musculus gastrocnemialis lateralis.

Procellariiformes-Passeriformes

Procellariiformes Fürbringer, 1888
Procellariidae Leach, 1820
Ardenna Reichenbach, 1853
Ardenna pacifica (Gmelin, 1789), wedge-tailed shearwater
Material: Nukuleka: NISP = 3, MNI = 1. Falevai: NISP = 3, MNI = 1. Ofu: NISP = 20, MNI = 2. Vuna:
NISP = 8, MNI = 1.

Procellariid magn. Ardenna pacifica (Gmelin, 1789)
Material: Otea: Ka1.05.12.18, pL cmc, NISP = 1, MNI = 1.

Procellariid sp. lge
?Pseudobulweria rostrata (Peale, 1849), Tahiti petrel
Material: Otea: Ka1.05.01.21, dL hum, NISP = 1, MNI = 1.

Pelecaniformes Sharpe, 1891
Ardeidae Leach, 1820
Ardea Linnaeus, 1758
Ardea modesta J. E. Gray, 1831, white heron/eastern great egret
Material: Falevai: Ka2.05.13.14, pL fem (lacking proximal three-quarters of trochanter), NISP = 1,
MNI = 1. Vuna: Pa1.04.11.08a, dR tib, NISP = 1, MNI = 1.

Charadriiformes Huxley, 1867
Laridae Rafinesque, 1815
Thalasseus Boie, 1822
Thalasseus bergii (M. H. Lichtenstein, 1823), crested tern
Material: Falevai: Ka2.05.03.10b, pL ulna, NISP = 1, MNI = 1. Vuna: Pa1.04.26.07, pt R cor, NISP = 1, MNI = 1.

Charadriidae Leach, 1820
Pluvialis Brisson, 1760
Pluvialis fulva (J. F. Gmelin, 1789), Pacific golden plover
Material: Falevai: Ka2.05.03.12a, juv R tmt, NISP = 1, MNI = 1. Ofu: Of1.05.03.07, dR hum,
NISP = 1, MNI = 1.

Psittaciformes Wagler, 1830
Psittaculidae Vigors, 1825
Eclectus Wagler, 1832
Eclectus infectus Steadman, 2006
Material: Hopoate: NISP = 1, MNI = 1.; Nukuleka: NISP = 1, MNI = 1. Ofu: NISP = 6, MNI = 2. Vuna:
NISP = 2, MNI = 1.

?Eclectus infectus Steadman, 2006
Material: Ofu: NISP = 6, MNI = 1 (does not change MNI for certainly identified material of this taxon).

Vini Lesson, 1833
Vini sp. indet
Material: Vuna: NISP = 3, MNI = 1.

Passeriformes
Undetermined passerines
Material: Hopoate: NISP = 4, MNI = 1. Nukuleka: NISP = 2, MNI = 1. Ofu: NISP = 4, MNI = 1. Vuna:
NISP = 9, MNI = 2.
Remarks: In the absence of appropriate comparative material, no attempt was made to identify these few specimens.

Fred