The region from New Guinea through Oceania sustains the world’s most diverse set of columbids. We describe osteological characters of the hindlimb (femur, tibiotarsus, tarsometatarsus) that divide the Papuan-Oceanic pigeons and doves into three groups based on functional morphology: “arboreal” (Hemiphaga, Ducula, Ptilinopus, Drepanoptila, Gymnophaps), “intermediate” (Columba, Macropygia, Reinwardtoena), and “terrestrial” (Gallicolumba [includes Alopecoenas], Trugon, Microgoura, Goura, Chalcophaps, Geopelia, Henicophaps, Caloenas, Didunculus, Otidiphaps). The arboreal and terrestrial groups are each distinctive osteologically, especially in the tibiotarsus and tarsometatarsus, which are short relative to the femur in the arboreal group, and long relative to the femur in the terrestrial group. The intermediate pigeons are more similar to arboreal than to terrestrial pigeons, but nonetheless fit in neither group. To estimate the phylogenetic relationships among or within these three groups is somewhat tentative using hindlimb osteology alone, although all five genera of arboreal pigeons have independent molecular evidence of relatedness, as do most of the genera of terrestrial pigeons.
Using the hindlimb and other osteological data as a framework, we describe a new extinct genus and species of pigeon, Tongoenas burleyi, from Holocene archaeological and Pleistocene paleontological sites on six islands (Foa, Lifuka, `Uiha, Ha`afeva, Tongatapu, and `Eua) in the Kingdom of Tonga. Tongoenas was a large-sized member of the “arboreal” pigeon group, with osteological characters that relate it to Ducula, Gymnophaps, and Hemiphaga (generally canopy frugivores) rather than with the “terrestrial” pigeons (more ground-dwelling and granivorous) such as Gallicolumba, Trugon, Microgoura, Goura, etc. (others listed above). Among volant columbids, living or extinct, only the species of Goura (from New Guinea) are larger than Tongoenas. From most of the same prehistoric sites, we also report new material of the nearly as large, extinct pigeon Ducula shutleri Worthy & Burley, recently described from islands in the Vava`u Group of Tonga. Thus, D. shutleri also was widespread in Tonga before human impact. The prehistoric anthropogenic loss in Tonga of Tongoenas burleyi, Ducula shutleri, and other columbids undoubtedly had a negative impact on the dispersal regimes of Tongan forest trees. At first human contact about 2850 years ago, at least nine species of columbids in six genera inhabited the Tongan islands, where only four species in three genera exist today.
Balouet, J.-C. & Olson, S.L. (1989) Fossil birds from late Quaternary deposits in New Caledonia. Smithsonian Contributions to Zoology, 469, 1–38.
Baptista, L.F., Trail, P.W. & Horblit, H.M. (1997) Family Columbidae (Pigeons and Doves). In: del Hoyo, J., Elliott, A. & Sargatal, J. (Eds.), Handbook to the Birds of the World. Vol. 4. Sandgrouse-Cuckoos. Lynx Edicions, Barcelona, pp. 60–243.
Baumel, J.J., King, A.S., Lukas, A.M., Breazile, J.E. & Evans, H.E. (Eds.) (1993) Nomina anatomica avium. Academic Press, London, 779 pp.
Beehler, B.M., Pratt, T.K. & Zimmerman, D.A. (1986) Birds of New Guinea. Princeton University Press, Princeton, 528 pp.
Beichle, U. (1987) Lebensraum, bestand und nahrungsaufnahme der zahntaube, Didunculus strigirostris. Journal für Ornithologie, 128, 75–89.
Brown, R.W. (1956) Composition of Scientific Words. Smithsonian Institution, Washington, D.C., 882 pp.
Burley, D.V. (1994) Settlement pattern and Tongan prehistory: reconsiderations from Ha`apai. Journal of the Polynesian Society, 103 (4), 379–411.
Burley, D.V. (1999) Lapita settlement to the East: new data and changing perspectives from Ha`apai (Tonga) prehistory. In: Galipaud, J.-C. & Lilley, I. (Eds.), The Pacific from 5000 to 2000 BP: Colonisation and transformations. Paris: Institut de Recherche pour le Développement, pp. 189–200.
Burley, D.V. (2016) Reconsideration of sea level and landscape from first Lapita settlement at Nukuleka, Kingdom of Tonga. Archaeology in Oceania, 51 (2), 84–90.
Burley, D.V., Barton, A., Dickinson, W.R., Connaughton, S.P. & Taché, K. (2010) Nukuleka as a founder colony for West Polynesian settlement: new insights from recent excavations. Journal of Pacific Archaeology, 1 (2), 128–144.
Burley, D.V., Nelson, E. & Shutler, R. (1995) Rethinking Tongan Lapita chronology in Ha`apai. Archaeology in Oceania, 30 (3), 132–134.
Cain, A.J. (1954) Animal Species and their Evolution. Hutchinson University Library, London, 210 pp.
Claessens, L.P.A.M., Meijer, H.J.M., Hume, J.P. & Rijsdijk, K.F. (Eds.) (2015) Anatomy of the Dodo (Raphus cucullatus L., 1758): an osteological study of the Thirioux specimens. Society of Vertebrate Paleontology Memoir, 15, 1–187.
Coates, B.J. (1985) The Birds of Papua New Guinea. Vol. 1, Non-passerines. Dove Publications, Brisbane, 576 pp.
Coates, B.J. & Bishop, K.D. (1997) A Guide to the Birds of Wallacea. Dove Publications, Alderley, 536 pp.
Diamond, J.M. (1972) Avifauna of the Eastern Highlands of New Guinea. Nuttall Ornithological Club, Cambridge, Massachusetts, 438 pp.
Diamond, J.M. (1974) Colonization of exploded volcanic islands by birds: the supertramp strategy. Science, 184 (4138), 803–806.
Dickinson, W.R., Burley, D.V. & Shutler, R. (1999) Holocene paleoshoreline record in Tonga: geomorphic features and archaeological implications. Journal of Coastal Research, 15 (3), 682–700.
Dunning, J.B. Jr. (2008) CRC Handbook of Avian Body Masses. 2nd Edition. CRC Press, Boca Raton, 666 pp.
Franklin, J. & Steadman, D.W. (2008) Prehistoric species richness of birds on oceanic islands. Oikos, 117 (12), 1885–1891.
Gibb, G.C. & Penney, D. (2010) Two aspects along the continuum of pigeon evolution: A South-Pacific radiation and the relationship of pigeons within Neoaves. Molecular Phylogenetics and Evolution, 56 (2), 698–706.
Gifford, E.W. (1941) The taxonomy and habits of pigeons. Auk, 58 (2), 242–245.
Goodwin, D. (1959) Taxonomy of the genus Columba. Bulletin of the British Museum (Natural History) Zoology, 6, 1–23.
Goodwin, D. (1960) Taxonomy of the genus Ducula. Ibis, 102 (4), 526–535.
Goodwin, D. (1983) Pigeons and Doves of the World. Cornell University Press, Ithaca, New York, 363 pp.
Howard, H. (1929) The avifauna of the Emeryville Shellmound. University of California Publications in Zoology, 32, 301–394.
Johnson, K.P. & Clayton, D.H. (2000) Nuclear and mitochondrial genes contain similar phylogenetic signal for pigeons and doves (Aves: Columbiformes). Molecular Phylogenetics and Evolution, 14 (1), 141–151.
Koopman, K.F. & Steadman, D.W. (1995) Extinction and biogeography of bats on `Eua, Kingdom of Tonga. American Museum Novitates, 3125, 1–13.
Mayr, E. & Diamond, J. (2001) The birds of northern Melanesia: speciation, ecology, and biogeography. Oxford University Press, New York, 548 pp.
McNab, B.K. (2000) The influence of body size, climate, and distribution on the energetics of pigeons. Comparative Biochemistry and Physiology, 127, 309–329.
McNab, B.K. (2002) Minimizing energy expenditure facilitates vertebrate persistence on oceanic islands. Ecology Letters, 5 (5), 693–704.
Moyle, R.G., Jones, R.M. & Andersen, M.J. (2013) A reconsideration of Gallicolumba (Aves: Columbidae) relationships using fresh source material reveals pseudogenes, chimeras, and a novel phylogenetic hypothesis. Molecular Phylogenetics and Evolution, 66 (3), 1060–1066.
Pereira, S.L., Johnson, K.P., Clayton, D.H. & Baker, A.J. (2007) Mitochondrial and nuclear DNA sequences support a Cretaceous origin of Columbiformes and a dispersal-driven radiation in the Paleogene. Systematic Biology, 56 (4), 656–672.
Peters, J.L. (1937) Checklist of the Birds of the World. Vol. 3. Harvard University Press, Cambridge, Massachusetts, 311 pp.
Pratt, H.D., Bruner, P.L. & Berrett, D.G. (1987) A field guide to the birds of Hawaii and the tropical Pacific. Princeton University Press, Princeton, New Jersey, 528 pp.
Pregill, G.K. (1993) Fossil lizards from the Late Quaternary of `Eua, Tonga. Pacific Science, 47 (2), 101–114.
Pregill, G.K. & Steadman, D.W. (2004) South Pacific iguanas: human impacts and a new species. Journal of Herpetology, 38 (1), 15–21.
Salvadori, T. (1893) Columbae, or pigeons. In: Sharpe, R.B. (Ed.), Catalogue of the birds in the British Museum. Vol. 21. Order of the Trustees, London, 676 pp.
Shapiro, B., Sibthorpe, D., Rambaut, A., Austin, J., Wragg, G.M., Bininda-Emonds, O.R.P., Lee, P.L.M. & Cooper, A. (2002) Flight of the dodo. Science, 295 (5560), 1683.
Soares, A.E.R., Novak, B.J., Haile, J., Heupink, T.H., Fjeldsa, J., Gilbert, M.T.P., Poinar, H., Church, G.M. & Shapiro, B. (2016) Complete mitochondrial genomes of living and extinct pigeons revise the timing of the columbiform radiation. BMC Evolutionary Biology, 16 (230), 1–9.
Steadman, D.W. (1989) New species and records of birds (Aves: Megapodiidae, Columbidae) from an archeological site on Lifuka, Tonga. Proceedings of the Biological Society of Washington, 102 (3), 537–552.
Steadman, D.W. (1992) New species of Gallicolumba and Macropygia (Aves: Columbidae) from archaeological sites in Polynesia. Los Angeles County Museum of Natural History, Science Series, 36, 329–348.
Steadman, D.W. (1993) Biogeography of Tongan birds before and after human impact. Proceedings of the National Academy of Science USA, 90 (3), 818–822.
Steadman, D.W. (1995) Prehistoric extinctions of Pacific island birds: biodiversity meets zooarchaeology. Science, 267 (5201), 1123–1131.
Steadman, D.W. (1997) The historic biogeography and community ecology of Polynesian pigeons and doves. Journal of Biogeography, 24 (6), 737–753.
Steadman, D.W. (1998) Status of land birds on selected islands in the Ha’apai Group, Kingdom of Tonga. Pacific Science, 52 (1), 14–34.
Steadman, D.W. (1999) The prehistory of vertebrates, especially birds, on Tinian, Aguiguan, and Rota, Northern Mariana Islands. Micronesica, 31 (2), 59–85.
Steadman, D.W. (2006a) Extinction and biogeography of tropical Pacific birds. University of Chicago Press, Chicago, Illinois, xiv + 594 pp.
Steadman, D.W. (2006b) An extinct species of tooth-billed pigeon (Didunculus) from the Kingdom of Tonga, and the concept of endemism in insular landbirds. Journal of Zoology, 268 (3), 233–241.
Steadman, D.W. (2008) Doves (Columbidae) and cuckoos (Cuculidae) from the Early Miocene of Florida. Bulletin of the Florida Museum of Natural History, 48 (1), 1–16.
Steadman, D.W. & Freifeld, H.B. (1998) Distribution and relative abundance of landbirds in the Vava`u Group, Kingdom of Tonga. Condor, 100 (4), 609–628.
Steadman, D.W. & Freifeld, H.B. (1999) The food habits of Polynesian pigeons and doves: a systematic and biogeographic review. Ecotropica, 5, 13–33.
Steadman, D.W., Plourde, A. & Burley, D.V. (2002a) Prehistoric butchery and consumption of birds in the Kingdom of Tonga, South Pacific. Journal of Archaeological Science, 29 (6), 571–584.
Steadman, D.W., Pregill, G.K. & Burley, D.V. (2002b) Rapid prehistoric extinction of iguanas and birds in Polynesia. Proceedings of the National Academy of Sciences, 99 (6), 3673–3677.
Thibault, J.-C. & Cibois, A. (2017) Birds of Eastern Polynesia: A Biogeographic Atlas. Lynx Edicions, Barcelona, 440 pp.
Wallace, A.R. (1865) On the pigeons of the Malay Archipelago. Ibis, New Series, 1, 365–400.
Watling, D. (2001) A Guide to the Birds of Fiji & Western Polynesia. Environmental Consultants Fiji Ltd., Suva, 272 pp.
Whistler, W.A. (2002) The Samoan Rainforest: A Guide to the Vegetation of the Samoan Archipelago. Isle Botanica, Honolulu, 168 pp.
Worthy, T.H. (2000) The fossil megapodes (Aves: Megapodiidae) of Fiji with descriptions of a new genus and two new species. Journal of the Royal Society of New Zealand, 30 (4), 337–364.
Worthy, T.H. (2001) A giant flightless pigeon gen. et sp. nov. and a new species of Ducula (Aves: Columbidae), from Quaternary deposits in Fiji. Journal of the Royal Society of New Zealand, 31 (4), 763–794.
Worthy, T.H. & Burley, D.V. (2020) Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society, 189 (3), 998–1045.
Worthy, T.H. & Wragg, G.M. (2008) A new genus and species of pigeon (Aves: Columbidae) from Henderson Island, Pitcairn Group. In: Clark, G., Leach, F. & O’Connor, S. (Eds.), Terra Australis, 29, pp. 499–510.
Wragg, G.M. & Worthy, T.H. (2006) A new species of extinct imperial pigeon (Ducula: Columbidae) from Henderson Island, Pitcairn Group. Historical Biology, 18 (2), 127–140.
Wright, N.A. & Steadman, D.W. (2012) Insular avian adaptations on two Neotropical continental islands. Journal of Biogeography, 39 (10), 1891–1899.
Wright, N.A., Steadman, D.W. & Witt, C.C. (2016) Predictable evolution toward flightlessness in volant island birds. Proceedings of the National Academy of Sciences, 113 (17), 4765–4770.