taxonID	type	description	language	source
7E2E97170246C21C3F6457A6FA7FFCB1.taxon	materials_examined	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. Remarks: The bones of this medium-sized megapode were identified to genus by the features described by Worthy (2000) and Worthy et al. (2015) and were compared directly with specimens of Megapodius reinwardt Dumont, 1823. Their size (Table 2) matches those of M. alimentum measured by Worthy (2000), who also provides sizes of bones of other megapodes.	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E97170246C21C3D2557ECFA84FE41.taxon	discussion	Remarks: Megapodes or scrubfowl of the genus Megapodius are widely distributed through Indonesia, Melanesia and western Polynesia and include many taxa that have been treated variously as subspecies or species (Mayr, 1938; White & Bruce, 1986; Jones et al., 1995). Now 13 extant species are recognized (Dickinson & Remsen, 2013). Several extinct species reveal a much greater Holocene diversity: Megapodius molistructor Balouet & Olson, 1989 from New Caledonia, Megapodius amissus Worthy, 2000 from Viti Levu, Fiji and M. alimentum from the Lau Group, Fiji and Tonga (Steadman, 1989 a; Worthy, 2000) fill the gap in extant distribution of the genus between Vanuatu and Tonga. In Tonga, the genus is now represented only by M. pritchardii, which is restricted to Niuafo'ou in northern Tonga (Jones et al., 1995; Dickinson & Remsen, 2013). Formerly, it was once widespread in Tonga, and its range also included American Samoa and Niue (Steadman, 1993, 1994, 1995, 2006 a; Steadman et al., 2000, 2002). To date, no other megapode genera are known from the Tongan region (Steadman, 2006 a). The nearest such to Tonga are present in Fiji, where a giant flightless form, Megavitiornis, existed (Worthy, 2000), and Vanuatu, where a large volant form, Mwalau, also co-existed with Megapodius (Worthy et al., 2015). Species of Megapodius are easily distinguished from those of other megapode genera by numerous features, as listed by Worthy (2000) and Worthy et al. (2015).	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E97170246C21F3F1355E2FD4AFCB0.taxon	materials_examined	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. Remarks: These specimens include those of the size identified as M. alimentum above, but which are not UF 57795 is a tarsometatarsus from Lifuka (Tonga) referred to M. alimentum by D. Steadman, with measurements from Worthy (2000). ‘ Lakeba’ shows data from Worthy (2000) for bones referred to M. alimentum from Lakeba in the Lau Group (Fiji). Abbreviations: CorSW, minimum shaft width of coracoid; FemDW, distal width of femur; FemPW, proximal width of femur; FemSW, width of femur at mid-shaft; HumDW, distal width of humerus; HumSW, width of humerus at mid-shaft; TibDW, distal width of condyles of tibiotarsus; TibSW, width of tibiotarsus at mid-shaft; TmtPW, proximal width of tibiotarsus; TmtSW, width of tarsometatarsus at mid-shaft; UlnaDW, maximum distal width of ulna; UlnaSW, width of ulna at mid-shaft. sufficiently complete to be certain of a specific referral. Addition of these remains to those identified as M. alimentum increases the MNI for Ofu by one, but leaves MNI for other sites unchanged.	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E97170246C21F3F1355E2FD4AFCB0.taxon	materials_examined	Material: Hopoate: NISP = 2, MNI = 1. Vuna: PNISP = 1, MNI = 1.	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E97170246C21F3F1355E2FD4AFCB0.taxon	materials_examined	Material: Ofu: NISP = 1, MNI = 1. Vuna: NISP = 1, MNI = 1. Remarks: These specimens are of the small size typical of this species (the smallest of the genus; see Worthy, 2000) and are likely to belong to M. pritchardii.	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E97170245C21F3D6955D6FA67FB87.taxon	materials_examined	Material: Hopoate: NISP = 4, MNI = 1. Falevai: NISP = 2. Ofu: Of 1.05.12.07, ungual phal; NISP = 1. Vuna: NISP = 8, MNI = 2. Remarks: These specimens are either juvenile, undiagnostic elements or too fragmentary for identification to species level. MEGAPODE GENUS & SPECIES INDET. LARGE SP. Material: Ofu: Of 1.05.22.07, dL hum (SW = 7.85 mm, D W = 1 8.8 m m); N I S P = 1, M N I = 1. Vu n a: Pa 1.04.18.10 a, four frags L cor; NISP = 1, MNI = 1. Remarks: These specimens are bigger than specimens for M. alimentum and about the same size as specimens of Macrocephalon maleo Müller, 1846 or that expected of M. molistructor, for which no humerus is known. On the distal humerus, the fossa brachialis is deep, and the origin of m. extensor carpi radialis has two scars aligned level with the most proximal point of condylus dorsalis. The pars dorsalis is round, elevated off the surface and protuberant craniodorsally on the dorsal margin, but extends less dorsally than the epicondylus dorsalis, from which it is separated by a shallow notch, as seen in cranial aspect. The pars ventralis is elongated and lies ventrad and extends slightly distal to the former. In Macrocephalon maleo AMNH 12013, the pars dorsalis origin of m. extensor carpi radialis is prominent dorsally of the shaft, but lies distal to the pars ventralis and is more prominent than the epicondylus dorsalis. Also, the shaft is more robust (SW = 9.3 mm, DW = 18.5 mm), together, precluding referral of the Tongan bone to this genus and species. The distal humerus of Mwalau from Vanuatu is not known; therefore, comparison to the Tongan sample cannot be made, but it was a bigger bird than all extant megapodes (Worthy et al., 2015). In M. reinwardt (ANWC 22869) and M. alimentum specimens above, the pars dorsalis of the origin of m. extensor carpi radialis is also located immediately and entirely distal to the pars ventralis and on the dorsal facies and forms a low processus supracondylaris dorsalis that projects further dorsally than the epicondylus dorsalis. Therefore, Of 1.05.22.07 cannot be referred to Megapodius. Steadman referred numerous bones to M. molistructor but stated that they ‘ were somewhat aberrant among Megapodius and may, with further study, warrant its own genus’ (Steadman, 2006 a: 292). The coracoid fragments represent a species about as large as the humerus, but little more can be said. Steadman (2006 a: 292) noted that M. molistructor was scarce in the Ha'apai Group relative to M. alimentum, which accords with our observations of the sites reported herein. Therefore, the bones listed here probably belong to Steadman's (2006 a) ‘ M. molistructor ’, but assessing their relationship among megapodes is outside the scope of this report.	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E97170245C21E3F05533FFDACFE01.taxon	materials_examined	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. Remarks: This is a common species, especially in later archaeological sites in Oceania. In these assemblages, it was a common component of the Lapita levels, confirming the early spread of Gallus into Oceania. CF. GALLUS GALLUS (LINNAEUS, 1758), RED JUNGLEFOWL Material: Otea: Ka 1.05.04.13, pt pR tmt. NISP = 1, this specimen does not add to the MNI for Gallus. Measurements: See Appendix 1. Sample sizes were too small to assess inter-island size variation effectively, but a perusal of the data showed no trends; therefore, all data were pooled. These chickens are about the same size as M. alimentum, the other common galliform in the sites, but are more variable, reflecting marked sexual dimorphism in Gallus. GALLIFORM FAMILY INDET	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E97170244C21E3CCB52D3FE98F8A1.taxon	discussion	Remarks: Recent molecular analyses have revealed that the ground dove genus Gallicolumba, sensu Dickinson (2003), was paraphyletic (Jønsson et al., 2011; Moyle et al., 2013); therefore, it has been separated into Gallicolumba for the bleeding-hearted doves and relatives and Alopecoenas for the ground doves, including all those east of Papua New Guinea (Dickinson & Remsen, 2013). Alopecoenas stairi (Gray, 1856) is the only extant species of the genus found in the Fiji – Samoa – Tonga region (Dickinson & Remsen, 2013). The following specimens were compared with bones of this species from Fiji (NMNZ S 38223). Diagnostic features for bones of this genus were given by Worthy & Wragg (2003, 2008 and Worthy et al. (2015).	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E97170244C21E3CCB52D3FE98F8A1.taxon	materials_examined	Material: Hopoate: NISP = 8, MNI = 3. Nukuleka: NISP = 1, MNI = 1. Ofu: NISP = 3, MNI = 2. Vuna: NISP = 6, MNI = 2.	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E97170244C2113FAB5703FD96FEA7.taxon	discussion	Remarks: The genus Caloenas is represented by one extant species, the Nicobar pigeon Caloenas nicobarica (Linnaeus, 1758). This is a large pigeon, with spectacular plumage, that is restricted to insular parts of Southeast Asia through Indonesia and New Guinea, extending to the Solomons (Dickinson & Remsen, 2013). The extinct species Caloenas canacorum Balouet & Olson, 1989 was described from New Caledonia based on three coracoids, part of a sternum, two scapulae and a distal humerus, which reveal a bird some 25 % larger than the Nicobar pigeon (Balouet & Olson, 1989). Steadman (1989, 2006 a) referred specimens from Tonga (archaeological sites on Lifuka, Ha'ano and Ha'afeva in the Ha'apai Group) to Caloenas cf. nicobarica. In addition, there is a single specimen of a third species, Caloenas maculata (Gmelin, 1789), collected historically from an unknown location before 1783. This specimen is a bird 32 cm long, smaller than C. nicobarica (40 cm), and has been confirmed as a member of Caloenas based on molecular data (Heupink et al., 2014). Tongan specimens referred to C. canacorum cannot be C. maculata, because they are of similar size to the New Caledonian bones, and thus are from a much larger taxon. Thespecimenslistedbelowallrepresentalargepigeon distinct from Ducula, and thus differ substantially from the giant form described below in that genus. They are identified as being from a species of Caloenas because of the combination of features observed in the coracoid, tibiotarsus and tarsometatarsus (Fig. 3). Coracoid: with the cotyla scapularis elongated laterosternally into an acute angle, and the tuberculum A (Worthy, 2012) globose, dorsally prominent over the sulcus supracoracoideus and with a relatively narrow separation from the cotyla scapularis. Sternally, the impressio m. sternocoracoidei is pneumatic and widely separated from the medial margin by a dorsoventrally thickened (inflated) section of the corpus extending from the angulus medialis. Tibiotarsus: distal end, craniocaudal depth of condylus medialis greater than distal width; proximal attachment for retinaculum extensorium tibiotarsi (ptRET) close to condylus medialis (separation less than proximodistal length of condyle); distal attachment for retinaculum extensorium tibiotarsi (dtRET) is wider than long; the sulcus extensorius is bounded laterally by a rounded ridge (not sharp crest). Tarsometatarsus: elongate, but fossa parahypotarsalis medialis nearly reaches fossa metatarsi I; foramen vascularia proximalia both small; tuberositas m. tibialis cranialis deep set in sulcus extensorius, not prominent; anterior facies adjacent to sulcus extensorius flat to slightly convex; and trochlea metatarsi IV not deeply notched distoplantarly and ends proximad to trochlea metatarsi II.	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E97170244C21E3CCF54A0FD77FBAE.taxon	discussion	Remarks: The columbid bones in the Vava'u assemblages can be separated into three broad size classes: small taxa (Ptilinopus and Gallicolumba), medium-sized taxa [Ducula pacifica (Gmelin, 1789) and Di. placopedetes] and large taxa (Caloenas sp. and a large species of Ducula). For columbids, the coracoid and tarsometatarsus are the most diagnostic elements (Worthy, 2012; Worthy et al., 2015; Rigal et al., 2018) and, fortunately, are well represented in these samples. Diagnostic characters for columbid bones are presented at length by Rigal et al. (2018), Worthy (2012), Worthy & Wragg (2003, 2008, Wragg & Worthy (2006), Worthy et al. (2015).	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E9717024DC2173F94547BFAD7FB79.taxon	discussion	Remarks: The genus Didunculus is represented by one extant species, the tooth-billed pigeon, Didunculus strigirostris (Jardine, 1845), a large colourful pigeon, notable for its stout, hooked bill, that historically was restricted to the islands of Savai'i, Upolu and Nu'utele in Western Samoa (Steadman, 2006 a, b). Steadman (2006 b) described an extinct species, Di. placopedetes, from Late Holocene fossil deposits on ' Eua and Tongatapu in Tonga. This was also a common species in the Teouma fauna on Efate, Vanuatu (Worthy et al., 2015). Diagnostic features distinguishing this species from the similar-sized Columba vitiensis Quoy & Gaimard, 1830 and Du. pacifica and from the larger C. nicobarica were presented by Worthy et al. (2015). The present Tongan material did not differ from the Vanuatu material. DIDUNCULUS PLACOPEDETES STEADMAN, 2006, TONGAN TOOTH-BILLED PIGEON	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E9717024CC20A3D335729FC62FCD5.taxon	discussion	Remarks: This genus is widely distributed from Asia, through the Philippines, Melanesia and across Polynesia (Dickinson & Remsen, 2013). The widespread Du. pacifica is the only extant member of the genus known from Tonga, although other extant species are known to the west in Fiji, Ducula latrans (Peale, 1849), and to the east in Society Islands and Tuamotu Archipelago, Ducula aurorae (Peale, 1849) (Dickinson & Remsen, 2013). In addition to such medium-sized pigeons, there existed formerly a suite of larger species of Ducula from New Caledonia to the Marquesas, although only the species at these two geographical extremes still survive: Du. goliath (New Caledonia) and Du. galeata (Marquesas). Extinct populations or species of large Ducula, from west to east, include Ducula sp. cf. Du. goliath on Efate in Vanuatu (Worthy et al., 2015), an undescribed Ducula sp. on Viti Levu and Ducula lakeba Worthy, 2001 on the Lau Group, Fiji (Worthy, 2001), Ducula david Balouet & Olson, 1987 on Uvea in Wallis and Futuna (Balouet & Olson, 1987), extirpated populations referred to as Du. galeata on the Cook Islands (Mangaia) and on the Society Islands (Huahine) (Steadman, 1989 a, b, 1995, 1997, 2006 a), an undescribed species from Rapa Island in the Austral Group (Tennyson & Anderson, 2012), Ducula tihonireasini Rigal et al., 2018 on Mangareva (Rigal et al., 2018) and Ducula harrisoni Wragg & Worthy, 2006 on Henderson Island (Wragg & Worthy, 2006). In addition to these populations or taxa, a large undescribed species of Ducula has been reported from several islands in Tonga [Steadman, 1989 a, 1993 (as Du. david), 1995, 1997, 2006 a]. IDENTIFYING TRAITS Species of Ducula are easily distinguished on most limb bones from columbids of a similar size by the following diagnostic features (Worthy, 2001; Wragg & Worthy, 2006; Worthy & Wragg, 2008; Rigal et al., 2018). The elements coracoid, tibiotarsus and tarsometatarsus are the most diagnostic; therefore, key features are listed here. Coracoid: The sulcus m. supracoracoidei is not pneumatic; the tuberculum A (see terminology of Worthy, 2012) is large, abuts the facies articularis humeralis, does not abruptly overhang the sulcus m. supracoracoidei and is widely separated from the insertion for the ligamentum acrocoraco-acromion acrocoracoid (ilaa); the tuberculum for ilaa has little projection over the sulcus m. supracoracoidei; the facies articularis clavicularis extends ventrally from the tuberculum for ilaa as a flattened facies to a large, sternally directed projection for the insertion for the ligamentum acrocoraco-procoracoideum (ilap) on its ventral margin; there is no crest overhanging the sulcus m. supracoracoidei connecting ilaa and ilap; sternally, the impressio m. sternocoracoidei extends close to the medial margin and has a large pneumatic foramen penetrating towards the angulus medialis; the facies articularis sternalis dorsalis is broad, especially medially, and a crest extends cranially from the angulus medialis, enclosing a distinct flattened area dorsally between it and the impressio m. sternocoracoidei and the facies articularis sternalis. Tibiotarsus: The distal condylar width is about the same caudally and cranially; the depth of the condylus medialis is about equal to the distal condylar width; the medial tuberosity for the proximal attachment of the retinaculum extensorium tibiotarsi (ptRET) is prominent and is separated from the condylus medialis by a distance greater than the proximodistal height of that condyle; the distal attachment point for the retinaculum extensorium tibiotarsi (dtRET) forms a marked prominence on the lateral side of the sulcus extensorius proximal to the pons supratendineus; a sharp crest bounds the lateral margin of the sulcus extensorius proximal to the pons supratendineus; the medial attachment point for the retinaculum m. fibularis (mtRMF) laterad of the sulcus extensorius is poorly marked; the lateral attachment of the retinaculum m. fibularis (ltRMF) is aligned parallel to the shaft on the lateral facies; and it lacks a foramen penetrating the corpus cranially, close to the proximal margin of the condylus lateralis. Tarsometatarsus: This is relatively short, with those of species of Columba shorter, and those of Alopecoenas, Caloenas, Didunculus, Gallicolumba and Goura all relatively longer; the cotyla medialis in medial view is projected dorsally; the medial foramen of the foramina vascularia proximalia is elongate and much larger than its lateral counterpart; the foramina vascularia proximalia are widely separated by a low ridge in a sulcus bounded by prominent ridges; the foramen vasculare proximale medialis lies close (less than the foramen width) to the medial edge of the shaft; the tuberositas m. tibialis cranialis is distal to the foramen vasculare proximale medialis in the sulcus extensorius and is therefore close to the medial margin; and the lateral profile of the shaft leading to trochlea metatarsi IV is slightly concave.	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E9717024CC20A3D335729FC62FCD5.taxon	materials_examined	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. Remarks: Ducula pacifica is widespread in the tropical Pacific, with a range extending from the Bismarcks and New Guinea east through the Solomons, Vanuatu, Fiji, Tonga and Samoa to Niue and the Cook Islands (Steadman, 2006 a; Dickinson & Remsen, 2013). In Fiji, Du. pacifica is found only on small islands, but the endemic Du. latrans typical of the larger islands is sympatric with Du. pacifica on some smaller islands (Watling et al., 2001; Steadman, 2006 a). In Tonga, Du. pacifica is the only resident member of this genus now, but Steadman (2006 a: 335) stated that it ‘ coexisted … with two larger, extinct or extirpated congeners, D. latrans and D. undescribed sp., even on small, flat islands in the Ha'apai Group’. However, based on the data available to us, Du. pacifica and Du. latrans broadly overlap in all skeletal measurements (Appendices 2 and 3); therefore, we do not know how the two species were separated. In the Tongan fossil sample, although the bones are rather fragmented, as is typical of archaeological faunas, all fall within a narrow size range, in part captured by measurements below and in Appendices 2 and 3, and no evidence was seen to suggest that two taxa might be represented. Therefore, all bones are assumed to represent the extant species of Ducula in Tonga, i. e. Du. pacifica. The other large species of Ducula is much larger and is considered separately below. Ancient mounds and oral traditions recording the construction of platforms from which to snare Pacific pigeons show the cultural significance of these birds in prehistoric times throughout Tonga (Burley, 1996). Measurements: See Appendices 2 and 3. Coracoid: length from base of cotyla scapularis to top of acrocoracoideus, mean 12.1 mm, range 11.5 – 13.2 mm, SD = 0.60, N = 10; minimum shaft width, mean 3.6 mm, range 3.3 – 3.9 mm, SD = 0.16, N = 17. Ulna: SW 3.6, 3.8 mm; width of condylus dorsalis, 6.5, 7.0 mm. Carpometacarpus: PW 10.1 mm. Femur: PW 8.4, 8.9, 9.0, 9.1 mm. Tarsometatarsus: shaft width fossa metatarsi I, mean 4.3 mm, range 4.3 mm, N = 3; depth of trochlea metatarsi III, mean 3.8 mm, range 3.2 – 4.9 mm, SD = 0.58, N = 6. 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. Remarks: These specimens are either too incomplete or otherwise undiagnostic for confident generic attribution among columbids and are therefore referred only to columbids, but their size conforms with that of Du. pacifica. DUCULA SHUTLERI WORTHY & BURLEY SP. NOV., SHUTLER'S FRUIT PIGEON (FIG. 4) h t t p: / / z o o b a n k. o r g / u r n: l s i d: z o o b a n k. o r g: a c t: C 7 BC 1177 - FC 61 - 4 BA 8 - 8 BBC-C 770 C 6884 A 20 Holotype: NMNZ S. 48354, field number Of 1.05.12.08, left tarsometatarsus, collected by David Burley et al. on 6 June 2005 (Fig. 4 A). Paratypes: NMNZ S. 48355, field number Of 1.05.21.09, distal right tarsometatarsus preserved distal to metatarsal facet; NMNZ S. 48356, field number Of 1.05.22.04, left tarsometatarsus minus trochlea; NMNZ S. 48357, field number Of 1.05.22.09, distal left tarsometatarsus (Fig. 4 B). 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 1990 s and was a part of Burley's Tongan field projects in the 1990 s and early 2000 s. Type locality: Layer 8, 70 – 80 cm depth, Unit 12, Ofu site, Ofu Island, Vava'u Group, Tonga. Diagnosis: A species of Ducula with tarsometatarsus much longer than the largest extant species (Du. galeata of the Marquesas and Du. goliath of New Caledonia) or the extinct Du. tihonireasini of Mangareva and Du. david of Uvea, but of equal length to those of Du. harrisoni of Henderson Island and Du. lakeba of Fiji (see Table 7). Tarsometatarsus is more robust than those of Du. harrisoni and Du. lakeba and all other large species of Ducula except that of Du. david, as defined by proximal width and maximal shaft width across the articular facet in fossa metatarsi I as a proportion of length (Table 7). The dorsal medial lineae intermusculari is weakly marked distal to the metatarsal facet, and the shaft is relatively compressed dorsal to fossa metatarsi I. Measurements (in millimetres): Holotype NMNZ S. 48354 (Of 1.05.12.08). Total length, 47.6; length to proximal margin foramen vasculare distale, 37.0; length to distal side of the metatarsal facet, 28.8; length from posterior edge cotyla medialis to proximal end of fossa metatarsi 1 (length 2 of Wragg & Worthy, 2006), 23.2; length from posterior edge of cotyla medialis to distal end of medial prominence of fossa metatarsi 1 (length 3 of Wragg & Worthy, 2006), 27.4; proximal width, 13.8; shaft width at metatarsal facet, 6.2; least shaft width distal to metatarsal facet, 5.5; width of trochlea metatarsi III, 4.3; depth of trochlea metatarsi III, 5.0. Paratypes: See Table 7 and NMNZ S. 48355 (Of 1.05.21.09), dR tmt, depth of trochlea metatarsi III, 5.0; NMNZ S. 48356 (Of 1.05.22.04), L tmt, preserved length, 41.2; NMNZ S. 48357 (Of 1.05.22.09), 1 dL tmt, preserved length, 36.3; depth of trochlea metatarsi III, 5.1. Referred material: Ducula shutleri Hopoate: Hop. 14. TP 2.06, sL tmt; NISP = 1, MNI = 1. Falevai: Ka 2.05.03.15 a, L scap; Ka 2.05. XX. 00 FE 0 a, sR tmt; NISP = 2, MNI = 1. Ofu: Of 1.05.11.05, R scap; Of 1.05.12.06, L cor, sR tmt; Of 1.05.12.07, L scap, dL cmc, sL tmt; Of 1.05.12.07, pR fem; Of 1.05.12.08, R scap, cran and sternal pts L cor, 2 sR cor, dL tib; Of 1.05.12.09, dR tib, dR tmt; Of 1.05.12.10, ant stern, R scap, cran and sternal pts 1 R cor, sternal pts LandR cor, pL fem; Of 1.05.20.06, pL fem; Of 1.05.20.06, L scap; Of 1.05.20.07, R tib (missing proximal end and distal condyles); Of 1.05.20.07, sternal pt L, cran pt L cor; Of 1.05.20.07, L fem; Of 1.05.20.08, dL tib, sR tmt, pR ulna, R MII. 1; Of 1.05.20.09, 3 pL 1 dR fem; Of 1.05.20.09, 2 L scap; Of 1.05.20.10, cran pt R cor, L scap, dL tib; Of 1.05.21.06, R (- d) fem; Of 1.05.21.08, sL tmt; Of 1.05.21.08, pR dL rad; Of 1.05.21.09, R scap; Of 1.05.21.10, s + dR tmt, sR tmt; Of 1.05.21.10, s + pL sL ulna; Of 1.05.21.10, R scap; Of 1.05.21.10, dR ulna; Of 1.05.21.10, pR fem; Of 1.05.21.11, cran pt R cor; Of 1.05.22.04, sR cor; Of 1.05.22.05, pLdL cmc; Of 1.05.22.07, pR rad, dL cmc, L scap, 1 sR cor, sR tib, sL tmt; Of 1.05.22.08, pLdL rad, L scap; Of 1.05.22.09, 2 pL 3 dL tmt, 1 R one cran pt R one sternal pt R cor; Of 1.05.22.09, 2 pL rad, R scap; Of 1.05.22.10, two cran pt L 1 R cor, pt pL dR tmt; Of 1.05.22.10, 3 pL 1 dL fem, 2 sR tib, 1 dL cmc; including type material NISP = 93, MNI = 11. Vuna: Pa 1.04.12.08, pL fem; Pa 1.04.12.10, R scap; Pa 1.04.22.04, pL fem medullary bone; Pa 1.04.23.09, dR tmt; Pa 1.04.24.09, pR fem; NISP = 5, MNI = 2. Tentatively referred material: Identified as ‘ Columbid lge, cf. Ducula ’. Ofu: Of 1.05.12.05, pLdR rad, dLdR ulna, two ulnar shaft frags; Of 1.05.12.06, sL ulna, 2 sR tib; Of 1.05.12.08, d + sR rad, pR rad; d + sL sL sR d + sR cmc; Of 1.05.12.09, 3 dL cmc, sL ulna, sL tib; Of 1.05.12.10 a, sR hum, sL ulna; Of 1.05.14.08, sL fem; Of 1.05.15.05, dR rad; Of 1.05.20.06, sR fem; Of 1.05.22.05, L scap, dL rad; Of 1.05.22.06, 3 pL two pt shaft ulnae, two pts 1 L cor, pt R cmc, 2 R scap, 1 dR rad, 2 sR 1 sL tib, sLsR fem, pt sR tib; Of 1.05.22.09, 1 dL rad, sLsR tib, pmx; Of 1.05.22.10, 1 dL 2 dR cmc; NISP = 51, does not alter Ofu MNI. Vuna: Pa 1.04.12.11, sR tmt; Pa 1.04.17.08 b, sR cor; Pa 1.04.24.09, sL tmt; NISP = 3, MNI for Vuna unchanged by these. Description and comparison In addition to the diagnostic features described above, Du. shutleri has the following features. Humerus: Humeri are represented by a part of a right shaft Of 1.05.12.10 a with a minimum dorsoventral width of 8.5 mm, which lacks diagnostic features. However, it is larger and more inflated craniocaudally than the Caloenas specimens described above, e. g. the dR humerus in lot Of 1.05.12.10. The specimen tentatively referred to Du. shutleri is larger than MNZ S. 37316, a specimen of an undetermined large species of Ducula from Vitilevu, available specimens of Du. lakeba (see Worthy, 2001: table 8) and those of Du. harrisoni, which apparently had somewhat diminutive wings (Wragg & Worthy, 2006). Ulna: The most informative ulnar specimens are a well-preserved proximal third of a bone (Of 1.05.20.08, Fig. 4 F, PW = 11.2 mm, least SW = 6.3 mm) and a distal right ulna (Of 1.05.21.10, DW = width 11.0 mm, width of condylus dorsalis 10.1 mm). These reveal that the ulna is rather larger than those referred to Caloenas carpometacarpus Of 1.05.22.05, in ventral view. H, left coracoid missing most of processus acrocoracoideus Of 1.05.22.10, in dorsal view. I, omal end of left coracoid Of 1.05.22.10, in dorsal view. Scale bars: 10 mm. Abbreviations: artic., articularis; dtRET, the distal attachment point for the retinaculum extensorium tibiotarsi; ilaa, insertion for the ligamentum acrocoraco-acromion acrocoracoid; ilap, insertion for the ligamentum acrocoraco-procoracoideum; proc., processus; ptRET, the medial tuberosity for the proximal attachment of the retinaculum extensorium tibiotarsi; tub. A, tuberculum A; tub. m., tuberositas musculus. Data are from Worthy (2001), Wragg & Worthy (2006) and Rigal et al. (2018). The PW / L 2 and SWmf / L 2 ratios are given as percentages. Abbreviations: DW, distal width; Ldmf, length to distal side of the metatarsal facet from eminence intercotylaris; L 2, length from posterior edge cotyla medialis to proximal end of fossa metatarsi 1 (length 2 of Wragg & Worthy, 2006); L 3, length from posterior edge cotyla medialis to distal end of medial prominence of fossa metatarsi 1 (length 3 of Wragg & Worthy, 2006); PW, proximal width; SWdm, least shaft width distal to metatarsal facet; SWmf, shaft width at metatarsal facet; TL, total length; WTIII, width of trochlea of metatarsi III.	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E97170254C20E3CCB5483FAE7FCC8.taxon	discussion	Remarks: Material was referred to Ptilinopus by general similarity of form and size to compared specimens of Ptilinopus and especially to Ptilinopus regina Swainson, 1825 SAM B 37060, NMV B 16328, including the following features from Worthy et al. (2015): coracoid (terminology after Worthy, 2012) tuberculum A separated by a distinct groove from and less dorsally protuberant than ilaa; ilaa offset cranially from ilap, meaning that in medial view, the facies articularis clavicularis is longer craniosternally than dorsoventrally wide; ilaa and ilap linked by a slightly arched crest lacking a noticeable notch; pneumatic foramina present under facies articularis clavicularis; and processus procoracoideus short. In the Tongan region, two Ptilinopus species are found, the smaller Ptilinopus perousii Peale, 1849 and larger Ptilinopus porphyraceus (Temminck, 1821) (see Worthy & Anderson, 2009; Dickinson & Remsen, 2013). Skeletal material of Pt. perousii was not available to us, but the following material was all of a size similar to that of Pt. regina SAM B 37060 and NMV B 16328 and rather smaller than Pt. porphyraceus, e. g. MNZ O. 16391 (see Goodwin, 1967).	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E97170254C20E3CCB5483FAE7FCC8.taxon	materials_examined	Material: Hopoate: NISP = 9, MNI = 2. Nukuleka: NISP = 2, MNI = 1. COLUMBID SP. INDET. SMALL PIGEON Material: Hopoate: NISP = 5, MNI = 1. Vuna: NISP = 1, MNI = 1. Remarks: These specimens are too fragmentary for certain generic referral and are unlikely to increase the MNI for small taxa (Alopecoenas and Ptilinopus) known at the generic level. 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. Remarks: These specimens are too fragmentary for certain generic referral and are unlikely to increase the MNI for taxa known at the generic level.	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E97170254C2013F96531EFD8BFEFA.taxon	materials_examined	Material: Hopoate: Hop. 1 4. TP 1.1 0, dR hum; NISP = 1, MNI = 1. Remarks: Cacomantis flabelliformis is resident in the tropical Pacific from Papua New Guinea to Fiji and in Australia (Higgins, 1999; Dickinson & Remsen, 2013). This Tongan specimen was a good match in both size and morphology for SAM B. 55124, a specimen from South Australia. It might represent a vagrant from or an extension of the former range of the population of the Fijian form of this species.	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E9717025BC2013D31572BFCF6F99A.taxon	materials_examined	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. Measurements (in millimetres): Ka 2.05. XX. 00 FE 0 b, dL fem, SW = 3.8, DW = 8.9; Of 1.05.03.07, dR tib, DW = 5.9; Of 1.05.08.06, sR hum, SW = 3.0; Of 1.05.08.09 a, pR tib, PW articular surface = 7.7; Ka 1.05.10.14, dL fem, DW = 7.5; Ka 1.05.10.15, dR hum, DW = 6.2; Ka 1.05.12.15, dR tib, DW = 6.2; Pa 1.04.19.08, dR tib, DW = 6.4. Remarks: These specimens are indistinguishable from Hypotaenidia philippensis (= Gallirallus philippensis, sensu Kirchman, 2012), which includes 22 subspecies (Dickinson & Remsen, 2013). In describing the prehistoric fossil bones of Gallirallus from Tonga, Kirchman & Steadman (2005) attributed most from Ha'apai and Tongatapu to Gallirallus philippensis. However, from ' Eua, they noted that all remains that post-dated human arrival were from Gallirallus philippensis, but that in older layers, a now extinct species (Gallirallus vekamatolu) was the sole one present. Material of Hypotaenidia philippensis ecaudata (J. F. Miller, 1783), the subspecies in the Tongan region, was not available to us, but the Tongan bones were qualitatively similar to and about the same size as SAM B. 36299, an example of the Australian form Hypotaenidia philippensis mellori (Mathews, 1912), with which they were compared. All bones fell within the size range of those given for Samoa, Tonga and Vanuatu for this sexually size-dimorphic species (Kirchman & Steadman, 2005) and were much smaller than for those of the following species. HYPOTAENIDIA VAVAUENSIS WORTHY & BURLEY SP.	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E97170263C2393FED5475FAE8FA1B.taxon	materials_examined	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. Remarks: We follow Sangster et al. (1998) and Gill et al. (2010) in recognizing the swamphens from Australia, Moluccas, New Guinea through Fiji, Tonga, Samoa and Niue as a distinct species. It is a common, widespread species in the tropical Pacific (Steadman, 2006 a; Worthy et al., 2015). The coracoid is distinguished from those of other rails in the region by its elongate crista procoracoideus, which extends close to the crista medialis, and less medially projected processus acrocoracoideus. The tibiotarsi are smaller than those referred to H. vavauensis and differ by, in lateral aspect, the caudal side of the condylus lateralis extending further proximal than the cranial side, having a thicker margin to the medial side of the sulcus extensorius, and a wider incisura intercotylaris such that the condylus medialis lies wholly mesad of the canalis extensorius.	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E97170263C2383F87539AFDB9FB69.taxon	materials_examined	Material: Nukuleka: NISP = 3, MNI = 1. Falevai: NISP = 3, MNI = 1. Ofu: NISP = 20, MNI = 2. Vuna: NISP = 8, MNI = 1. Remarks: This species is abundant in the tropical Pacific, and the specimens matched the reference specimen, MV B. 12741, well. PROCELLARIID MAGN. ARDENNA PACIFICA (GMELIN, 1789) Material: Otea: Ka 1.05.12.18, pL cmc, NISP = 1, MNI = 1. Remarks: This worn specimen lacks the processus extensorius, is the size of MV B. 12741, but differs with a more robust os metacarpale minus. PROCELLARIID SP. LGE	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E97170263C2383F87539AFDB9FB69.taxon	materials_examined	Material: Otea: Ka 1.05.01.21, dL hum, NISP = 1, MNI = 1. Measurements: Maximum SW = 5.2 mm, est. DW (flexor process missing) 11.0 mm. Remarks: This specimen compares in size with Ps. rostrata, NMNZ O. 24691, with DW = 11.7 mm and SW = 5.1 mm. As in Ps. rostrata (NMNZ O. 23900, 24691), the impression for the brachialis anticus within the fossa brachialis is well defined distally and near circular, whereas it is poorly defined distally and oval in Pterodroma.	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E97170262C2383D0552ABFB08F9B3.taxon	materials_examined	Material: Falevai: Ka 2.05.13.14, pL fem (lacking proximal three-quarters of trochanter), NISP = 1, MNI = 1. Vuna: Pa 1.04.11.08 a, dR tib, NISP = 1, MNI = 1. Remarks: Ardea modesta is treated as a distinct species, following Gill et al. (2010) and references therein. It is distributed from Asia to Australasia including the Solomons, where it is ‘ mostly vagrant’ (Dickinson & Remsen, 2013). These specimens were compared with the respective elements from multiple specimens each of Ardea modesta, Egretta sacra (Gmelin, 1789), Egretta novaehollandiae (Latham, 1790) and Nycticorax caledonicus Gmelin, 1789. The proximal femur is distinctive, although the proximal three-quarters of the trochanter is lost, because it preserves the distal part of the trochanter and the scars for m. iliotrochanterica cranialis and the m. ischiofemoralis. Its width (6.5 mm) at the distal end of the trochanter is greater than in Ny. caledonicus (5.4 – 6.4 mm, N = 10) and much bigger than that of Eg. novaehollandiae (e. g. 5.0 mm, SAM B. 32854) or Eg. sacra. The specimen shows that the crista trochanteris extends distally well past the level of the scar for m. iliotrochanterica cranialis, as seen in Ardea and Egretta, rather than being relatively short and ending level with that scar in Nycticorax. Also, in Ardea and Egretta, there is a shallow notch, seen in lateral aspect, caudally slightly distal to the obturator impression, but this notch is much deeper and more distally placed in Nycticorax. In all features, the Tongan bone matched Ardea modesta, e. g. SAM B. 31393. The distal tibiotarsus Pa 1.04.11.08 a shares with Ardea modesta and Eg. novaehollandiae some features and differs from Nycticorax in several ways: the shaft is relatively more slender; the medial margin of the pons supratendineus is narrowly separated from the shaft margin (in Nycticorax, the separation is wider, being absolutely wider than the width of the sulcus extensorius); the condylus lateralis is more cranially projecting and forms a near right angle with the shaft proximally (less projecting, slopes distally at proximal margin with shaft); in anterior view, the condylus lateralis projects laterad of the shaft (not so); in distal aspect, the incisura intercondylaris is broad, wider than the width of condylus lateralis and flat bottomed (narrower, round bottomed); the notch in condylus medialis distally is poorly marked (well marked); and in lateral view, the scar for the fibular retinaculum ends well proximad to the level of the proximal side of condylus lateralis (same level). The size of Pa 1.04.11.08 a matches that of Ardea modesta (Table 18) and is considerably larger than Eg. sacra or Eg. novaehollandiae. Therefore, the Tongan specimens cannot be referred to a species of Nycticorax and fit in qualitative features and in size with Ardea modesta, and thus are referred to this taxon, which is not a resident of Tonga. These specimens, therefore, probably represent vagrants from areas more west, such as the Solomons.	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E97170261C23B3CD45176FABCFC1F.taxon	materials_examined	Material: Ofu: NISP = 6, MNI = 1 (does not change MNI for certainly identified material of this taxon). Remarks: The genus Eclectus is represented by the extant Ec. roratus (S. Müller, 1776), a polytypic species (nine subspecies) ranging from the Moluccas through to New Guinea, the Solomons and northern Australia (Dickinson & Remsen, 2013). The extinct species Ec. infectus was described on archaeological specimens from Tonga and from Malakula in Vanuatu (Steadman, 2006 c). Many more specimens attributed to this form were described from Teouma, on Efate in Vanuatu (Worthy et al., 2015). The bones described here were qualitatively the same as those from Vanuatu.	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E97170261C23B3CCA539CFDE9F910.taxon	materials_examined	Material: Hopoate: NISP = 1, MNI = 1.; Nukuleka: NISP = 1, MNI = 1. Ofu: NISP = 6, MNI = 2. Vuna: NISP = 2, MNI = 1. Remarks: All from Lapita layers.	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
7E2E97170261C23B3D7B551AFD8AFAAC.taxon	materials_examined	Material: Falevai: Ka 2.05.03.12 a, juv R tmt, NISP = 1, MNI = 1. Ofu: Of 1.05.03.07, dR hum, NISP = 1, MNI = 1. Remarks: This is a common migrant species to Oceania (Dickinson & Remsen, 2013). The specimens were a good match qualitatively and by size with NMV B. 20033.	en	Worthy, Trevor H., Burley, David V. (2020): Prehistoric avifaunas from the Kingdom of Tonga. Zoological Journal of the Linnean Society 189: 998-1045
