APTERYGIFORMES Kiwi

Symbols and Abbreviations
➤ Indicates a species (cf. subspecies)
BMNH, Natural History Museum (bird section), Tring, United Kingdom
NMNZ, Museum of New Zealand Te Papa Tongarewa, Wellington

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Order APTERYGIFORMES: Kiwi

The Checklist Committee (2010) placed kiwi in the order Casuariiformes, as they were then considered sister to cassowaries + emus (A. Cooper et al. 1992, 2001; A. Cooper 1997; Haddrath & Baker 2001). More recent studies have revealed kiwi to be sister to the extinct elephant birds (Aepyornithidae) from Madagascar (M. Phillips et al. 2010; Mitchell, Llamas et al. 2014; Grealy et al. 2017; Yonezawa et al. 2017), and most authorities recognise seven orders within the Palaeognathae. We follow Cracraft (2013), Dickinson & Remsen (2013), Clements et al. (2019), de Moya et al. (2019), Urantówka et al. (2020), and F. Gill et al. (2021) in returning kiwi in their own order (Apterygiformes), which was the recommendation of the Checklist Committee in 1953, 1970, and 1990.

Family APTERYGIDAE G.R. Gray: Kiwi

Apteryginae G.R. Gray, 1840: List Gen. Birds (1^st edition): 63 – Type genus Apteryx Shaw, 1813.

Genus Apteryx Shaw

Apteryx Shaw, 1813: Nat. Miscell. 24(286): pls 1057–1060 – Type species (by monotypy) Apteryx australis Shaw.

Apternyx Swainson, 1837: Nat. Hist. Classif. Birds 1: 119. Unjustified emendation.

Apternix Agassiz, 1846: Nomen. Zool. Index Univ. Aves 2. Unjustified emendation.

Pseudapteryx Lydekker, 1891: Cat. Fossil Birds Brit. Museum: 218 – Type species (by monotypy) Pseudapteryx gracilis Lydekker = Apteryx owenii Gould.

Stictapteryx Iredale & Mathews, 1926: Bull. Brit. Ornith. Club. 46: 76 – Type species (by original designation) Apteryx owenii Gould.

Kiwi Verheyen, 1960: Bull. Roy. Soc. d’Anvers 15: 10. Unnecessary nomen novum for Stictapteryx Iredale & Mathews, 1935.

Kiwi were once throughout the main islands of New Zealand: North, Little Barrier / Hauturu, Great Barrier / Aotea, South, D’Urville, and Stewart / Rakiura Islands; probably originally in all vegetated habitats (Worthy & Holdaway 2002). Increasingly restricted since European settlement to residual forests and adjacent scrub and rough farmland. North Island brown kiwi (Apteryx mantelli), but not other forms, have colonised exotic forests (Germano et al. 2018).

Since compilation of the 1990 Checklist (Checklist Committee 1990), several studies have been published on the molecular biology of kiwi (e.g. A. Baker et al. 1995; Burbidge et al. 2003; Shepherd & Lambert 2008; Shepherd et al. 2012; Weir et al. 2016; Scofield et al. 2021; Shepherd, Tennyson et al. 2021; Bemmels et al. 2021). Kiwi, particularly brown kiwi, are marked by mostly allopatric genetic diversity in both modern and extinct populations. This is not closely associated with morphological differences, making delineation of species limits difficult (Shepherd & Lambert 2008). For brown kiwi we follow Holdaway et al. (2001) and Tennyson, Palma et al. (2003) in recognising a North Island species (A. mantelli) and two extant South Island species (A. rowi and A. australis – the latter having two subspecies). Several historical names were not based on localised specimens or adequate descriptions and are unable to be referred to known taxa. These include:

Dromiceius Novae-Zelandiae, Lesson, 1828: Manuel d’Ornith. 2: 210 – Bay of Islands (and also Apteryx australis novae-zelandiae (Lesson), A. a. novaezealandiae [sic] (Lesson), and Dromiceius novaezealandiae – see Mathews 1935 and 1937a, and Lee & Bruce 2019).

Apteryx major Ellman, 1861: The Zoologist 19: 7468 – New Zealand.

Apteryx fusca Potts, 1873: Trans. N.Z. Inst. 5: 196 – West Coast. Not Apteryx fusca Rowley, 1875.

The authorship of Apteryx and Apteryx australis is restricted to Shaw, following ICZN (1916), and as supported by Dickinson et al. (2006). Apteryx haastii Potts, 1872 was used for great spotted kiwi until 2021, when the two syntype specimens for A. haastii were shown to be hybrids between A. owenii and A. rowi (Shepherd, Tennyson et al. 2021).

➤ Apteryx mantelli Bartlett
North Island Brown Kiwi | Kiwi-nui

Apteryx Mantelli Bartlett, 1852: Proc. Zool. Soc. London 1850 (18): 275, pl. 30, figs 3–4; pl. 31, fig. 2 – North Island, restricted to Ohakune (fide Shepherd et al. 2009, Bull. Brit. Ornith. Club 129: 197).

Apteryx mantellii Bartlett; G.R. Gray 1862, Ibis 4: 233. Unjustified emendation.

Apteryx australis var. Mantelli Bartlett; Finsch 1872, Journ. für Ornith. 20: 263.

Apteryx bulleri Sharpe, 1888: Proc. Wellington Phil. Soc.: 6 – North Island.

Apteryx australis mantelli Bartlett; Checklist Committee 1953, Checklist N.Z. Birds: 13.

Apteryx mantelli Bartlett; Holdaway et al. 2001, New Zealand Journ. Zool. 28(2): 125, 175.

Apteryx apteryx mantelli Bartlett; Grealy, Phillips, Miller, Gilbert, Rouillard, Lambert, Bunce & Haile 2017, Mol. Phyl. Evol. 109: 154. Error for “Apteryx australis mantelli Bartlett”.

Apteryx mantelli is treated as a full species because of the marked differences in plumage noted in its original description and significant genetic differences (A. Baker et al. 1995; Burbidge et al. 2003). In this, we follow Holdaway et al. (2001), Worthy & Holdaway (2002), and Tennyson, Palma et al. (2003). Originally recorded throughout the North Island. Now restricted to isolated and declining populations in Northland, Taranaki, western slopes of Ruapehu, King Country, inland northern Hawke’s Bay, Urewera, and Coromandel; introduced to Ponui, Kawau, Kapiti (1910–40), and Hauturu / Little Barrier Islands (Miskelly & Powlesland 2013; Germano et al. 2018). Birds introduced to the latter were from Taupo in 1913 (Oliver 1955: 49), but lice from Hauturu / Little Barrier Island birds indicate survival of the former resident population reported by Reischek (1887) (see Palma 1991). Genetic studies have revealed a high degree of genetic differentiation between remnant populations (A. Baker et al. 1995; Burbidge et al. 2003; Shepherd & Lambert 2008; Weir et al. 2016; Bemmels et al. 2021). Common in Late Pleistocene and Holocene deposits and in middens throughout much of the North Island (Worthy & Holdaway 2002). However, mtDNA from bones from Martinborough (Wairarapa) and Lake Poukawa (Hawke’s Bay) clustered with A. rowi (Shepherd & Lambert 2008). This suggests that the now extinct brown kiwi populations from the south of the North Island may have been A. rowi not A. mantelli.

➤ Apteryx rowi Tennyson, Palma, Robertson, Worthy & Gill
Rowi | Okarito Brown Kiwi

Apteryx australis australis Shaw & Nodder [sic]; Checklist Committee 1953, Checklist N.Z. Birds: 13. In part.

Apteryx sp. Okarito Form; Worthy & Holdaway 2002, Lost World of Moa: 219.

Apteryx rowii Burbidge, Colbourne, Robertson & Baker, 2003 (Apr.): Conservation Genetics 4: 172, 176. Nomen nudum.

Apteryx rowii Marsh, 2003 (5 Jul.): New Zealand Listener 5: 29. Nomen nudum.

Apteryx rowi Tennyson, Palma, Robertson, Worthy & Gill, 2003: Rec. Auck. Inst. Museum 40: 57 – South Okarito Forest, South Westland.

Until recently, confined to Okarito, South Island. Brown kiwi have been recorded from Okarito since at least 1867 under the names of roa, rohi, and rowi, and have been recognised as distinct from South Island brown kiwi since the 1950s (for nomenclatural history see Tennyson, Palma et al. 2003). Genetic studies of mtDNA supported morphological differences and suggested that the Okarito brown kiwi is the sister taxon of the North Island brown kiwi (A. Baker et al. 1995; Burbidge et al. 2003; Shepherd & Lambert 2008; Weir et al. 2016; Bemmels et al. 2021), and not closely related to A. australis. Closer relationship to A. mantelli than to A. australis is also supported by evidence from lice (Palma & Price 2004). Endangered; c. 450 birds remaining in 2018 (Germano et al. 2018). Translocated to Mana and Blumine Islands during 2010–12 (Miskelly & Powlesland 2013).

Studies of ancient mtDNA sequences (cytochrome-b, control region, derived from bones found in natural deposits) indicates that extinct brown kiwi populations north of Okarito on the South Island West Coast (Buller, Takaka Hill) and from Martinborough (Wairarapa) and Lake Poukawa (Hawke’s Bay), in the south-east North Island, form a single clade with extant A. rowi that is sister to remaining North Island brown kiwi (Shepherd & Lambert 2008; Weir et al. 2016). This suggests that A. rowi had a former range from Okarito, up the South Island West Coast and into the southern North Island.

➤ Apteryx australis Shaw
Southern Brown Kiwi | Tokoeka

Southern brown kiwi was formerly abundant throughout Stewart Island / Rakiura and at low elevations in southern and south-eastern South Island; however, its South Island range had contracted to Fiordland by 1893 (Marchant & Higgins 1990; Weir et al. 2016; Scofield et al. 2021). Several studies combining modern and ancient DNA sequences have demonstrated that populations of A. australis at Haast, Fiordland, and Stewart Island, and extinct populations in Southland and east of the Southern Alps / Kā Tiritiri o te Moana formed a single clade with deep diversity (Shepherd & Lambert 2008; Weir et al. 2016; Scofield et al. 2021; Undin et al. 2021). These include the populations of “large” kiwi from eastern areas that Worthy (1997d, 1998b) found to have significantly smaller and stouter leg bones than extant A. australis. The Stewart Island / Rakiura birds form a monophyletic clade divergent from South Island birds (Burbidge et al. 2003; Shepherd & Lambert 2008; Weir et al. 2016; Scofield et al. 2021; Bemmels et al. 2021), supporting the subspecies status attributed them below. Also present on Secretary, Resolution, Cooper, and Long Islands, Fiordland, with introduced populations on Parrot and Indian Islands, Dusky Sound (Miskelly & Powlesland 2013). Introduced to Kapiti Island where the stock is now hybrid between A. australis and A. mantelli (Checklist Committee 1990). Scofield et al. (2021) demonstrated that the type specimen of Apteryx australis must have been collected on Stewart Island / Rakiura. This means that the Stewart Island brown kiwi (formerly known as A. australis lawryi) becomes the nominate subspecies. There are no scientific names available for the brown kiwi found in Fiordland and near Haast.

Apteryx australis subsp.
South Island Brown Kiwi | Tokoeka

Apteryx australis; Hutton 1871, Cat. Birds N.Z.: 23. Not Apteryx australis Shaw.

Apteryx australis australis Shaw & Nodder [sic]; Checklist Committee 1953, Checklist N.Z. Birds: 13. Not Apteryx australis Shaw.

Apteryx “eastern South Is.”; Holdaway et al. 2001, New Zealand Journ. Zool. 28: 125.

Apteryx “East South Is”; Holdaway et al. 2001, New Zealand Journ. Zool. 28: 175.

Apteryx (Eastern South Island)”; Holdaway et al. 2001, New Zealand Journ. Zool. 28: 185.

Apteryx sp. Eastern Kiwi; Worthy & Holdaway 2002, Lost World of Moa: 220.

Apteryx australis “Haast”; Miskelly et al. 2008, Notornis 55: 126.

Apteryx australis (Northern Fiordland); Miskelly et al. 2008, Notornis 55: 127.

Apteryx australis australis; Checklist Committee 2010, Checklist N.Z. Birds: 21. Not Apteryx australis Shaw.

Apteryx australis “northern Fiordland”; Robertson et al. 2013, Conservation status New Zealand birds, 2012: 2.

Apteryx australis “northern”; Robertson et al. 2017, Conservation status New Zealand birds, 2016: 5.

Apteryx australis “southern Fiordland”; Robertson et al. 2021, Conservation status Aotearoa New Zealand birds, 2021: 6.

Apteryx australis “Northern Fiordland”; Robertson et al. 2021, Conservation status Aotearoa New Zealand birds, 2021: 23.

South Island; Haast River to Arawata River, and Fiordland (C. Robertson et al. 2007). Disjunct populations, the result of postulated recent and ongoing population declines, are treated as distinct management units by the Department of Conservation (Burbidge et al. 2003; Weir et al. 2016; Germano et al. 2018; Bemmels et al. 2021; Scofield et al. 2021; Shepherd, Tennyson et al. 2021; Undin et al. 2021). Holocene remains widespread throughout the South Island (Worthy 1997d, 1998b; Worthy & Holdaway 2002).

Apteryx australis australis Shaw
Stewart Island Brown Kiwi | Rakiura Tokoeka

Apteryx australis Shaw, 1813: Nat. Miscell. 24(286): pls 1057–1060 – New Zealand, restricted to Stewart Island / Rakiura (fide Scofield et al. 2021, Cons. Gen. doi: 10.1007/s10592-021-01349-y: 5).

Apternyx australis (Shaw); Swainson 1837, in D. Lardner, The Cabinet Cyclopaedia 2(92): 346. Unjustified emendation.

Apteryx maxima Buller, 1891: Trans. N.Z. Inst. 24: 602 – Stewart Island. Junior primary homonym of Apteryx maxima Sclater & Hochstetter, 1861.

Apteryx lawryi Rothschild, 1893: Bull. Br. Ornith. Club 1: 61 – Stewart Island.

Apteryx Australis Shaw; Mathews 1930, Emu 29: 278.

Apteryx australis lawryi Rothschild; Checklist Committee 1953, Checklist N.Z. Birds: 13.

Stewart Island / Rakiura (main island, and Ulva Island in Paterson Inlet / Whaka a Te Wera), in forest and scrub; widespread and locally common (Marchant & Higgins 1990). Several midden records.

➤ Apteryx owenii Gould
Little Spotted Kiwi | Kiwi Pukupuku

Apteryx Owenii Gould, 1847 (12 Jun.): The Literary Gazette 1586: 433 – Middle [South Island], restricted to Nelson District (fide Oliver 1930, New Zealand Birds, 1^st edition: 61).

Apteryx Oweni Gould; Anon. 1870, Cat. Colonial Mus.: 74. Unjustified emendation.

Apteryx mollis Potts, 1873: Trans. N.Z. Inst. 5: 196 – Martins Bay.

Apteryx fusca Rowley, 1875: Ornith. Miscellany 2: 8. South Island. Nomen novum for “dark coloured A. owenii”.

Apteryx oweni Gould; Travers 1883, Trans. Proc. N.Z. Inst. 15: 186. Unjustified emendation.

Pseudapteryx gracilis Lydekker, 1891: Cat. Fossil Birds Brit. Museum: 218, fig. 53A – New Zealand.

Apteryx oweni occidentalis Rothschild, 1893: Bull. Br. Ornith. Club 1: 61 – “west coast of the South and North Islands”, restricted to Dusky Sound, Fiordland (fide Hartert 1927, Novit. Zool. 34: 31). Unjustified emendation.

Apteryx occidentalis Rothschild; Buller 1896, Trans. N.Z. Inst. 28: 358.

Apteryx owenii owenii Gould; Mathews & Iredale 1913, Ibis 1 (10^th series): 205.

Stictapteryx owenii iredalei Mathews, 1935: Bull. Br. Ornith. Club 55: 180 – North Island.

Apteryx oweni Gould; Checklist Committee 1953, Checklist N.Z. Birds: 13. Unjustified emendation.

Kiwi owenii (Gould); Verheyen 1960, Bull. Roy. Soc. d’Anvers 15: 10.

Apteryx owenii Gould; Checklist Committee 1990, Checklist Birds N.Z.: 8.

On European settlement, rare in the North Island: a specimen in BMNH from Mount Hector, Tararua Range (Buller 1876a); reported from Wangapopo Range, near Pirongia, King Country, by Reischek in Feb. 1882 (Reischek 1930: 177; the species name A. owenii was also recorded in the original German text, i.e. Reischek 1924). On European settlement, common in the South Island throughout forest areas of Marlborough, Nelson, Westland, and Fiordland (e.g. Potts 1873). Rapidly declined in the late 19^th Century. Few verified mainland records after 1930 (Ramstad et al. 2021). An adult breeding female from Banjo Creek, Westhaven Inlet, Jul. 1978 (NMNZ OR.023036; Worthy & Holdaway 1994: 307) and an adult female from Smyth River, South Westland, 1978 (NMNZ OR.023043) are A. owenii, not A. maxima (contra Marchant & Higgins 1990: 81). A. owenii also survived on D’Urville Island until the last birds were transferred to Long Island, Queen Charlotte Sound, in 1982 and 1987 (Ramstad et al. 2021). Common on Kapiti Island, where introduced in 1912 (Miskelly & Powlesland 2013; Ramstad et al. 2021). Kapiti birds have different lice from those of all mainland skins examined (Pilgrim & Palma 1982; Marchant & Higgins 1990: 85). Successfully transferred in the 1980s from Kapiti to Red Mercury (Whakau) Island (Coromandel), Hen Island / Taranga (Hauraki Gulf) and Long Island (Marlborough Sounds). In 1993 transferred from Kapiti to Tiritiri Matangi Island (Hauraki Gulf), in 2000 to Karori Sanctuary / Zealandia, Wellington, in 2008 to Chalky Island, Fiordland, in 2009 to Motuihe Island (Hauraki Gulf), and in 2015 to Anchor Island, Fiordland (Miskelly & Powlesland 2013; H. Robertson 2015). Frequent in Late Pleistocene and Holocene deposits, and in middens, throughout North and South Islands (Worthy & Holdaway 2002). Bruce & McAllan (1990), as first revisers, selected The Literary Gazette as the original publication of the name Apteryx owenii. See comments under rowi (A. rowi) regarding the status of Potts’ kiwi (A. haastii).

➤ Apteryx maxima Sclater & Hochstetter
Great Spotted Kiwi | Roroa

Apteryx maxima Sclater & Hochstetter, 1861: The Natural History Review: 506 – near Charleston, West Coast.

Apteryx maxima Verreaux [sic]; Anon. 1870, Cat. Colonial Mus.: 74.

Apteryx maxima Hutton, 1871: Cat. Birds N.Z.: 23, 75 – Westland. Junior primary homonym and junior synonym of Apteryx maxima Sclater & Hochstetter, 1861.

Apteryx maximus Verreaux [sic]; Rothschild 1893, Ibis 5 (6^th series): 576.

Apteryx grandis Grieve, 1913: Proc. Roy. Phys. Soc. Edinburgh 19: 63 – South Island.

Apteryx owenii maxima Hutton; Mathews 1935, Bull. Br. Ornith. Club 55: 180.

Apteryx maxima Sclater & Hochstetter, 1861; Palma, Worthy & Tennyson 2003, Tuhinga 14: 7. As nomen oblitum.

Apteryx haastii of authors. Not Apteryx haastii Potts, 1872: Ibis 2 (3^rd series): 35 – West Coast (= hybrid between A. owenii and A. rowi; Shepherd et al. 2021).

Apteryx maxima Sclater & Hochstetter, 1861; Shepherd, Tennyson, Robertson, Colbourne & Ramstad 2021, Avian Research 12:24: 11. Resurrected from nomen oblitum.

North-western South Island (C. Robertson et al. 2007). Extends across the divide (in high altitude beech forest) at various points between Arthur’s Pass and the Hope River (Germano et al. 2018). Bones in deposits are largely indeterminate, as they are generally morphologically inseparable from those of Apteryx australis (e.g. Worthy 1997d), but remains likely to be this species are known in high-altitude sites in north-west Nelson (Mount Owen, Mount Arthur) and in sites in the Honeycomb Hill cave system, Oparara. Some bones have been identified as A. maxima by DNA-typing (Shepherd & Lambert 2008), but this method is limited to relatively few well-preserved specimens. Not recorded from the North Island. Nineteen individuals from Gouland Downs were introduced to Hauturu / Little Barrier Island in 1915 in a failed bid to establish an island population (Oliver 1955). More recently, translocated to St Arnaud, Nelson Lakes in 2004–06, Flora Saddle, Mt Arthur in 2010–16, and Nina Valley, Lewis Pass in 2011–12 (Miskelly & Powlesland 2013; Toy & Toy 2020).

Until 2021, great spotted kiwi were known by the species name Apteryx haastii; however, the two syntype specimens of Apteryx haastii are hybrids between A. owenii and A. rowi (Shepherd, Tennyson et al. 2021, and see under A. rowi above). Apteryx maxima was made a nomen oblitum by Palma et al. (2003) because at that time Apteryx haastii and Apteryx maxima were considered to be synonyms. However, as they clearly are not synonyms (Shepherd, Tennyson et al. 2021), Article 23.11 of the ICZN Code (1999) does not apply. From accounts in Grieve (1913) it is clear that in the late 1880s Apteryx maxima was called A. grandis by various commercial collectors, notably James Dall.