APPENDIX 1 Fossil birds

Symbols and Abbreviations

➤Indicates a species (cf. subspecies)
† Indicates an extinct taxon
BMNH, Natural History Museum (bird section), Tring, United Kingdom
NMNZ, Museum of New Zealand Te Papa Tongarewa, Wellington

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Fossil Birds of New Zealand

This Appendix lists birds recorded as fossils in New Zealand from sediments older than the middle Pleistocene (≥1 Ma). It therefore includes material from the Kaimatira Pumice Sand of the Kai Iwi Group (Oxygen Isotope Stage 25–27, c. 1 Ma) found at Marton (Worthy 1997a; Tennyson & Tomotani 2021b). All younger extinct birds are members of the Recent fauna; all are species that persisted to human arrival and are covered in the main text. Before 2007, the pre-Pleistocene record of birds in New Zealand comprised mainly penguins, as reviewed by Fordyce (1991b). Until then, the record of the Tertiary terrestrial avifauna was restricted to two undescribed anatids from the Miocene lacustrine deposits at St Bathans, Otago (Fordyce 1991b). Subsequent investigations in this area have recovered a rich avifauna comprising at least 40 taxa (Tennyson, Worthy et al. 2013; Mather et al. 2019). In addition to the diverse taxa named below, the St Bathans assemblage includes moa (Dinornithiformes), geese (Anatidae: Anserinae), an eagle (Accipitridae), and several passerines (Passeriformes). Elsewhere, G. Mayr & Scofield (2015) attributed a partial humerus and a partial carpometacarpus from Paleocene marine sediments of the Waipara Greensand, North Canterbury, to a tropicbird. The remains were too fragmentary to be assigned to family, genus or species. Ages of taxa are described using the New Zealand geological timescale (R. Cooper 2004).

Order APTERYGIIFORMES: Kiwi

Family APTERYGIDAE G.R. Gray: Kiwi

Genus †Proapteryx T. Worthy, J. Worthy, Tennyson, Salisbury, Hand & Scofield

Proapteryx T. Worthy, J. Worthy, Tennyson, Salisbury, Hand & Scofield, 2013: Proc. 8th Int. Meeting Soc. Avian Paleont. Evol.: 67 – Type species (by original designation) Proapteryx micromeros T. Worthy, J. Worthy, Tennyson, Salisbury, Hand & Scofield.

➤ †Proapteryx micromeros T. Worthy, J. Worthy, Tennyson, Salisbury, Hand & Scofield
St Bathans Kiwi

Proapteryx micromeros T. Worthy, J. Worthy, Tennyson, Salisbury, Hand & Scofield, 2013: Proc. 8th Int. Meeting Soc. Avian Paleont. Evol.: 67 – St Bathans, Central Otago.

Known from two fossil remains only, from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage, Otago (Worthy, Worthy, Tennyson, Salisbury et al. 2013).

Genus Apteryx Shaw

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

➤ †Apteryx littoralis Tennyson & Tomotani
Marton Kiwi

Apteryx littoralis Tennyson & Tomotani, 2021 [2022]: Historical Biology 34: 353 – Marton, North Island.

Known from a Pleistocene (1 Ma) epoch tarsometatarsus found in shallow marine sediment north of Marton (Tennyson & Tomotani 2021b).

Order ANSERIFORMES: Duck-like Birds

Suborder ANSERES: Swans, Geese, and Ducks

Family ANATIDAE Leach: Swans, Geese, and Ducks

Anatidae Leach, 1819: Eleventh room. In Synopsis Contents British Museum 15th edition, London: 67 – Type genus Anas Linnaeus, 1758.

Subfamily ANSERINAE Vigors: Swans and Geese

Anserina Vigors, 1825: Zoological Journ. 2: 404 – Type genus Anser Brisson, 1760.

Genus †Notochen T. Worthy, Scofield, Hand, De Pietri & Archer

Notochen T. Worthy, Scofield, Hand, De Pietri & Archer, 2022: Zootaxa 5168: 45 – Type species (by original designation) Notochen bannockburnensis T. Worthy, Scofield, Hand, De Pietri & Archer.

➤ †Notochen bannockburnensis T. Worthy, Scofield, Hand, De Pietri & Archer
Bannockburn Swan

Notochen bannockburnensis T. Worthy, Scofield, Hand, De Pietri & Archer, 2022: Zootaxa 5168: 45 – St Bathans, Central Otago.

Known from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage from the lower Bannockburn Formation, Manuherikia Group; near St Bathans, Otago (Worthy, Scofield, Hand et al. 2022).

Subfamily OXYURINAE J.C. Phillips: Stiff-tailed Ducks

Oxyurinae J.C. Phillips, 1926: Nat. Hist. Ducks 4: 201 – Type genus Oxyura Bonaparte, 1827.

When described by Worthy et al. (2007), Manuherikia was placed in the extinct subfamily Dendrocheninae, created by Livezey & Martin (1988), and Dunstanetta in Tribus incertae sedis. However, Worthy & Lee (2008) and Worthy (2009) conducted extensive phylogenetic analyses of Manuherikia and Dunstanetta and found no support for Dendrocheninae, and that both the New Zealand genera are basal in a clade inclusive of traditional oxyurines. Accordingly, they advocated that Oxyurinae be expanded to encompass an enlarged set of taxa that included the fossil European Oligo-Miocene Mionetta and New Zealand Manuherikia and Dunstanetta, as well as the modern genera Malacorhynchus, Stictonetta, Thalassornis, Nomonyx, Oxyura, and Biziura. We follow this course here.

Genus †Manuherikia T. Worthy, Tennyson, Jones, McNamara & Douglas

Manuherikia T. Worthy, Tennyson, Jones, McNamara & Douglas, 2007: Journ. Syst. Palaeont. 5(1): 9 – Type species (by original designation) Manuherikia lacustrina T. Worthy, Tennyson, Jones, McNamara & Douglas.

➤ †Manuherikia lacustrina T. Worthy, Tennyson, Jones, McNamara & Douglas
Manuherikia Duck

Manuherikia lacustrina T. Worthy, Tennyson, Jones, McNamara & Douglas, 2007: Journ. Syst. Palaeont. 5(1): 10, figs 4G, H, 5D, H, I, 6D – St Bathans, Central Otago.

Known from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage from the lower Bannockburn Formation, Manuherikia Group; near St Bathans, Otago (Worthy et al. 2007).

➤ †Manuherikia minuta T. Worthy, Tennyson, Jones, McNamara & Douglas
Minute Manuherikia Duck

Manuherikia minuta T. Worthy, Tennyson, Jones, McNamara & Douglas, 2007: Journ. Syst. Palaeont. 5(1): 12, figs 4B, D, 5B, E, F, 6A – St Bathans, Central Otago.

Known from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage from the lower Bannockburn Formation, Manuherikia Group; near St Bathans, Otago (Worthy et al. 2007).

➤ †Manuherikia douglasi T. Worthy, Tennyson, Hand & Scofield
Douglas’ Duck

Manuherikia douglasi T. Worthy, Tennyson, Hand & Scofield, 2008: Journ. Royal Soc. N.Z. 38(2): 100, fig. 1A–H – St Bathans, Central Otago.

Known from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage from the lower Bannockburn Formation, Manuherikia Group; near St Bathans, Otago (Worthy et al. 2007, 2008).

➤ †Manuherikia primadividua T. Worthy, Scofield, Salisbury, Hand, De Pietri, Blokland & Archer
St Bathans Diving Duck

Manuherikia primadividua T. Worthy, Scofield, Salisbury, Hand, De Pietri, Blokland & Archer, 2022: Geobios 70: 96 – St Bathans, Central Otago.

Known from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage from the lower Bannockburn Formation, Manuherikia Group; near St Bathans, Otago (Worthy, Salisbury, Hand et al. 2022a). The stratigraphic location of the fossils indicate that they are younger than M. lacustrina, which is found in lower beds within the same formation (Worthy et al. 2019).

Genus †Dunstanetta T. Worthy, Tennyson, Jones, McNamara & Douglas

Dunstanetta T. Worthy, Tennyson, Jones, McNamara & Douglas, 2007: Journ. Syst. Palaeont. 5(1): 17 – Type species (by monotypy) Dunstanetta johnstoneorum T. Worthy, Tennyson, Jones, McNamara & Douglas.

➤ †Dunstanetta johnstoneorum T. Worthy, Tennyson, Jones, McNamara & Douglas
Johnstones’ Duck

Dunstanetta johnstoneorum T. Worthy, Tennyson, Jones, McNamara & Douglas, 2007: Journ. Syst. Palaeont. 5(1): 18, figs 7C, 8C, 9E – St Bathans, Central Otago.

Known from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage from the lower Bannockburn Formation, Manuherikia Group; near St Bathans, Otago (Worthy et al. 2007).

Subfamily TADORNINAE Reichenbach: Shelducks

Tribe TADORNINI Reichenbach: Shelducks

Tadorninae Reichenbach, 1849: Avium Syst. Nat.: 10 – Type genus Tadorna J.D.D. Fleming, 1822.

Genus †Miotadorna T. Worthy, Tennyson, Jones, McNamara & Douglas

Miotadorna T. Worthy, Tennyson, Jones, McNamara & Douglas, 2007: Journ. Syst. Palaeont. 5(1): 14 – Type species (by monotypy) Miotadorna sanctibathansi T. Worthy, Tennyson, Jones, McNamara & Douglas.

➤ †Miotadorna sanctibathansi T. Worthy, Tennyson, Jones, McNamara & Douglas
St Bathans Shelduck

Miotadorna sanctibathansi T. Worthy, Tennyson, Jones, McNamara & Douglas, 2007: Journ. Syst. Palaeont. 5(1): 14, figs 7A, D, 8A, D, 9A, B – St Bathans, Central Otago.

Known from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage from the lower Bannockburn Formation, Manuherikia Group; near St Bathans, Otago (Worthy et al. 2007).

➤ †Miotadorna catrionae Tennyson, Greer, Lubbe, Marx, Richards, Giovanardi & Rawlence
Catrionas Shelduck

Miotadorna catrionae Tennyson, Greer, Lubbe, Marx, Richards, Giovanardi & Rawlence, 2022: Taxonomy 2: 139 – St Bathans, Central Otago.

Known from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage from the lower Bannockburn Formation, Manuherikia Group; near St Bathans, Otago (Tennyson et al. 2022). Worthy, Scofield, Hand et al. (2022) suggested that the referred material was from large males of M. sanctibathansi. It is included here as a distinct species pending further information.

Subfamily ANATINAE Leach: Ducks

Tribe ANATINI Leach: Typical Ducks

Anatidae Leach, 1819: Eleventh room. In Synopsis Contents British Museum 15th edition, London: 67 – Type genus Anas Linnaeus, 1758.

Genus †Matanas T. Worthy, Tennyson, Jones, McNamara & Douglas

Matanas T. Worthy, Tennyson, Jones, McNamara & Douglas, 2007: Journ. Syst. Palaeont. 5(1): 19 – Type species (by monotypy) Matanas enrighti T. Worthy, Tennyson, Jones, McNamara & Douglas.

➤ †Matanas enrighti T. Worthy, Tennyson, Jones, McNamara & Douglas
Enright’s Duck

Matanas enrighti T. Worthy, Tennyson, Jones, McNamara & Douglas, 2007: Journ. Syst. Palaeont. 5(1): 20, figs 4A, C, 5A, C, 6C – St Bathans, Central Otago.

Known from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage from the lower Bannockburn Formation, Manuherikia Group; near St Bathans, Otago (Worthy et al. 2007).

Order †ODONTOPTERYGIFORMES: Pseudotoothed Birds

We follow Bourdon (2005), Bourdon et al. (2010), and G. Mayr et al. (2021) in placing the family Pelagornithidae in its own order, and in using the common name ‘Pseudotoothed Birds’ for the order and family.

Family †PELAGORNITHIDAE Fürbringer: Pseudotoothed Birds

Pelagornithinae Fürbringer, 1888: Untersuch. Morph. Syst. Vögel 2: 1565, footnote – Type genus Pelagornis Lartet, 1857.

Common names are based on Bourdon et al. (2010). Fossil bones from Tangahoe Formation, Hawera, North Island, of middle Pliocene age were described as the first record of pseudodontornids, possibly of Pseudodontornis [now Pelagornis fide G. Mayr & Rubilar-Rogers 2010], from the North Island (McKee 1985). McKee subsequently reported the discovery of isolated pseudodontorn bones of Pliocene age from Hawera and Hawke’s Bay, in the Society of Avian Paleontology and Evolution Newsletter (SAPE) as follows: SAPE 4, 1990; SAPE 5, 1991; SAPE 10, 1996, first Miocene (10–9 Ma) North Island record; SAPE 12, 1998, two pelagornithid humeri; SAPE 13, 1999; SAPE 16, 2002. However, all remain undescribed.

Genus †Protodontopteryx G. Mayr, De Pietri, Love, Mannering & Scofield

Protodontopteryx G. Mayr, De Pietri, Love, Mannering & Scofield, 2021: Papers Palaeont. 7: 218 – Type species (by original designation) Protodontopteryx ruthae G. Mayr, De Pietri, Love, Mannering & Scofield.

➤ †Protodontopteryx ruthae G. Mayr, De Pietri, Love, Mannering & Scofield
Ruth’s Pseudotoothed Bird

Protodontopteryx ruthae G. Mayr, De Pietri, Love, Mannering & Scofield, 2021: Papers Palaeont. 7: 219 – Waipara River, North Canterbury.

Described from a partial skeleton found in an early Paleocene (c. 62–61.5 Ma) deposits at the Waipara River (G. Mayr et al. 2021).

Genus †Pelagornis Lartet

Pelagornis Lartet, 1857: Compt. Rend. Séa. Acad. Sci., Paris 44: 740 – Type species (by monotypy) Pelagornis miocaenus Lartet.

Pseudodontornis Lambrecht, 1930: Geol. Hungarica, Ser. Palaeont. 7: 10 – Type species (by subsequent designation) Odontopteryx longirostris Spulski = Pelagornis longirostris (Spulski).

Osteodontornis Howard, 1957: Santa Barbara Mus. Nat. Hist. Dept. Geol. Bull. 1: 3 – Type species (by original designation) Osteodontornis orri Howard = Pelagornis orri (Howard).

Neodontornis Harrison & Walker, 1976: Tertiary Res. Spec. Pap. 2: 22 – Type species (by original designation) Pseudodontornis stirtoni Howard & Warter = Pelagornis stirtoni (Howard & Warter).

We follow G. Mayr & Rubilar-Rogers (2010) in synonymising Neodontornis with Pelagornis.

➤ †?Pelagornis miocaenus Lartet
Miocene Pseudotoothed Bird

Pelagornis miocaenus Lartet, 1857: Compt. Rend. Séa. Acad. Sci., Paris 44: 740 – France.

Known from the middle Miocene of France (C. Harrison & Walker 1976). In New Zealand: middle to late Miocene, near mouth of Waipara River, North Canterbury (Scarlett 1972). The assignment by Harrison & Walker (1976) of the single proximal humerus described by Scarlett (1972) to Pelagornis miocaenus was tentative.

➤ †Pelagornis stirtoni (Howard & Warter)
Stirton’s Pseudotoothed Bird

Pseudodontornis stirtoni Howard & Warter, 1969: Rec. Cant. Museum 8: 348 – Motunau Beach, North Canterbury.

Neodontornis stirtoni (Howard & Warter); C. Harrison & Walker 1976: Tertiary Res. Spec. Pap. 2: 22.

Pelagornis stirtoni (Howard & Warter); G. Mayr & Rubilar-Rogers 2010: Journ. Vert. Paleont. 30: 1327.

Age uncertain, within the range Otaian to Waipipian Stage (early Miocene–Pliocene; 21.7–3.0 Ma), probably late Pliocene; Motunau Beach, North Canterbury (H. Howard & Warter 1969).

Order PHOENICOPTERIFORMES: Flamingoes and Palaelodids

Family †PALAELODIDAE Stejneger: Palaelodids

Genus †Palaelodus Milne-Edwards

Palaelodus Milne-Edwards, 1863: Ann. Sci. Nat. 4: 157, 158 – Type species (by subsequent designation) Palaelodus ambiguous Milne-Edwards.

➤ †Palaelodus aotearoa T. Worthy, Tennyson, Archer & Scofield
New Zealand Palaelodus

Palaelodus aotearoa T. Worthy, Tennyson, Archer & Scofield, 2010: Rec. Aust. Mus. 62: 80 – St Bathans, Central Otago.

Known only from fossil remains from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage, Otago (Worthy, Tennyson et al. 2010).

Order COLUMBIFORMES: Pigeons and Doves

Suborder COLUMBAE: Pigeons and Doves

Family COLUMBIDAE Illiger: Pigeons and Doves

Columbini Illiger, 1811: Prodromus Syst. Mamm. Avium: 243 – Type genus Columba Linnaeus, 1758.

Subfamily RAPHINAE Wetmore: Fruit Doves

Raphidae Wetmore, 1930: Proc. U.S. Nat. Mus 76(24): 5 – Type genus Raphus Brisson, 1760.

Genus †Deliaphaps De Pietri, Scofield, Tennyson, Hand & T. Worthy

Deliaphaps De Pietri, Scofield, Tennyson, Hand & T. Worthy, 2017: Paleontología y Evolución de las Aves: 57 – Type species (by original designation) Deliaphaps zealandiensis De Pietri, Scofield, Tennyson, Hand & T. Worthy.

➤ †Deliaphaps zealandiensis De Pietri, Scofield, Tennyson, Hand & T. Worthy
Zealandian Dove

Deliaphaps zealandiensis De Pietri, Scofield, Tennyson, Hand & T. Worthy, 2017: Paleontología y Evolución de las Aves: 57 – Manuherikia River, Otago.

Known only from a carpometacarpus and two tentatively referred bones from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage, Otago (De Pietri et al. 2017). Deliaphaps displays a combination of features also present in the extant genera Caloenas, Didunculus, and Goura (De Pietri et al. 2017). Based on placement of these genera within Raphinae (Pereira et al. 2007; Dickinson & Remsen 2013; Nowak et al. 2019), we include Deliaphaps in Raphinae.

Genus †Rupephaps T. Worthy, Hand, J. Worthy, Tennyson & Scofield

Rupephaps T. Worthy, Hand, J. Worthy, Tennyson & Scofield, 2009: Auk 126: 651 – Type species (by original designation) Rupephaps taketake T. Worthy, Hand, J. Worthy, Tennyson & Scofield.

➤ †Rupephaps taketake T. Worthy, Hand, J. Worthy, Tennyson & Scofield
St Bathans Pigeon

Rupephaps taketake T. Worthy, Hand, J. Worthy, Tennyson & Scofield, 2009: Auk 126: 651 – St Bathans, Central Otago.

Known from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage from the lower Bannockburn Formation, Manuherikia Group; near St Bathans, Otago (Worthy et al. 2009). De Pietri et al. (2018) considered Rupephaps to be part of a clade including Ptilinopus and Hemiphaga.

Order APODIFORMES: Swifts, Hummingbirds, and Owlet-nightjars

Family AEGOTHELIDAE Bonaparte: Owlet-nightjars

Aegothelinae Bonaparte, 1853: Compt. Rend. Séa. Acad. Sci., Paris 37(18): 645 – Type genus Aegotheles Vigors & Horsfield, 1827.

Genus Aegotheles Vigors & Horsfield

Aegotheles Vigors & Horsfield, 1826: Trans. Linn. Soc. London 15(1): 194 – Type species (by monotypy) Caprimulgus novaehollandiae Latham = Aegotheles cristatus (White).

➤ †Aegotheles zealandivetus T. Worthy, Scofield, Salisbury, Hand, De Pietri & Archer
St Bathans Owlet-nightjar

Aegotheles zealandivetus T. Worthy, Scofield, Salisbury, Hand, De Pietri & Archer, 2022: Journ. Ornith. 163: 646 – Manuherikia River, St Bathans, Otago.

Worthy et al. (2007) referred a specimen from the St Bathans assemblage to Aegotheles sp. but did not name it. The species was named after further referable material was found (Worthy, Salisbury, Hand et al. 2022b). From the Altonian Stage (early Miocene; 19–16 Ma), lower Bannockburn Formation, Manuherikia Group; near St Bathans, Otago.

Family APODIDAE Olphe-Galliard: Swifts and Swiftlets

Apodidae Olphe-Galliard, 1887: Contrib. Faune Ornith. Europe Occidentale 22: 90 – Type genus Apus Scopoli 1777.

Genus Collocalia G.R. Gray

Collocalia G.R. Gray, 1840: List Gen. Birds: 8 – Type species (by original designation) Hirundo esculenta Linnaeus = Collocalia esculenta (Linnaeus).

➤ Collocalia sp. indet.
(swiftlet)

Collocalia sp. indet. T. Worthy, Tennyson, Jones, McNamara & Douglas, 2007: Journ. Syst. Palaeont. 5(1): 26, fig. 16A.

Worthy et al. (2007) referred a specimen from the St Bathans assemblage to Collocalia sp. but did not name it. From the Altonian Stage (early Miocene; 19–16 Ma), lower Bannockburn Formation, Manuherikia Group; near St Bathans, Otago.

Order GRUIFORMES: Rails and Cranes

Family †APTORNITHIDAE Bonaparte: Adzebills

Genus †Aptornis G.A. Mantell

Aptornis G.A. Mantell, 1848 (2 Feb.): Quart. Journ. Geol. Soc. London 4: 233 – Type species (by monotypy) Dinornis otidiformis Owen = Aptornis otidiformis (Owen).

➤ †?Aptornis proasciarostratus T. Worthy, Tennyson & Scofield
St Bathans Adzebill

?Aptornis proasciarostratus T. Worthy, Tennyson & Scofield, 2011: Journ. Ornith. 152: 671 – St Bathans, Central Otago.

Known only from fossil remains from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage, Otago (Worthy et al. 2011a).

Family RALLIDAE Rafinesque: Rails, Gallinules, and Coots

Subfamily RALLINAE Rafinesque: Rails, Gallinules, and Coots

Rallia Rafinesque, 1815: Analyse de la Nature: 70 – Type genus Rallus Linnaeus, 1758.

Genus †Priscaweka Mather, Tennyson, Scofield, De Pietri, Hand, Archer, Handley & T. Worthy

Priscaweka Mather, Tennyson, Scofield, De Pietri, Hand, Archer, Handley & T. Worthy, 2018 [2019]: Journ. Syst. Palaeont. 17: 432 – Type species (by original designation) Priscaweka parvales Mather, Tennyson, Scofield, De Pietri, Hand, Archer, Handley & T. Worthy.

➤ †Priscaweka parvales Mather, Tennyson, Scofield, De Pietri, Hand, Archer, Handley & T. Worthy
Bannockburn Crake

Priscaweka parvales Mather, Tennyson, Scofield, De Pietri, Hand, Archer, Handley & T. Worthy, 2018 [2019]: Journ. Syst. Palaeont. 17: 438 – St Bathans, Central Otago.

Known from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage from the lower Bannockburn Formation, Manuherikia Group; near St Bathans, Otago (Mather et al. 2018). Phylogenetic relationships uncertain, but many features shared with Gallirallus and related genera (Mather et al. 2018).

Genus †Litorallus Mather, Tennyson, Scofield, De Pietri, Hand, Archer, Handley & T. Worthy

Litorallus Mather, Tennyson, Scofield, De Pietri, Hand, Archer, Handley & T. Worthy, 2018 [2019]: Journ. Syst. Palaeont. 17: 441 – Type species (by original designation) Litorallus livezeyi Mather, Tennyson, Scofield, De Pietri, Hand, Archer, Handley & T. Worthy.

➤ †Litorallus livezeyi Mather, Tennyson, Scofield, De Pietri, Hand, Archer, Handley & T. Worthy
Livezey’s Rail

Litorallus livezeyi Mather, Tennyson, Scofield, De Pietri, Hand, Archer, Handley & T. Worthy, 2018 [2019]: Journ. Syst. Palaeont. 17: 441 – St Bathans, Central Otago.

Known from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage from the lower Bannockburn Formation, Manuherikia Group; near St Bathans, Otago (Mather et al. 2018). Phylogenetic relationships uncertain, but likely from the same lineage as Priscaweka (Mather et al. 2018).

Genus †Pleistorallus T. Worthy

Pleistorallus T. Worthy, 1997: Alcheringa 21: 74 – Type species (by original designation) Pleistorallus flemingi T. Worthy.

➤ †Pleistorallus flemingi T. Worthy
Fleming’s Rail

Pleistorallus flemingi T. Worthy, 1997: Alcheringa 21: 74 – Marton, Manawatu.

Known from the Kaimatira Pumice Sand of the Kai Iwi Group, Lower Castlecliffian, Oxygen Isotope Stage 25–27, c. 1 Ma, near Marton. Known from the holotype tibiotarsus and paratype femur. Relationships to Recent rails unresolved, but most like Gallirallus group (Worthy 1997a).

Order CHARADRIIFORMES: Waders, Gulls, & Terns

Suborder CHARADRII

Superfamily CHIONOIDEA Lesson

Family INCERTAE SEDIS

Genus †Neilus De Pietri, Scofield, Hand, Tennyson & T. Worthy

Neilus De Pietri, Scofield, Hand, Tennyson & T. Worthy, 2016: Journ. Roy. Soc. N.Z. 46: 186 – Type species (by original designation) Neilus sansomae De Pietri, Scofield, Hand, Tennyson & T. Worthy.

➤ †Neilus sansomae De Pietri, Scofield, Hand, Tennyson & T. Worthy
Sansom’s Plover

Neilus sansomae De Pietri, Scofield, Hand, Tennyson & T. Worthy, 2016: Journ. Roy. Soc. N.Z. 46: 186 – St Bathans, Central Otago.

Known only from fossil remains from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage, Otago (De Pietri, Scofield, Hand et al. 2016).

Suborder SCOLOPACI

Superfamily THINOCOROIDEA Sundevall

Family INCERTAE SEDIS

Genus †Hakawai De Pietri, Scofield, Tennyson, Hand & T. Worthy

Hakawai De Pietri, Scofield, Tennyson, Hand & T. Worthy, 2015 [2016]: Journ. Syst. Palaeont. 14: 605 – Type species (by original designation) Hakawai melvillei De Pietri, Scofield, Tennyson, Hand & T. Worthy.

➤ †Hakawai melvillei De Pietri, Scofield, Tennyson, Hand & T. Worthy
New Zealand Lake-wanderer

Hakawai melvillei De Pietri, Scofield, Tennyson, Hand & T. Worthy, 2015 [2016]: Journ. Syst. Palaeont. 14: 605 – St Bathans, Central Otago.

Known only from fossil remains from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage, Otago (De Pietri, Scofield, Tennyson et al. 2015 [2016]).

Order PHAETHONTIFORMES: Tropicbirds

Family INCERTAE SEDIS: Fossil tropicbirds

In addition to Clymenoptilon novaezealandicum listed below, G. Mayr & Scofield (2015) and G. Mayr et al. (2023) referred to a smaller, as yet undescribed, fossil tropicbird specimen from the Waipara Greensand, Waipara River, Canterbury.

Genus †Clymenoptilon G. Mayr, De Pietri, Love, Mannering, Crouch, Reid & Scofield

Clymenoptilon G. Mayr, De Pietri, Love, Mannering, Crouch, Reid & Scofield, 2023: Alcheringa 47: 316 – Type species (by original designation) Clymenoptilon novaezealandicum Mayr, De Pietri, Love, Mannering, Crouch, Reid & Scofield.

➤ †Clymenoptilon novaezealandicum G. Mayr, De Pietri, Love, Mannering, Crouch, Reid & Scofield
Zealandian Tropicbird

Clymenoptilon novaezealandicum G. Mayr, De Pietri, Love, Mannering, Crouch, Reid & Scofield, 2023: Alcheringa 47: 316 – Waipara River, Canterbury.

Described from a partial skeleton including the skull, vertebral column, right wing, pectoral girdle elements, and pelvis from the lower part of the Stormont Member, Waipara Greensand (late early Paleocene to early late Paleaocene, c. 62–58 Ma), in the Waipara River valley, Canterbury (G. Mayr et al. 2023).

Order SPHENISCIFORMES: Penguins

We follow J. Clarke et al. (2003) and Ksepka et al. (2006, 2012) in restricting the family Spheniscidae to the common ancestor of living penguin taxa plus Marplesornis, and the descendants of this common ancestor. Several subfamilies have been used in the past for fossil penguins (e.g. Simpson 1971, 1975). If families are intended to be monophyletic, at least 12 new families would be required to contain the known diversity among fossil penguins, based on phylogenetic diagrams presented by Ksepka et al. (2012) and G. Mayr et al. (2018). We have therefore placed all fossil penguin taxa, other than Marplesornis and modern genera, in Families incertae sedis.

New Zealand has a rich record of fossil penguins (24 named species in 20 genera) extending from the Paleocene through to the Holocene, as reviewed by Fordyce (1991a,b) and Fordyce & Jones (1990). Rich records of fossil penguins are also known from the Eocene of Seymour Island, Antarctica (e.g. Myrcha et al. 2002 and references therein); Patagonia in Argentina (Simpson 1972a, 1981; Cione & Tonni 1981; Cozzuol et al. 1993; Acosta Hospitaleche et al. 2004); Chile (Walsh & Hume 2001); and Perú (Noriega & Tambussi 1989; Stucchi 2002; Stucchi et al. 2003; Acosta Hospitaleche & Stucchi 2005). To date, no species or genera overlap in ranges between New Zealand and South America (Acosta Hospitaleche et al. 2004) and there is only limited co-occurrence of genera in Seymour Island, Australia and New Zealand (Fordyce 1991a,b; Myrcha et al. 2002). Common names of fossil species follow Checklist Committee (2010), apart from the more recently named Rosie’s penguin, Stilwell’s penguin, Waipara Crossvallia penguin, Bice’s penguin, Maxwell’s penguin, Waitaki penguin, Grebneff’s penguin, and dawn crested penguin, plus Huxley’s penguin used here for the first time.

Families INCERTAE SEDIS: Fossil penguins

Genus †Waimanu Jones, Ando & Fordyce

Waimanu Jones, Ando & Fordyce, 2006: in Slack et al., Molec. Biol. Evolution 23(6): 1145 – Type species (by original designation) Waimanu manneringi Jones, Ando & Fordyce.

➤ †Waimanu manneringi Jones, Ando & Fordyce
Mannering’s Penguin

Waimanu manneringi Jones, Ando & Fordyce, 2006: in Slack et al., Molec. Biol. Evolution 23(6): 1145 – Waipara River, North Canterbury.

Known from the Teurian Stage (late early Palaeocene; 61.6–60.5 Ma), basal Waipara Greensand, Waipara River, Canterbury (Slack et al. 2006).

Genus †Muriwaimanu G. Mayr, De Pietri, Love, Mannering & Scofield

Muriwaimanu G. Mayr, De Pietri, Love, Mannering & Scofield, 2018: Journ. Vert. Paleont. 37(6: e1398169-3): 3 – Type species (by original designation) Waimanu tuatahi Ando, Jones & Fordyce.

➤ †Muriwaimanu tuatahi (Ando, Jones & Fordyce)
Waipara Penguin

Waimanu tuatahi Ando, Jones & Fordyce, 2006: in Slack et al., Molec. Biol. Evolution 23(6): 1146 – Waipara River, North Canterbury.

Muriwaimanu tuatahi (Ando, Jones & Fordyce); G. Mayr et al. 2018, Journ. Vert. Paleont. 37(6: e1398169-3): 3.

Known from the Teurian Stage (late early Palaeocene; 60–58 Ma), middle-upper Waipara Greensand, Waipara River, Canterbury (Slack et al. 2006). Originally named as Waimanu tuatahi, this fossil species was considered to have sufficiently distinct tarsometatarsus morphology to be placed in its own genus by G. Mayr et al. (2018).

Genus †Sequiwaimanu G. Mayr, De Pietri, Love, Mannering & Scofield

Sequiwaimanu G. Mayr, De Pietri, Love, Mannering & Scofield, 2018: Journ. Vert. Paleont. 37(6: e1398169-3): 3 – Type species (by original designation) Sequiwaimanu rosieae G. Mayr, De Pietri, Love, Mannering & Scofield.

➤ †Sequiwaimanu rosieae G. Mayr, De Pietri, Love, Mannering & Scofield
Rosie’s Penguin

Sequiwaimanu rosieae G. Mayr, De Pietri, Love, Mannering & Scofield, 2018: Journ. Vert. Paleont. 37(6: e1398169-3): 4 – Waipara River, North Canterbury.

Described from a partial skeleton found in a late Teurian stage (late early Paleocene) deposit (61 Ma) at the Waipara River (G. Mayr et al. 2018).

Genus †Kupoupou Blokland, Reid, T. Worthy, Tennyson, Clarke & Scofield

Kupoupou Blokland, Reid, T. Worthy, Tennyson, Clarke & Scofield, 2019: Palaeontologia Electronica 22.3.78: 8 – Type species (by original designation) Kupoupou stilwelli Blokland, Reid, T. Worthy, Tennyson, Clarke & Scofield.

➤ †Kupoupou stilwelli Blokland, Reid, T. Worthy, Tennyson, Clarke & Scofield
Stilwell’s Penguin

Kupoupou stilwelli Blokland, Reid, T. Worthy, Tennyson, Clarke & Scofield, 2019: Palaeontologia Electronica 22.3.78: 16 – Maunganui Beach, Chatham Island.

Described by Blokland et al. (2019) from several fossils found in late early-middle Paleocene (62.5–60 Ma) deposits east of Tahatika Creek, north-western Chatham Island.

Genus †Crossvallia Tambussi, Reguero, Marenssi & Santillana

Crossvallia Tambussi, Reguero, Marenssi & Santillana, 2005: Geobios 38: 669 – Type species (by original designation) Crossvallia unienwillia Tambussi, Reguero, Marenssi & Santillana.

One species known from Seymour Island, Antarctica (Tambussi et al. 2005) and another New Zealand specimen tentatively referred to this genus.

➤ †?Crossvallia waiparensis G. Mayr, De Pietri, Love, Mannering & Scofield
Waipara Crossvallia Penguin

?Crossvallia waiparensis G. Mayr, De Pietri, Love, Mannering & Scofield, 2020: Alcheringa 44: 195 – Waipara River, North Canterbury.

Described from a fossil found in a Teurian Paleocene deposit (62–58 Ma), Waipara River, Canterbury (G. Mayr et al. 2020).

Genus †Kumimanu G. Mayr, Scofield, De Pietri & Tennyson

Kumimanu G. Mayr, Scofield, De Pietri & Tennyson, 2017: Nature Communications 8: 1927 (doi.org/10.1038/s41467-017-01959-6) 3 – Type species (by original designation) Kumimanu biceae G. Mayr, Scofield, De Pietri & Tennyson.

➤ †Kumimanu biceae G. Mayr, Scofield, De Pietri & Tennyson
Bice’s Penguin

Kumimanu biceae G. Mayr, Scofield, De Pietri & Tennyson, 2017: Nature Communications 8: 1927 (doi.org/10.1038/s41467-017-01959-6) 3 – Hampden Beach, Otago.

Described from a partial skeleton from the late Paleocene Moeraki Formation (60–55 Ma) near Oamaru (G. Mayr et al. 2017).

➤ †Kumimanu fordycei Ksepka, Field, Heath, Pett, Thomas, Giovanardi & Tennyson
Fordyce’s Penguin

Kumimanu fordycei Ksepka, Field, Heath, Pett, Thomas, Giovanardi & Tennyson, 2023: Journal of Paleontology 2022.88: 3 – Hampden Beach, Otago.

Described from a partial skeleton from the late Paleocene Moeraki Formation (59.5–55.5 Ma) near Oamaru (Ksepka, Field et al. 2023).

Genus †Petradyptes Ksepka, Field, Heath, Pett, Thomas, Giovanardi & Tennyson

Petradyptes Ksepka, Field, Heath, Pett, Thomas, Giovanardi & Tennyson, 2023: Journal of Paleontology 2022.88: 7 – Type species (by original designation) Petradyptes stonehousei Ksepka, Field, Heath, Pett, Thomas, Giovanardi & Tennyson.

➤ †Petradyptes stonehousei Ksepka, Field, Heath, Pett, Thomas, Giovanardi & Tennyson
Stonehouse’s Penguin

Petradyptes stonehousei Ksepka, Field, Heath, Pett, Thomas, Giovanardi & Tennyson, 2023: Journal of Paleontology 2022.88: 7 – Hampden Beach, Otago.

Described from a humerus and part femur (plus three other referred specimens containing multiple skeletal elements) from the late Paleocene Moeraki Formation (59.5–55.5 Ma) near Oamaru (Ksepka, Field et al. 2023).

Genus †Kaiika Fordyce & Thomas

Kaiika Fordyce & Thomas, 2011: New Zealand Journ. Geol. Geophys. 54: 45 – Type species (by original designation) Kaiika maxwelli Fordyce & Thomas.

➤ †Kaiika maxwelli Fordyce & Thomas
Maxwell’s Penguin

Kaiika maxwelli Fordyce & Thomas, 2011: New Zealand Journ. Geol. Geophys. 54: 45 – Waihao River, South Canterbury.

Known from a single fossil humerus cast from the early Eocene (c. 54 Ma), Waihao River, South Canterbury (Fordyce & Thomas 2011).

Genus †Palaeeudyptes Huxley

Palaeeudyptes Huxley, 1859: Quart. Journ. Geol. Soc. London 15: 675 – Type species (by monotypy) Palaeeudyptes antarcticus Huxley.

Eosphaeniscus Wiman, 1905: Bull. Geological Inst. Uppsala 6: 250 – Type species (by monotypy) Eosphaeniscus Gunnari Wiman = Palaeeudyptes gunnari (Wiman).

Several indeterminate specimens from New Zealand, Australia, southern Chile, and Seymour Island (Antarctic Peninsula) have been referred to the genus (Hector 1872; Marples 1952, 1962; Simpson 1971, 1975; Fordyce 1991a,b; Sallaberry et al. 2010; Ksepka et al. 2012; T. Park & Fitzgerald 2012). Two extralimital fossil species, Palaeeudyptes klekowskii Myrcha, Tatur & del Valle, 1990 and P. gunnari (Wiman, 1905) have been named from the Eocene La Meseta formation of Seymour Island, Antarctica (Myrcha et al. 2002).

➤ †Palaeeudyptes antarcticus Huxley
Huxley’s Penguin

Palaeeudyptes antarcticus Huxley, 1859: Quart. Journ. Geol. Soc. London 15: 675 – Kakanui, near Oamaru.

Huxley (1859) named Palaeeudyptes antarcticus from a tarsometatarsus (BMNH A1048) from Kakanui, North Otago. Oliver (1930) followed Hector (1872) in referring a second specimen (NMNZ S.1449, which includes a humerus) from Seal Rock (near Punakaiki) to the same taxon, for which Oliver coined the name ‘Narrow-flippered Penguin’. This nomenclature was followed by Checklist Committee (2010). Ksepka et al. (2012) considered that NMNZ S.1449 could not be assigned to a currently recognised genus, and placed it in “Sphenisciformes indet.”. Further, Ksepka et al. (2012) accepted two specimens only within a much constrained Palaeeudyptes antarcticus: the holotype, and a second tarsometatarsus from Burnside Quarry, Dunedin (OM GL430 (C47:17)). Therefore “Narrow-flippered penguin” is not an appropriate name for a taxon known from leg elements only. A name derived from Huxley’s specific epithet antarcticus is also not appropriate for a penguin known from the Otago Province only. We consider that a more appropriate name for P. antarcticus is Huxley’s penguin, in recognition of Thomas Huxley’s contribution in recognising and naming this, the first named fossil penguin.

Age uncertain within the range Whaingaroan Stage (early Oligocene; 34.3 Ma) to Waitakian Stage (late Oligocene–early Miocene; 21.7 Ma) (Simpson 1971; Fordyce & Jones 1990; Fordyce 1991a,b). Records of this taxon from Australia (Simpson 1957; Brodkorb 1963) are now referred to Palaeeudyptes sp. indet. (Simpson 1971; R. Jenkins 1974). The referral of humeri from Seymour Island to this species (Tambussi et al. 2006) has been challenged by Jadwiszczak (2006b).

➤ †Palaeeudyptes marplesi Brodkorb
Marples’ Penguin

Palaeeudyptes cf. antarcticus Huxley, 1859; Marples 1952, New Zealand Geol. Surv. Pal. Bull. 20: 31.

Palaeeudyptes marplesi Brodkorb, 1963: Bull. Florida State Museum (Biol. Sci.) 7: 231 – Burnside, near Dunedin, Otago.

Known from a Kaiatan or Runangan Stage (late Eocene; 37.0–34.3 Ma) deposit at Burnside, Otago (Brodkorb 1963; Simpson 1971, 1975; Fordyce 1991a,b). A specimen from the Upper Eocene, Blanche Point Marls, Christie’s Beach, Adelaide, South Australia (South Australian Museum P.10870; Simpson 1957), was referred first to this taxon (Brodkorb 1963) but later to Palaeeudyptes sp. indet. (Simpson 1971).

Genus †Pachydyptes Oliver

Pachydyptes Oliver, 1930: New Zealand Birds, 1st edition: 85 – Type species (by original designation) Pachydyptes ponderosus Oliver.

Pachydyptes simpsoni Jenkins, 1974 from Australia was regarded as conspecific with Anthropornis nordenskjoeldii Wiman, 1905 from Seymour Island (Fordyce 1991a; Vickers-Rich 1991). However, Simpson (1971: 347) considered Anthropornis Wiman, 1905 to be tentatively distinct from Pachydyptes. More recently, Ksepka & Clarke (2010) and T. Park & Fitzgerald (2012) concluded the generic position of Pachydyptes simpsoni to be uncertain.

➤ †Pachydyptes ponderosus Oliver
New Zealand Giant Penguin

Palaeeudyptes antarcticus; Hector 1873, Trans. Proc. N.Z. Inst. 5: 438. Not Palaeeudyptes antarcticus Huxley, 1859.

Pachydyptes ponderosus Oliver, 1930: New Zealand Birds, 1st edition: 86 – Fortification Hill, Oamaru.

Pachydyptes ponderosa Oliver; Lowe 1939, Ibis 3 (14th series): 282. Unjustified emendation.

Anthropornis ponderosus (Oliver); Lowe 1939, Ibis 3 (14th series): 291.

Anthropornis ponderosa (Oliver); Lowe 1939, Ibis 3 (14th series): 291. Unjustified emendation.

Anthropornis (Pachydyptes) ponderosus (Oliver); Lowe 1939, Ibis 3 (14th series): 292.

Anthropornis nordenskjöldii; Lowe 1939, Ibis 3 (14th series): 293. Not Anthropornis nordenskjoeldii Wiman, 1905.

Runangan Stage (late Eocene; 36.0–34.3 Ma); near Oamaru, Otago (Marples 1952; Brodkorb 1963; Simpson 1971, 1975; Fordyce 1991a,b).

Genus †Kairuku Ksepka, Fordyce, Ando & Jones

Kairuku Ksepka, Fordyce, Ando & Jones, 2012: Journ. Vert. Paleont. 32: 239 – Type species (by original designation) Kairuku waitaki Ksepka, Fordyce, Ando & Jones.

➤ †Kairuku waewaeroa Giovanardi, Ksepka & Thomas
Junats’ Penguin

Kairuku waewaeroa Giovanardi, Ksepka & Thomas, 2021: Journ. Vert. Paleont. 41: 4 – Kawhia Harbour.

Described from a single skeleton from Kawhia Harbour, from the Oligocene, 35–27 Ma (Giovanardi et al. 2021). Junats is a constriction of the Hamilton Junior Naturalist Club, who were involved in the discovery of the holotype; Junats’ penguin should have an initial capital.

➤ †Kairuku waitaki Ksepka, Fordyce, Ando & Jones
Waitaki Penguin

Kairuku waitaki Ksepka, Fordyce, Ando & Jones, 2012: Journ. Vert. Paleont. 32: 239 – Waihao River, South Canterbury.

Described from a single skeleton from the Waihao River, South Canterbury, from the late Oligocene, 27–26 Ma (Ksepka et al. 2012).

➤ †Kairuku grebneffi Ksepka, Fordyce, Ando & Jones
Grebneff’s Penguin

Kairuku grebneffi Ksepka, Fordyce, Ando & Jones, 2012: Journ. Vert. Paleont. 32: 245 – Waipati catchment, North Otago.

Known from two fossils from the late Oligocene, 27–26 Ma, from the Waipati catchment of the Maerewhenua River near Duntroon, North Otago, and a specimen from the bank of the Waihao River, South Canterbury (Ksepka et al. 2012).

Genus †Platydyptes Marples

Platydyptes Marples, 1952: New Zealand Geol. Surv. Pal. Bull. 20: 37 – Type species (by original designation) Pachydyptes novaezealandiae Oliver = Platydyptes novaezealandiae (Oliver).

➤ †Platydyptes marplesi Simpson
Simpson’s Penguin

Platydyptes novaezealandiae (Oliver, 1930); Marples 1952, New Zealand Geol. Surv. Pal. Bull. 20: 38. In part.

? Platydyptes marplesi Simpson, 1971: Bull. Am. Mus. Nat. Hist. 144 (5): 354 – Waitaki Valley, north Otago.

? Platydyptes marplesigi; Simpson 1975, in Stonehouse (Ed.) Biology of Penguins: 31. Unjustified emendation.

“Platydyptes” marplesi Simpson; Checklist Committee 1990, Checklist Birds N.Z.: 64.

Known from a Duntroonian Stage (late Oligocene; 27.3–25.2 Ma) deposit; probably Wharekuri, Otago (Simpson 1971; Fordyce 1991a,b). Simpson (1971) assigned this species to Platydyptes but considered this assignation “quite doubtful”.

➤ †Platydyptes novaezealandiae (Oliver)
Wide-flippered Penguin

Pachydyptes novaezealandiae Oliver, 1930: New Zealand Birds, 1st edition: 86 – Oamaru district.

Pachydyptes novae-zelandiae Oliver; Lowe 1939, Ibis 3 (14th series): 282. Unjustified emendation.

Platydyptes novaezealandiae (Oliver); Marples 1952, New Zealand Geol. Surv. Pal. Bull. 20: 38. In part.

Known from a Duntroonian to Waitakian Stage (late Oligocene–early Miocene; 27.3–21.7 Ma) deposit; Oamaru, Otago (Simpson 1971, Fordyce 1991b).

➤ †Platydyptes amiesi Marples
Amies’ Penguin

Platydyptes amiesi Marples, 1952: New Zealand Geol. Surv. Pal. Bull. 20: 39 – Hakataramea Valley, South Canterbury.

Known from Duntroonian to Waitakian Stage (late Oligocene–early Miocene; 27.3–21.7 Ma) deposits; Hakataramea (Canterbury), and Duntroon (Otago) (Marples 1952; Fordyce 1991a,b). Known from many specimens (Fordyce 2003).

Genus †Archaeospheniscus Marples

Archaeospheniscus Marples, 1952: New Zealand Geol. Surv. Pal. Bull. 20: 40 – Type species (by original designation) Archaeospheniscus lowei Marples.

Notodyptes Marples, 1953: Falkland Islands Dependencies Survey Scientific Reports 5: 11 – Type species (by original designation) Notodyptes wimani Marples = Archaeospheniscus wimani (Marples).

An extralimital fossil species, Archaeospheniscus wimani (Marples, 1953), is known from the Eocene in La Meseta Formation of Seymour Island, Antarctica (Myrcha et al. 2002; Jadwiszczak 2006a).

➤ †Archaeospheniscus lowei Marples
Lowe’s Penguin

Archaeospheniscus lowei Marples, 1952: New Zealand Geol. Surv. Pal. Bull. 20: 41 – Duntroon, North Otago.

Known from Duntroonian Stage (late Oligocene; 27.3–25.2 Ma) deposits; Duntroon, Otago (Marples 1952; Fordyce 1991a,b).

➤ †Archaeospheniscus lopdellorum Marples
Lopdells’ Penguin

Archaeospheniscus lopdelli Marples, 1952: New Zealand Geol. Surv. Pal. Bull. 20: 41 – Duntroon, North Otago.

Archaeospheniscus lopdellorum Marples, 1952; Tennyson et al. 2010, Notornis 57: 54. Emendation.

Known from Duntroonian Stage (late Oligocene; 27.3–25.2 Ma) deposits; Duntroon, Otago (Marples 1952; Fordyce 1991a,b). The referral of humeri from Seymour Island, Antarctica, to this species (Tambussi et al. 2006) has been challenged by Jadwiszczak (2006b).

The specific name should be A. lopdellorum (Lopdells’ penguin) rather than A. lopdelli (Lopdell’s penguin) (see Tennyson et al. 2010).

Genus †Duntroonornis Marples

Duntroonornis Marples, 1952: New Zealand Geol. Surv. Pal. Bull. 20: 42 – Type species (by original designation) Duntroonornis parvus Marples.

➤ †Duntroonornis parvus Marples
Duntroon Penguin

Duntroonornis parvus Marples, 1952: New Zealand Geol. Surv. Pal. Bull. 20: 42 – Duntroon, North Otago.

Known from Duntroonian Stage (late Oligocene; 27.3–25.2 Ma) deposits; Duntroon, Otago (Marples 1952; Fordyce 1991a,b). Possibly also Waitakian Stage (late Oligocene–early Miocene; 25.2–21.7 Ma) deposits at Hakataramea Valley, South Canterbury (Fordyce & Jones 1990).

Genus †Korora Marples

Korora Marples, 1952: New Zealand Geol. Surv. Pal. Bull. 20: 43 – Type species (by original designation) Korora oliveri Marples.

➤ †Korora oliveri Marples
Oliver’s Penguin

Korora oliveri Marples, 1952: New Zealand Geol. Surv. Pal. Bull. 20: 43 – Hakataramea Valley, South Canterbury.

Known from Waitakian Stage (late Oligocene–early Miocene; 25.2–21.7 Ma) deposits; Hakataramea Valley, South Island (Marples 1952; Fordyce & Jones 1990; Fordyce 1991a,b).

Family SPHENISCIDAE Bonaparte: Penguins

Spheniscidae Bonaparte, 1831: Giornale Arcadico di Scienze, Lettere ed Arti 49: 62 – Type genus Spheniscus Brisson, 1760.

We follow J. Clarke et al. (2003) and Ksepka et al. (2006, 2012) in restricting the family Spheniscidae to the common ancestor of living penguin taxa plus Marplesornis, and the descendants of this common ancestor.

Scarlett (1984) proposed the name Tereingaornis moisleyi for Waipipian Stage (Pliocene; 3.6–3.0 Ma), Hawke’s Bay penguin fossils that he considered to be closely related to, if not congeneric with, Spheniscus. McKee (1988) referred two Taranaki fossils to the same genus and species. However, Thomas, Ksepka et al. (2020) showed that fossils referred to Tereingaornis could not be distinguished from either Eudyptes or Megadyptes, and so we regard both Tereingaornis and T. moisleyi as nomina dubia.

Genus †Marplesornis Simpson

Marplesornis Simpson, 1972: Rec. Cant. Museum 9: 162 – Type species (by original designation) Palaeospheniscus novaezealandiae Marples = Marplesornis novaezealandiae (Marples).

➤ †Marplesornis novaezealandiae (Marples)
Harris’ Penguin

Palaeospheniscus novaezealandiae Marples, 1960: Rec. Cant. Museum 7: 194 – Motunau Beach, North Canterbury.

Marplesornis novaezealandiae (Marples); Simpson 1972, Rec. Cant. Museum 9: 162.

Age uncertain, within the range Otaian to Waipipian Stage (early Miocene–Pliocene; 21.7–3.0 Ma), probably late Pliocene; Motunau, North Canterbury (Simpson 1972b; Fordyce 1991a,b).

Genus Aptenodytes J.F. Miller

Aptenodytes J.F. Miller, 1778: Icones Animalium 4: pl. 23 – Type species (by monotypy) Aptenodytes patagonicus J.F. Miller.

➤ †Aptenodytes ridgeni Simpson
Ridgen’s Penguin

Aptenodytes ridgeni Simpson, 1972: Rec. Cant. Museum 9: 167 – Motunau Beach, North Canterbury.

Age uncertain within the range Otaian to Waipipian Stage (early Miocene–Pliocene; 21.7–3.0 Ma), probably late Pliocene; Motunau, North Canterbury (Simpson 1972b; Fordyce 1991a,b).

Genus Pygoscelis Wagler

Pygoscelis Wagler, 1832: Isis von Oken, Heft 2: col. 281 – Type species (by monotypy) Aptenodytes papua J.R. Forster = Pygoscelis papua (J.R. Forster).

➤ †Pygoscelis tyreei Simpson
Tyree’s Penguin

Pygoscelis tyreei Simpson, 1972: Rec. Cant. Museum 9: 166 – Motunau Beach, North Canterbury.

Age uncertain within the range Otaian to Waipipian Stage (early Miocene–Pliocene; 21.7–3.0 Ma), probably late Pliocene; Motunau, North Canterbury (Simpson 1972b; Fordyce 1991a,b).

Genus Eudyptes Vieillot

Eudyptes Vieillot, 1816 (Apr.): Analyse Nouv. Ornith. Elem.: 67, 70 – Type species (by subsequent designation) Aptenodytes chrysocome J.R. Forster = Eudyptes chrysocome (J.R. Forster).

➤ †Eudyptes atatu Thomas, Tennyson, Scofield & Ksepka
Dawn Crested Penguin

Eudyptes atatu Thomas, Tennyson, Scofield & Ksepka, 2020: Proc. R. Soc. B 287(1932): 2 – southern Taranaki.

Described from multiple well-preserved late Pliocene (3.36–3.06 Ma) fossils from southern Taranaki (Thomas, Tennyson et al. 2020).

Genus Eudyptula Bonaparte

Eudyptula Bonaparte, 1856: Compt. Rend. Séa. Acad. Sci., Paris 42: 775 – Type species (by monotypy) Aptenodytes minor J.R. Forster = Eudyptula minor (J.R. Forster).

The reference to a “Eudyptula fossil in New Zealand about 24 mya” (A. Baker et al. 2006) appears to refer to a specimen of a small penguin that has no close affinity with Eudyptula, as discussed by Fordyce & Jones (1990), Fordyce (1991b), and Acosta Hospitaleche et al. (2004). Simpson (1975) claimed possible late Pliocene records of the extant E. minor (localities not stated); however, Thomas et al. (2023) found no referable material, and suggested that the record(s) likely referred to Pleistocene infill deposits.

➤ †Eudyptula wilsonae Thomas, Tennyson, Marx & Ksepka
Wilson’s Penguin

Eudyptula wilsonae Thomas, Tennyson, Marx & Ksepka, 2023: Journal of Paleontology 97: 712 – Tangahoe Formation, southern Taranaki.

Known from two late Pliocene (3.36–3.06 Ma) skulls found on the south Taranaki coast (Thomas et al. 2023). The age of these fossils precedes the proposed molecular divergence between E. minor minor and E. m. novaehollandiae at 1.34 Ma (Cole et al. 2022).

Order PROCELLARIIFORMES: Albatrosses, Petrels, and Shearwaters

Family DIOMEDEIDAE G.R. Gray: Albatrosses

Genus †Plotornis Milne-Edwards

Plotornis Milne-Edwards, 1874: Ann. Soc. Géol. 11(3): 4, 5 – Type species (by monotypy) Plotornis delfortrii Milne-Edwards.

➤ †Plotornis archaeonautes Ksepka, Tennyson, Richards & Fordyce
Hakataramea Albatross

Plotornis archaeonautes Ksepka, Tennyson, Richards & Fordyce, 2023: Journ. Roy. Soc. NZ. 54: 647 – Mount Harris Formation, Hakataramea Quarry, South Canterbury (online 13 November 2023).

Known from an early Miocene (22.7–22.0 Ma) partial skeleton and two other fragments found in the Hakataramea Quarry, Hakataramea River valley, South Canterbury (Ksepka, Tennyson et al. 2023). The authors placed Plotornis in family “Pan-Diomedeidae” outside the clade of extant albatrosses (Ksepka, Tennyson et al. 2023). We include it within Diomedeidae as their Fig. 4 shows branching within Pan-Diomedeidae to be shallower than within Procellariidae.

Genus †Aldiomedes G. Mayr & Tennyson

Aldiomedes G. Mayr & Tennyson, 2020: Ibis 162: 724 – Type species (by original designation) Aldiomedes angustirostris G. Mayr & Tennyson.

➤ †Aldiomedes angustirostris G. Mayr & Tennyson
Alastair’s Albatross

Aldiomedes angustirostris G. Mayr & Tennyson, 2020: Ibis 162: 724 – Ohawe Beach, south Taranaki.

Known only from a fossil skull found in a late Pliocene (3.4–3.0 Ma) deposit on the south Taranaki coast (G. Mayr & Tennyson, 2020).

Family PROCELLARIIDAE Leach: Fulmars, Petrels, Prions, and Shearwaters

Genus Macronectes Richmond

Ossifraga Hombron & Jacquinot, 1844: Compt. Rend. Séa. Acad. Sci., Paris 18: 356 – Type species (by monotypy) Procellaria gigantea Gmelin. Junior homonym of Ossifraga Wood, 1835.

Macronectes Richmond, 1905: Proc. Biol. Soc. Washington 18: 76. Nomen novum for Ossifraga Hombron & Jacquinot, 1844.

➤ †Macronectes tinae Tennyson & Salvador
Tina’s Giant Petrel

Macronectes tinae Tennyson & Salvador, 2023: Taxonomy 3: 61 – Tangahoe Formation, Hāwera, southern Taranaki.

Known from a late Pliocene (3.36–3.06 Ma) skull and humerus found on the south Taranaki coast (Tennyson & Salvador 2023).

Genus Procellaria Linnaeus

Procellaria Linnaeus, 1758: Syst. Nat., 10th edition 1: 131 – Type species (by subsequent designation) Procellaria aequinoctialis Linnaeus.

➤ †Procellaria altirostris Tennyson & Tomotani
Deep-billed Petrel

Procellaria altirostris Tennyson & Tomotani, 2021: Pap. Avulsos Zool. 61: e20216116: 4 – Ohawe Beach, south Taranaki.

Known from a late Pliocene (3.6–3.0 Ma) partial skeleton found on the south Taranaki coast (Tennyson & Tomotani 2021a).

Genus Ardenna Reichenbach

Ardenna Reichenbach, 1853: Handb. Spec. Ornith 3: iv – Type species (by original designation and monotypy) Puffinus major (Faber) = Ardenna gravis (O’Reilly).

N. Henderson & Gill (2010) described a shearwater skull similar in size and proportions to “Puffinus pacificus” (now Ardenna pacifica) from a mid-Pliocene deposit near Taihape, North Island.

➤ †Ardenna davealleni Tennyson & Mannering
Pom’s Shearwater

Ardenna davealleni Tennyson & Mannering, 2018: Tuhinga 29: 3 – Ohawe Beach, south Taranaki.

Known from two late Pliocene (3.4–3.0 Ma) partial skeletons found on the south Taranaki coast (Tennyson & Mannering 2018).

Genus Pelecanoides La Cépède

Pelecanoides La Cépède, 1799: Tableaux Method. Mamm. Oiseaux: 13 – Type species (by monotypy) Procellaria urinatrix Gmelin = Pelecanoides urinatrix (Gmelin).

An indeterminate Pelecanoides specimen qualitatively distinct from modern taxa was reported from the Waiauan, middle Miocene, Double Corner Shell Beds of North Canterbury (Scofield et al. 2006). It is represented by elements not yet known for P. miokuaka and so direct comparison is not yet possible.

➤ †Pelecanoides miokuaka T. Worthy, Tennyson, Jones, McNamara & Douglas
Miocene Diving Petrel

Pelecanoides miokuaka T. Worthy, Tennyson, Jones, McNamara & Douglas, 2007: Journ. Syst. Palaeont. 5(1): 8, figs 3A, D – St Bathans, Central Otago.

Known from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage, from the lower Bannockburn Formation, Manuherikia Group; near St Bathans, Otago (Worthy et al. 2007).

Order PELECANIFORMES: Pelicans, Herons, and Ibises

Family ARDEIDAE Leach: Herons & Bitterns

Subfamily ARDEINAE Leach: Herons and Egrets

Genus †Matuku Scofield, T. Worthy & Tennyson

Matuku Scofield, T. Worthy & Tennyson, 2010: Rec. Aust. Mus. 62: 93 – Type species (by original designation) Matuku otagoense Scofield, T. Worthy & Tennyson.

➤ †Matuku otagoense Scofield, T. Worthy & Tennyson
St Bathans Heron

Matuku otagoense Scofield, T. Worthy & Tennyson, 2010: Rec. Aust. Mus. 62: 93 – St Bathans, Central Otago.

Known from fossil remains from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage, Otago (Scofield, Worthy et al. 2010).

Subfamily BOTAURINAE Reichenbach: Bitterns

Genus †Pikaihao T. Worthy, J. Worthy, Tennyson & Scofield

Pikaihao T. Worthy, J. Worthy, Tennyson & Scofield, 2013: Palaeont. Journ. 47: 1333 – Type species (by original designation) Pikaihao bartlei T. Worthy, J. Worthy, Tennyson & Scofield.

➤ †Pikaihao bartlei T. Worthy, J. Worthy, Tennyson & Scofield
Bartle’s Bittern

Pikaihao bartlei T. Worthy, J. Worthy, Tennyson & Scofield, 2013: Palaeont. Journ. 47: 1333 – St Bathans, Central Otago.

Known from fossil remains from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage, Otago (Worthy, Worthy, Tennyson & Scofield 2013).

Order Incertae sedis

Family ZEALANDORNITHIDAE T. Worthy, Scofield, Salisbury, Hand, De Pietri & Archer: False-colies

Genus †Zealandornis T. Worthy, Scofield, Salisbury, Hand, De Pietri & Archer

Zealandornis T. Worthy, Scofield, Salisbury, Hand, De Pietri & Archer, 2022: Journ. Ornith. 163: 650 – Type species (by original designation) Zealandornis relictus T. Worthy, Scofield, Salisbury, Hand, De Pietri & Archer.

➤ †Zealandornis relictus T. Worthy, Scofield, Salisbury, Hand, De Pietri & Archer
Zealandian False Colie

Zealandornis relictus T. Worthy, Scofield, Salisbury, Hand, De Pietri & Archer, 2022: Journ. Ornith. 163: 650 – Manuherikia River, St Bathans, Otago.

Known from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage from the lower Bannockburn Formation, Manuherikia Group; near St Bathans, Otago (Worthy, Salisbury, Hand et al. 2022b).

Order PSITTACIFORMES: Cockatoos, Parrots, and Parakeets

Family STRIGOPIDAE Bonaparte: Kākāpō, and Kākā and Kea

Subfamily INCERTAE SEDIS

Genus †Heracles T. Worthy, Hand, Archer, Scofield & De Pietri

Heracles T. Worthy, Hand, Archer, Scofield & De Pietri, 2019: Biology Letters 15: 3 – Type species (by original designation) Heracles inexpectatus T. Worthy, Hand, Archer, Scofield & De Pietri.

➤ †Heracles inexpectatus T. Worthy, Hand, Archer, Scofield & De Pietri
St Bathans Giant Parrot

Heracles inexpectatus T. Worthy, Hand, Archer, Scofield & De Pietri, 2019: Biology Letters 15: 3 – Manuherikia River, Otago.

Known from two partial tibiotarsi from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage, Otago (Worthy et al. 2019, 2021).

Subfamily NESTORINAE Bonaparte: Kākā and Kea

Nestoridae Bonaparte, 1849: Consp. Syst. Ornith.: 1 – Type genus Nestor Lesson, 1830.

Genus †Nelepsittacus T. Worthy, Tennyson & Scofield

Nelepsittacus T. Worthy, Tennyson & Scofield, 2011: Journ. Vert. Paleont. 31: 1104 – Type species (by original designation) Nelepsittacus minimus T. Worthy, Tennyson & Scofield.

We recognise genus Nelepsittacus as part of subfamily Nestorinae (cf. Worthy et al. 2011b, who treated Nestoridae as a family).

➤ †Nelepsittacus minimus T. Worthy, Tennyson & Scofield
Little St Bathans Parrot

Nelepsittacus minimus T. Worthy, Tennyson & Scofield, 2011: Journ. Vert. Paleont. 31: 1104 – St Bathans, Central Otago.

Known from fossil remains from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage, Otago (Worthy et al. 2011b).

➤ †Nelepsittacus donmertoni T. Worthy, Tennyson & Scofield
Merton’s Parrot

Nelepsittacus donmertoni T. Worthy, Tennyson & Scofield, 2011: Journ. Vert. Paleont. 31: 1110 – St Bathans, Central Otago.

Known from fossil remains from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage, Otago (Worthy et al. 2011b).

➤ †Nelepsittacus daphneleeae T. Worthy, Tennyson & Scofield
Lee’s Parrot

Nelepsittacus daphneleeae T. Worthy, Tennyson & Scofield, 2011: Journ. Vert. Paleont. 31: 1112 – St Bathans, Central Otago.

Known from fossil remains from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage, Otago (Worthy et al. 2011b).

Order PASSERIFORMES: Passerine (Perching) Birds

Family ACANTHISITTIDAE Sundevall: New Zealand Wrens

Genus †Kuiornis T. Worthy, Hand, Nguyen, Tennyson, J. Worthy, Scofield, Boles & Archer

Kuiornis T. Worthy, Hand, Nguyen, Tennyson, J. Worthy, Scofield, Boles & Archer, 2010: Journ. Vert. Paleont. 30: 482 – Type species (by original designation) Kuiornis indicator T. Worthy, Hand, Nguyen, Tennyson, J. Worthy, Scofield, Boles & Archer.

➤ †Kuiornis indicator T. Worthy, Hand, Nguyen, Tennyson, J. Worthy, Scofield, Boles & Archer
St Bathans Wren

Kuiornis indicator T. Worthy, Hand, Nguyen, Tennyson, J. Worthy, Scofield, Boles & Archer, 2010: Journ. Vert. Paleont. 30: 482 – St Bathans, Central Otago.

Known from fossil remains from the Altonian Stage (early Miocene; 19–16 Ma) St Bathans assemblage, Otago (Worthy, Hand et al. 2010).

INCERTAE SEDIS

Order (unknown)

Genus †Manu Marples

Manu Marples, 1946: Trans. Roy. Soc. N.Z. 76: 133 – Type species (by monotypy) Manu antiquus Marples.

The former placement of Manu in family Diomedeidae (order Procellariiformes), based on a part furcula (being the only material available) is uncertain (Olson 1985b). G. Mayr (2009) suggested that Manu antiquus was possibly a pelagornithid, but see G. Mayr & Rubilar-Rogers (2010).

➤ †Manu antiquus Marples
(bird of unknown affinities)

Manu antiquus Marples, 1946: Trans. Roy. Soc. N.Z. 76: 133 – Duntroon, North Otago.

Known from a middle to late Oligocene deposit, near Duntroon, North Otago (Marples 1946b).

Genus †Australornis G. Mayr & Scofield

Australornis G. Mayr & Scofield, 2014: Journ. Roy. Soc. N.Z. 44: 49 – Type species (by original designation) Australornis lovei G. Mayr & Scofield.

➤ †Australornis lovei G. Mayr & Scofield
Love’s Paleocene Seabird

Australornis lovei G. Mayr & Scofield, 2014: Journ. Roy. Soc. N.Z. 44: 49 – Waipara River, North Canterbury.

Known from one fossil only, from the Waipara River, North Canterbury, from a late early Paleocene deposit, 61.6–60.5 Ma (G. Mayr & Scofield 2014).