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Thesis Front Matter Tertiary fossil waterfowl (Aves: Anseriformes) of Australia and New Zealand Trevor H. Worthy Table of contents CHAPTER 1 1.1 1.2 1.3 1.3.1 1.3.2 1.3.3 1.3.4 1.3.4.1 1.3.4.2 1.3.4.3 1.4 1.5 1.6 1.7 1.8 CHAPTER 2 Journal of Systematic Palaeontology CHAPTER 3 ManuherikiaDunstanettaMiotadorna Palaeontology CHAPTER 4 Manuherikia Journal of the Royal Society of New Zealand CHAPTER 5 Zoological Journal of the Linnean Society CHAPTER 6 Journal of Vertebrate Paleontology CHAPTER 7 Emu CHAPTER 8 Tadorna Transactions of the Royal Society of South Australia CHAPTER 9 9.1 9.2 9.3 9.4 9.5 9.6 R 9.6.1 Presbyornis 9.6.2 CnemiornisCereopsis 9.6.3 Dendrocygna 9.6.4 Plectropterus 9.6.5 9.6.6 9.6.7 9.7 CHAPTER 10 Appendix 1 Proceeding of the National Academy of Sciences Appendix 2 Anser scaldiiBulletin of the British Ornithologist’s Club Abstract Anseranas CereopsisStictonetta MalacorhynchusBiziura DendrocygnaOxyura Manuherikia DunstanettaMatanasMiotadorna ManuherikiaDunstanetta Miotadorna MiotadornaTadorna ManuherikiaDunstanettaStictonettaMalacorhynchus OxyuraBiziuraManuherikia Cereopsis PinpanettaAustralotadorna Pinpanetta ManuherikiaDunstanetta StictonettaMionetta MalacorhynchusPinpanettaManuherikiaDunstanettaOxyura NomonyxBiziuraThalassornis Dendrocygna AnserpicaAnatalavis Pinpanetta Anseranas semipalmataCereopsis novaehollandiaeCygnus atratusTadorna tadornoidesBiziura lobataOxyura australisAnasA. castaneaAA. gracilisAythyaaustralis Tirarinetta kanunka TadornaT. tadornoides M Mionetta Pinpanetta Cygnus, Anas, Aythyain situ Declaration GGGGGGGGGG Acknowledgements AM AMNH ANWC BMNH CM FU MV RNMNZ NTM SAM QM USNM Université Claude Bernard This thesis is dedicated to my wife Jennifer P. Worthy 1 CHAPTER 1 1.1 Anseriform phylogeny The Anseriformes, or waterfowl, are now widely regarded as the sister group to the Galliformes (chickens and kin), together forming Galloanserae, as the sister group to the rest of Neornithes (e.g., Sibley et al. 1988; Sibley & Ahlquist 1990; Cracraft 2001; Cracraft et al. 2004; Fain & Houde 2004; Harrison et al. 2004). Extant members of Anseriformes are widely regarded as including the Anhimidae (screamers of South America), Anseranatidae (Magpie Goose, Australia), and true waterfowl Anatidae (swans, geese and ducks, cosmopolitan), for example, as treated by Delacour & Mayr (1945), Woolfenden (1961), Johnsgard (1968), Olson & Feduccia (1980), Olson (1985), Ericson (1997), Livezey (1997a), Dickinson (2003), and Kear (2005). Anhimidae is the basal family within Anseriformes (Livezey 1986, 1997a; Clarke et al. 2005; Livezey & Zusi 2007), and as their fossil record is so far restricted to South America (Alvarenga 1999), the group will not be considered further below. 1.2 The modern Australasian fauna The modern native Australian waterfowl fauna comprises two families: the monotypic Anseranatidae for Anseranas semipalmata (Magpie Goose), and the Anatidae, with 18 species in 12 genera (Condon 1975; Marchant & Higgins 1990; Christidis & Boles 1994, 2008). Of these, Cygnus atratus (Black swan), Biziura lobata (Musk Duck), Oxyura australis (Blue-billed duck), Stictonetta naevosa (Freckled Duck), Cereopsis novaehollandiae (Cape Barren Goose), Chenonetta jubata (Wood Duck), Malacorhynchus membranaceus (Pink-eared Duck) and Aythya australis (White-eyed Duck), are from genera that are monotypic in Australia. There are two species each in the tropical pygmy- geese (Nettapus), the tropical whistling ducks (Dendrocygna), and the shelducks (Tadorna), and four species in the cosmopolitan genus Anas. Of these, Anseranas, Cereopsis, Nettapus and Stictonetta are endemic Australian taxa. Biziura, Chenonetta, and Malacorhynchus are now endemic Australia taxa, but were part of the Recent fauna of New Zealand (see below), so are best listed as endemic Australasian taxa. In its Recent fauna (those that were extant until human intervention about 500 years ago), New Zealand had 17 species in 11 genera (Turbott 1990; Worthy 2002, 2005; Worthy 2 & Holdaway 2002; Worthy & Olson 2002). Three New Zealand genera are endemic: Cnemiornis (extinct geese, 2 spp), Hymenolaimus (Blue Duck) and Pachyanas (Chatham Duck). While Livezey (1989a, 1997a) considered Cnemiornis a primitive monotypic family basal to anserines, Worthy et al. (1997) used an enlarged morphological dataset and genetic data to show that it was the sister taxon to Cereopsis. The genera Chenonetta (=Euryanas), Oxyura (stiff-tailed ducks), Biziura, and Malacorhynchus are each known from New Zealand by a single now extinct species that has its sister taxon still extant in Australia (Olson 1977a; Worthy 1995, 2002; Worthy & Olson 2002). Of the extant fauna, Tadorna variegata (Paradise Shelduck) and the three teal (Anas chlorotis, A. nesiotis and A. aucklandica are closely related to A. castanea in Australia, and three other Anas species are shared with Australia (Marchant & Higgins 1990; Kear 2005). An undescribed extinct shelduck from the Chatham Islands, and Aythya novaeseelandiae and Mergus australis, both with Northern Hemisphere affinities, complete the fauna. Thus the Recent New Zealand and Australian faunas were intimately linked in recent anseriform evolution. Within this Australasian fauna many taxa, particularly the monotypic or less speciose genera are considered relatively basal in Anseriformes: Anseranatidae is accepted as the sister group to Anatidae; Dendrocygna (Whistling ducks, 2 Australian sp.) is the most basal anatid genus; and Oxyura, Biziura, and Stictonetta (Stictonettinae) are all now considered primitive anatids (Frith 1964; Madsen et al. 1988; Sibley & Ahlquist 1990; Christidis & Boles 1994; Sraml et al. 1996; Livezey 1997a; Johnson & Sorenson 1999; McCracken et al. 1999; Sorenson et al. 1999; Donne-Goussé et al. 2002; Dickinson 2003; McCracken & Sorenson 2005; Callaghan & Harshman 2005). After considering the photoperiodic breeding responses of Australian taxa, Kear & Murton (1976) concluded that Cereopsis was an ancient Australian resident, and further that swans and tadornines also originated in Southern Hemisphere temperate regions. Further, while the aberrant Malacorhynchus is often placed relatively basal within Anatinae (e.g. Marchant & Higgins 1990; Livezey 1997a; Dickinson 2003; Callaghan & Harshman 2005), feather protein evidence (Brush 1976), the limited genetic evidence so far available (Sraml et al. 1996), and morphological and behavioural evidence (Frith 1977; Olson & Feduccia 1980), suggest that it should be classified outside of Anatinae and before Tadorninae. Fullagar (in Kear (2005: 442), considered this taxon to be part of the old endemic component of Australia’s avifauna with no close relatives elsewhere. Christidis & Boles (2008) placed it after Tadorna and Chenonetta, along the anatine stem. 3 In this thesis, I recognise the following subfamilies within the Anatidae: Dendrocygninae (whistling ducks); Anserinae (swans and geese), in which I include the tribe Cereopsini for Cereopsis (Cape Barren Goose) and Cnemiornis (New Zealand fossil geese); Tadorninae (shelducks and sheldgeese); Anatinae (dabbling ducks and scaup); Merginae (sea ducks) and Oxyurinae (stiff-tailed ducks), following Turbott (1990). Oxyurinae traditionally includes stiff-tailed ducks of the genera Nomonyx, Oxyura, Biziura, and Thalassornis (Delacour & Mayr 1945). But, more recently, Biziura has been listed Aves incertae sedis (e.g., McCracken et al. 1999; Callaghan & Harshman 2005), and Thalassornis has been considered sister to Dendrocygna (Johnsgard 1968; Raikow 1971; Livezey 1997a; Callaghan & Harshman 2005). In this thesis, however, the hypothesis is advanced in Chapters 3 and 5, that the Oxyurinae should be expanded from Delacour & Mayr’s (1945) definition to include also Malacorhynchus (Australasian pink-eared ducks), Stictonetta (Freckled Duck), and the Oligo-Miocene fossil taxa Mionetta, Manuherikia and Dunstanetta). As detailed above, a substantial number of Australasian taxa are relatively basal in the anseriform radiation, perhaps implying a long history of evolution in the region. It will be instructive to determine when these relatively basal taxa originated and whether they have an older origin than Recent taxa in the Northern Hemisphere. Accordingly, the fossil record is analysed to reveal the evolutionary development of the Australasian waterfowl fauna. In so doing, valuable insights into the origin of Anseriformes in the Southern Hemisphere are obtained. At the onset of this project there were no data on the pre- Quaternary fossil waterfowl of Australasia, therefore, the overall objective of this thesis was to determine the diversity and history of fossil anseriforms in Australasia. First, a review of the global fossil record of anseriforms is necessary to provide a background against which to interpret the Australasian record; this is given below. Following this, an introduction to the avian fossil records of Australia and New Zealand sets the scene against which the waterfowl record is assessed. 1.3 Fossil History of Anseriformes There are several published reviews of the fossil record of waterfowl, which provide a good basis from which to understand
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