Quick viewing(Text Mode)

Thesis Front Matter

Thesis Front Matter

Tertiary waterfowl (Aves: )

of and 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 DunstanettaMiotadorna 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

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 9.6.1 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 scaldiiBulletin of the British Ornithologist’s Club

Abstract Anseranas CereopsisStictonetta DendrocygnaOxyura Manuherikia DunstanettaMatanasMiotadorna ManuherikiaDunstanetta Miotadorna MiotadornaTadorna ManuherikiaDunstanettaStictonettaMalacorhynchus OxyuraBiziuraManuherikia Cereopsis PinpanettaAustralotadorna Pinpanetta ManuherikiaDunstanetta

StictonettaMionetta MalacorhynchusPinpanettaManuherikiaDunstanettaOxyura NomonyxBiziuraThalassornis Dendrocygna Anserpica Pinpanetta Anseranas semipalmataCereopsis novaehollandiaeCygnus atratusTadorna tadornoidesBiziura lobataOxyura australisA. castaneaAA. gracilisaustralis Tirarinetta kanunka TadornaT. tadornoides Mionetta Pinpanetta

Cygnus, Anas, Aythyain situ

Declaration

Acknowledgements AM AMNH ANWC BMNH CM FU MV NMNZ 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 (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 ( of ), (Magpie , Australia), and true waterfowl (, geese and , 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 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 (), and the Anatidae, with 18 in 12 genera (Condon 1975; Marchant & Higgins 1990; Christidis & Boles 1994, 2008). Of these, Cygnus atratus (Black ), Biziura lobata (Musk ), Oxyura australis (Blue-billed duck), Stictonetta naevosa (), Cereopsis novaehollandiae (), 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 (Tadorna), and four species in the cosmopolitan 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 (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 () 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 ) 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 australis, both with Northern Hemisphere affinities, complete the fauna. Thus the Recent New Zealand and Australian faunas were intimately linked in recent anseriform .

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 (e.g. Marchant & Higgins 1990; Livezey 1997a; Dickinson 2003; Callaghan & Harshman 2005), 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 . 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); (swans and geese), in which I include the 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- 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 the diversity of fossil anseriforms. The first to attempt a world-wide review was Lambrecht (1933), who described all known fossil taxa of and figured many, with synonymies given. Two very useful works appeared in 1964 4

and probably are among the most often quoted palaeornithological works in existence. Howard’s (1964) chapter on the fossil anseriforms in Delacour (1964) listed in a consistent format most taxa then known, giving the primary taxon reference, and referred material, locality and age, description, measurements and remarks for all. Also in 1964, Brodkorb published the Anseriformes section of his monumental ‘Catalogue of Fossil Birds’, listing all taxa with full synonymies and age ranges (Brodkorb 1964), with additions in Brodkorb (1978). Since then, Bocheski (1997) listed all described European fossil species, indicating their current taxonomic status. Most recently, Mlíkovský (2002) published an extensively annotated work on the Cenozoic birds of , wherein all taxa are listed with complete synonymies and distributions. The work has been heavily criticized for its idiosyncratic classification, which does not follow any modern system, and for many novel taxonomic decisions made within it, often with little or no justification (Mourer- Chauviré 2004). Thus, new synonymies by Mlíkovský (2002) noted below may not withstand scrutiny. These criticisms notwithstanding, the work is very well referenced and indexed, with authors, countries, localities, collections, stratigraphic units, family- and genus-group names and species, each independently indexed, with an extensive bibliography, making it a most useful resource. These resources reveal the amount of palaeontological activity in different regions of the world, i.e. comparatively great amounts of effort in Europe and , little in South America, and nearly none in . The following review uses these sources as a basis from which to assess the taxonomic and stratigraphic history of anseriforms in the world.

1.3.1 The origin of Anseriformes

The oldest anseriform known is Vegavis iaai Clarke, Tambussi, Noriega, Erickson, & Ketchman, 2005 from a pre-KT boundary deposit about 66-68 Ma on Vega Island, western (Clarke et al. 2005). It was initially described as a possible presbyornithid (Noriega & Tambussi 1995) and is figured in Tambussi & Noriega (1996) and Tambussi & Acosta Hospitaleche (2007). Clarke et al. (2005) identified Vegavis as an anseriform more closely related to Anatidae than Anseranatidae and not a presbyornithid, although the Vegavis+Anatidae+Presbyornis was unresolved. However, it is confidently placed in Anseriformes and is more derived than Anhimidae and Anseranatidae showing that the basal of extant Anseriformes (Anhimidae, Anseranatidae, Anatidae) originated prior to the close of the . This date accords with genetic data presented 5

by authors such as Cooper & Penny (1997) and Harrison et al. (2004), which place the Galliformes-Anseriformes split in the .

1.3.2

The extinct Presbyornithidae includes several unnamed forms from the Late Cretaceous of North America (Hope 2002), and taxa from the Late Cretaceous of Mongolia (Kurochkin et al. 2002), and the Palaeocene and of Argentina, North America, and Europe (Olson 1985, 1994; Dyke 2001). Their relationships have been controversial on account of the morphological mosaic nature of their skeleton with Olson & Feduccia (1980) and Olson (1985) regarding them as transitional shorebirds related to , but it is now agreed that they are members of the Anseriformes and the sister group to Anatidae (Olson 1994; Ericson 1997, 2000; Livezey 1997a,b). No members of this group are known from Australasia.

1.3.3 Anseranatidae

The next most significant anseriform fossil is Anatalavis oxfordi Olson, 1999 from the Eocene (European Paleogene Reference Level MP 8) . This taxon, represented by most of the skeleton, was referred to Anseranatidae by Olson (1999). Anatalavis is otherwise known by Anatalavis rex (Shufeldt, 1915) from the Hornerstone Formation in New Jersey, , which is of Early Palaeocene age (Olson 1994; Olson & Parris 1987) or latest Cretaceous age (Parris & Hope 2002). Mlíkovský (2002) placed A. oxfordi into a new genus Nettapterornis, but this move was not supported by Mourer-Chauviré (2004) or Mayr (2005). Further, the referral of A. oxfordi to Anseranatidae was challenged by Dyke (2001) on the basis of a cladistic analysis, which suggested Anatalavis was the sister group of Presbyornithidae + Anatidae. Dyke’s analysis was criticised by Mayr (2005), however, on account of large numbers of missing data and non-inclusion of characters indicating specific anseranatid affinity signalled by Olson (1999).

From the latest (MP 29) in France, Anserpica kiliani Mourer-Chauviré, Berthet & Hugueney, 2004 was described as a member of Anseranatidae by Mourer- Chauviré et al. (2004), indicating greater diversity of this family in the early Tertiary of Europe than in Recent Australia. 6

1.3.4 Anatidae

1.3.4.1 Eocene-Oligocene taxa (55 – 23.8 Ma; MP 7-30)

No modern genera have been positively identified from deposits of this age.

Romainvillia stehlini Ledebinsky, 1927 from the Late Eocene (MP 20) of France was based on several bones (Brodkorb 1964; Howard 1964). It was suggested to be possibly an anseranatid (Olson 1999; Mlíkovský 2002), but a recent phylogenetic analysis by Mayr (in press) has determined that it is the sister to extant Anatidae and more derived than presbyornithids.

Eonessa anaticula Wetmore, 1938 from the Late Eocene of Utah, North America, known only from bones of a left wing, was the basis for the anatid subfamily Eonessinae. It was removed from Anseriformes and considered of indeterminate familial affinity by Olson & Feduccia (1980).

Harrison & Walker (1976) named Howardia eous on the basis of a partial from the Late Eocene (MP 17) and referred it to Anseriformes. Because the generic name was preoccupied, Harrison & Walker (1979a) subsequently proposed the new genus Palaeopapia for it and to which they also referred a partial . From material of Lower Oligocene age (MP 21-23), a second species in this genus, P. hamsteadiensis, based on a scapula, and a new genus and species, Paracygnopterus scotti, based on the humeral end of a coracoid, were described by Harrison & Walker (1979b). All three taxa are regarded as Aves incertae sedis by Dyke (2001) as no synapomorphies of Anseriformes had been identified in the . Harrison & Walker (1976) also named the new genus and species Petropluvialis simplex on two from the Late Eocene (MP 17) of England; they referred the taxon to Burhinidae, but this specimen is an anseriform similar to Romainvillia, as indicated by the presence of a procoracoidal foramen (Mayr & Smith 2001).

The lowermost Oligocene of Belgium has revealed several anseriform fossils including coracoids, one similar to Romainvillia and one similar to Paracygnopterus (Mayr & Smith 2001), though Mayr (in press) altered the latter referral to Anatidae, gen. et. sp. indet. 7

The best known Oligocene anseriform is Cygnopterus Lambrecht, although its phylogenetic affinities are uncertain. A single Middle Oligocene species Cygnopterus affinis (Van Beneden, 1883) from Belgium (MP 23-24), is generally considered to have anserine affinities (Lambrecht 1933; Brodkorb 1964; Howard 1964; Cheneval 1984; Olson 1985). Cheneval (1984) identified it as a swan in Cygnini, however Louchart et al. (2005) listed several differences that indicated Cygnopterus was an anserine, but ‘unrepresentative’ of swans. Mlíkovský (2002: 112) synonymised Palaeopapia hamsteadiensis with C. affinis, but this was an error, perhaps due to confusion of the figures in Harrison & Walker (1979b: Pl. 1), in which Headonornis hantoniensis (a presbyornithid) and Palaeopapia hamsteadensis (an anatid) are both shown, as noted by Mayr (in press). Cygnopterus lambrechti Kurochkin, 1968, based on a distal humerus from the Middle Oligocene of Kazakhstan (Kurochkin 1968), was synonymised with the Agnopterus turgaiensis Tugarinov, 1940, and thus is not an anseriform (Mlíkovský & Švec 1986). A third species from the Miocene is discussed below. The anserine affinities of Cygnopterus were questioned by Mayr (2005, in press), who noted especially that the referred coracoids of C. alphonsi are very similar to those of presbyornithids. Yet, the type tarsometatarsus of C. alphonsi is short and robust like those of anserines, unlike the elongate ones of presbyornithids. The phylogenetic affinities of Cygnopterus and its included species await re-examination of the material. All three Cygnopterus species were placed in Cygnopteridae by Callahan & Harshman (2005).

Two other ‘swans’ have been described from the Oligocene in Europe: Guguschia nailiae Aslanova & Burczak-Abramovicz, 1968, on a part skeleton from the Late Oligocene-Early Miocene of Azerbaijan (Aslanova & Burczak-Abramovicz 1968), and Cygnavus formosus Kurochkin, 1968, on a distal tibiotarsus from the Lower Oligocene of Kazakhstan (Kurochkin 1968; Mlíkovský & Švec 1986; Boev 2000). Louchart et al. (2005) noted that Guguschia had some similarities with Cygnopterus.

Several fossils of Oligocene age have been named in Anas. The species Anas oligocaena Tugarinov, 1940 from the Late Oligocene of Kazakhstan, was referred to Dendrochen in Dendrocygnini by Mlíkovský & Švec (1986) and Cheneval (1987). The taxon Anas benedeni Sharpe, 1899, a replacement name for the preoccupied Anas creccoides Beneden, 1871, based on fossils from the Lower Oligocene (MP 23-24) of Belgium, was found upon re-evaluation to be not assignable to Anseriformes and was relegated to Aves incertae sedis (Brodkorb 1962: 707). Anas basaltica Bayer, 1883, from 8

the early Oligocene (MP 23-24) of Czechia, is an indeterminate (Mlíkovský 2002: 70). The types of Anas skalicensis Bayer, 1883 are bones from the Czech Republic that are indeterminate at the ordinal level; they were regarded as Early Miocene age by Bocheski (1997), and Middle Oligocene (MP 23-24) age by Mlíkovský (2002: 251). In summary none of these Oligocene taxa are correctly attributed to Anas although some are anatids. No crown-group members of modern families are known from pre-Oligocene deposits (Mayr 2005).

From the Late Oligocene–Early Miocene of Argentina, an anseriform of undetermined subfamilial affinity within Anatidae was described as Cayaoa bruneti (Tonni 1980). Also from South America are the poorly known taxa Telornis impressus Ameghino and Loxornis clivus Ameghino (Brodkorb 1964). In a recent revision of some of the Oligocene taxa named by Ameghino, Agnolin (2004) transferred Telornis to Tadorninae and Aminornis excavatus from to Anseriformes: Anserinae, ?Cygnini. However, as these taxa are based on distal humerus and a cranial part of a coracoid, respectively, these referrals are tenuous.

1.3.4.2 Miocene (28.3-5.3 Ma; MN 1-17)

Only the modern genera Cygnus and Anser certainly make their first appearance by the Middle Miocene. By the Late Miocene, Dendrocygna, Oxyura, Mergus, and Bucephala have made appearances in the fossil record. The genus Anas Linnaeus, as now defined (e.g. Livezey 1997a), is not known with certainly from sediments older than the latest Miocene despite large collections of fossil anseriforms, implying that it evolved no earlier than 5-6 Ma.

The best known ducks of the Early-Middle Miocene of the Northern Hemisphere are relatively primitive forms that also do not belong in Anas. In Europe, the most abundant anatid is Anas blanchardi Milne-Edward, 1863 (Milne-Edwards 1863, 1867-71). Cheneval (1983, 1987) recognised that this taxon and associated anatids from Saint-Gérand-le-Puy should not be in Anas, and placed them in Dendrochen. Livezey & Martin (1988) restudied Anas blanchardi and erected for it the genus Mionetta in a new subfamily Dendrocheninae, taxonomically placed between Dendrocygninae and Thalassorninae. Mionetta blanchardi first appeared in the Late Oligocene (MP 30) (Mourer-Chauviré et al. 2004), but ranges across the Early Miocene (Neogene Mammal Faunal Zones MN 1-3) (Mlíkovský 2002). Of 9

the two taxa found with M. blanchardi, the relatively rare larger taxon, Anas consobrina Milne-Edward, 1867-71, was considered to represent bones of large individuals of Mionetta blanchardi by Livezey & Martin (1988), who also transferred the smaller Anas natator Milne-Edward, 1867-71 to Mionetta. The latter ranges in age from Late Oligocene (MP 30) (Mourer-Chauviré et al. 2004) to the Early Miocene (MN 2-3) (Mlíkovský 2002). The species Aythya (=Fuligula) arvernensis (Lydekker, 1891), considered in need of reassessment by Cheneval (1987), was included in the synonymy of Mionetta blanchardi by Mlíkovský (2002).

There three described Early Miocene waterfowl from the Flint Hill Fauna of South Dakota in North America (Brodkorb 1964). The unusual swan-sized diver Paranyroca magna A.H. Miller & Compton, 1939, was placed in a monotypic family Paranyrocidae by Miller & Compton (1939) on account of the hypotarsus having only two ridges, but it was later reduced to a subfamily of Anatidae (Brodkorb 1964). That this taxon is known only from tarsometatarsi precludes understanding its true relationships to other anatids, and its possession of two hypotarsal ridges is similar only to anhimids among anseriforms. Miller (1944) described Dendrochen robusta Miller, 1944 and the teal-size Querquedula integra Miller, 1944. Both were subsequently recognised as relatively primitive anatids, with Anas (=Querquedula) integra transferred to Dendrochen by Cheneval (1987), and Dendrochen made the type genus of Dendrocheninae by Livezey & Martin (1988). Other undetermined duck-sized anseriforms are reported from the Late Miocene of California (Miller 1952).

To date, there is only a single record of a putative Dendrocheninae in the southern Hemisphere: a humerus from the Late Miocene of Argentina (Noriega 1995).

Other than these dendrochenines, most other Early Miocene anatids are anserines. Anserines are represented in the Early Miocene of Europe (MN2a) by Cygnopterus alphonsi Cheneval, 1984 and Cygnavus senckenbergi Lambrecht, 1931, with both genera making their first appearance in the Oligocene (Lambrecht 1933; Kurochkin 1968; Cheneval 1987; Mlíkovský 2002). Mlíkovský (2002) synonymized Cygnopterus alphonsi with Cygnavus senckenbergi, an action not accepted by Louchart et al. (2005). As noted above, at least the referred coracoid of C. alphonsi is similar to that of presbyornithids (Mayr in press). Anas robusta Milne-Edward, 1867-71 (based on a -sized distal humerus from Sansan, France, MN 6), was placed in Anserobranta? by Cheneval (1987), but later transferred to Mionetta by Mlíkovský (2002). It is not like Mionetta, however, especially in the short distal extent of the flexor process (in which it is more similar to Cygnus), the conformation 10

of the brachial fossa, and by having the space between the dorsal condyle and the facet for the attachment of the anterior ligament greater than the width of the facet (Cheneval 1987, pl. 1, fig. 1), not less than as in Dendrocygnini (Woolfenden 1961: 6). It is best identified as an anserine in ‘Anserobranta?’.

The true swans, Cygnus, do not appear in Europe until the Middle Miocene (MN7) in Steinheimer Becken, with Cygnus atavus (Fraas, 1870), under which name Anas cygniformis Fraas, 1870 was included as a (Heizmann & Hesse 1995; Mlíkovský 2002). Heizmann & Hesse (1995) reported a further undetermined Cygnus species from the same site. Cygnus (=Cygnanser) csakvarensis (Lambrecht, 1933), from the Late Miocene (MN 10) of Hungary (Mlíkovský 2002), should however, be in Anser (Lambrecht 1933). Cygnus herrenthalsi van Beneden, 1871 from the Middle Miocene (MN 7-8) of Belgium, based on a phalanx is a nomen nudum, but made available by Lambrecht (1933) as Cygnus herenthalsi (Mlíkovský 2002: 253). Because a phalanx is a diagnostically poor element, the taxon should be considered Aves incertae sedis (Howard 1964; Mlíkovský 2002). Cygnus was first reported from the Miocene of North America by Wetmore (1943), but the first named species, Cygnus mariae Bickart, 1990, comes from the Late Miocene (Bickart 1990). The tribe Cygnini has recently been reported from the Late Miocene (c. 7 Ma) of Chad, Africa, represented by a new genus Afrocygnus (Louchart et al. 2005). Another ‘swan-like’ bird was described as Megalodytes morejohni Howard, 1992 from the Middle Miocene of California (Howard 1992), but it is a giant diving anatid not related to swans (Louchart et al. 2005). A large flightless anatid described as a flightless swan with affinities to Megalodytes has been reported from 11-12 Ma sediments in Japan (Hiroshige et al. 2001, 2004), but it has so far not been named nor been the subject of a phylogenetic analysis.

Geese, in the form of an undetermined species of Anser, are first known from the Middle Miocene (MN7) in Germany (Heizmann & Hesse 1995), and occur in the Late Miocene in Europe with Anser thraceiensis Burczak-Abramovicz & Nikolov, 1984 (MN 11-15) and other undetermined species (Howard 1964; Boev 2002; Mlíkovský 2002). Branta made a possible first appearance in Europe in the Late Miocene (MN 12); B. thessaliensis Boev & Koufos, 2006), was described from Greece (Boev & Koufos 2006), but no apomorphies for this genus were advanced. Both Anser and Branta appear in the Middle-Late Miocene of North America (Miller 1961; Bickart 1990), but are preceded by Presbychen abavus Wetmore, 1930 from the Miocene of California (Brodkorb 1964; Howard 1964). From the latest Miocene–Pliocene Big Sandy Formation of Arizona (6.1-4.6 11

Ma), Bickart (1990) named Anser arenosus, A. arizonae, and Branta woolfendeni, and listed two other indeterminate anserines, Anabernicula sp. and three undetermined species of Anas. Two ‘anserines’ are known from the Upper Sarmatian Hipparion Fauna (Upper- Middle Miocene) in the Caucasian region: Anser eldaricus Burczak-Abramovicz & Gadziev, 1978, from the area between Azerbaydzhan and Georgia, was described on the basis of a partial humeral head and three other fragments (Burczak-Abramovicz & Gadziev 1978), all so fragmentary that their generic and familial identifications must be questioned; Anser udabnensis Burczak-Abramovicz, 1957 (MN 9-11), from eastern Georgia, was based on a proximal ulna (Aslanova & Burczak-Abramovicz 1968; Burczak-Abramovicz & Gadziev 1978; Mlíkovský & Švec 1986).

The taxon Chenornis graculoides Portis, 1884 (MN 1-2) is sometimes listed in Anseriformes and/or Anserinae (Lambrecht 1933; Brodkorb 1964; Howard 1964), but Mlíkovský (2002) considered it Aves incertae sedis, with possible affinity to Phalacrocoracidae.

The tadornines are first known with certainty in Europe with the appearance of the genus Tadorna in the Early Pliocene (Mlíkovský 2002), although, the group may have an earlier record in Europe if Mlíkovský’s (2002) referral of Anserobranta tarabukuni Kurokin & Ganea, 1972, based on a proximal carpometacarpus from the Late Miocene (MN 9-11) of Moldavia, to is correct. Olson (1985) reported specimens of Tadornini from the Middle Miocene Calvert Formation of Maryland, United States, and undescribed fossils similar to Tadorna from the Nördlinger Ries, Middle Miocene of Germany, which remain undescribed (Ballmann, pers. comm. 24 November 2004). Until a description of this ‘tadornine’ is forthcoming its affinities remain unknown, but it may represent a Middle Miocene occurrence of Tadorninae in Europe.

Potentially, another tadornine is Anser scaldii, listed by Lambrecht (1933: 368) with a brief description stating that the taxon is based on a humerus the size of Tadorna casarca at 129 mm long, from the Middle Miocene (MN 7-8) of Belgium. Lambrecht (1933: 368) thus became the author of the name, and wrongly attributing it to van Beneden wherein it is a nomen nudum (Mlíkovský 2002: 125). Anser scaldii is redescribed in Appendix 2 herein, and it is confirmed as an anserine.

There are few other genera of anatids recognised from the Miocene of Europe. Mlíkovský (2002) erected Oxyura doksana (MN 4b) based on a cranial end of a left 12

coracoid from the Czech Republic initially described by Švec & Mlíkovský (1986). This generic attribution may be in doubt, however, as the cranial ends of Oxyura coracoids differ little from those of many anatid genera, including especially those of the Miocene dendrochenine taxa Mionetta and Manuherikia. A Mergus species was reported by Heizmann & Hesse (1995) from Steinheimer Becken (Middle Miocene, MN7) in Germany from abundant fossils. There is a Late Miocene (MN 10) record of Dendronessa sp. (Mlíkovský 2002). Sinanas diatomas Yeh, 1980, of Middle Miocene age from Shandong Province in China, is poorly described and was placed in Anatidae: subfamily incertae sedis by Mlíkovský & Švec (1986) pending re-evaluation of the type. From the Middle Miocene Calvert Formation (c. 14 Ma) of the eastern United States, Alvarez & Olson (1978) described Mergus miscellus Alvarez & Olson, 1978 based on an associated pelvis and left and right tibiotarsi and tarsometatarsi. This generic assignment was criticised by Livezey & Martin (1988), as it had a phenetic basis in which no synapomorphies were identified.

The few Miocene taxa named in Anas all probably are not Anas sensu stricto. The taxa Anas velox Milne-Edwards, 1867-71, whose lectotype is a right carpometacarpus from Sansan (MN 6), and Anas sansaniensis Milne-Edwards, 1867-71, whose lectotype is a distal left tibiotarsus (MN 6-7), are both doubtfully correctly placed in Anas (Mlíkovský 2002), a conclusion I agree with as these elements are of insufficient diagnostic utility for generic attribution. The referred coracoid of Anas velox has “à fosse pneumatique profonde et à bord externe de la facette glénoidale épais” (Cheneval 1987). This species thus cannot be in Anas because members of this genus have coracoids that lack pneumatic foramina within the acrocoracoid; this is contrary to the pneumatic state suggested by this description and as shown in Cheneval (1987, pl.1, fig. 3). Mlíkovský (2002: 118) synonymized Anas meyerii Milne-Edwards, 1867-71 of Germany (MN 7), which Howard (1964) considered indeterminate to genus, with A. velox. Anas luederitzensis Lambrecht, 1929 based on a proximal humerus from the Early Miocene of southwest Africa has a closed ventral pneumotricipital fossa (Howard 1964), so does not belong in Anas.

Several Late Miocene species named in Anas are not generically assignable, but are anatids, and so must be considered Anatidae: genus incertae sedis (Mlíkovský 2002). These include Anas albae Janossy, 1979, a carpometacarpus from Hungary (MN 13); Anas isarensis Lambrecht, 1933, a scapula from Aumeister (MN 9); and Anas eppelsheimensis Lambrecht, 1933, a cranial fragment of a coracoid from Eppelsheim, Germany (MN 9). Similarly, Mlíkovský (2002) considered the following Middle Miocene species to be 13

Anseriformes, genus incertae sedis: Anser brumeli Milne-Edwards, 1871 from France (MN 4); Anas oeningensis Meyer, 1865 from Germany (MN 7); and Anas risgoviensis Ammon, 1918 from Germany (MN 6).

From the Middle Miocene of Europe, other than species of ‘Anas’, the only described anatine is Aythya chauvirae Cheneval, 1987, from Sansan, France (MN 6). Its holotype is a femur, but the hypogeum includes a complete coracoid, a proximal and a distal humerus, a proximal and a distal ulna, and a distal femur (Cheneval 1987). The holotype, as figured in Cheneval (1987: pl. 1, fig. 7a,b) lacks the lateral expansion of the fibular condyle typical of Aythya. It does have a deep popliteal fossa, but so do all diving taxa (e.g. Mergus), and others, (e.g. Malacorhynchus), so this is not an apomorphy of Aythya. I have examined the referred specimens and have concluded that Aythya chauvirae is unlikely to belong in Aythya, and may well be a dendrochenine (sensu Livezey & Martin 1988), and that the hypogeum comprises two taxa (Worthy et al. 2007). From Late Miocene sediments at Lufeng, Yunnan, in the south of China, Lianhai (1985) briefly described Aythya shihubas and reported an undetermined Anas species.

The two species Eutelornis patagonica Ameghino, 1895 and Eoneornis australis Ameghino, 1895 from the Late Miocene of Argentina were based on scanty material and were considered Anseriformes incertae sedis (Lambrecht 1933; Howard 1964).

From ten Late Miocene to Early Pliocene (9.0-4.5 Ma) avifaunas from Florida, United States, Becker (1987) reported 13 anatid taxa: a Dendrocygna sp., a Branta sp., three indeterminate anserines, an indeterminate tadornine, two indeterminate anatines, two indeterminate Anas sp., one indeterminate Aythya sp., an Oxyura cf. O. dominica (Linnaeus, 1766), and the extinct Bucephala ossivallis Howard, 1963. Olson & Rasmussen (2001) reported a single specimen possibly of ?Anas from the Pungo River Formation, North Carolina, of Middle Miocene age (c. 14 Ma).

The accepted fossil anseriforms discussed above are listed in Tables 1 and 2 by geological age under relevant taxonomic headings. Excluding taxa whose affinity is unresolved, such as those under Anatidae incertae sedis, the following summary is made. Anseranatidae is represented by three taxa ranging from to Late Oligocene in age. Anserinae ranges from the Oligocene to the present, and some 25 taxa are known from the Oligocene and Miocene periods. Modern anserine genera appeared only in the Middle Miocene. Dendrocygninae has a single Late Miocene record. Dendrocheninae, ranged from 14

Late Oligocene to Middle Miocene, and Oxyura has been reported from the Late Miocene. Tadorninae may have members in the Middle Miocene, if undescribed material so referred withstands scrutiny, otherwise, its first definite occurrence is in the Late Miocene. Anatinae may have its first members by the Middle Miocene, if Megalodytes is correctly referred to this group, otherwise and more likely, it arose in the Late Miocene. Merginae may have a first record in the Middle Miocene, and Aythyinae appear first in the Late Miocene.

Table 1. A list of accepted fossil anseriform taxa (Anseriformes incertae sedis, Anseranatidae, and Anatidae incertae sedis) older than the Pliocene. Taxa are listed by geological age and under the family/subfamily they are considered herein to belong to. Anseriformes incertae Anseranatidae Anatidae incertae sedis sedis Paleocene Anatalavis rex Eocene Palaeopapia eous Anatalavis oxfordi Romainvillia stehlini Petropluvialis simplex Oligocene Palaeopapia hamsteadiensis ‘Anas’ oligocaena Paracygnopterus scotti Loxornis clivus Late Oligocene- Paranyroca magna Anserpica kiliani Cayaoa bruneti Early Miocene Middle Miocene Anas velox Anas sansaniensis Late Miocene Anser brumeli Sinanas diatomas Anas oeningensis Anas luederitzensis Anas risgoviensis Anas albae Eutelornis patagonica Anas isarensis Eoneornis australis Anas eppelsheimensis

15

Table 2. A list of accepted fossil taxa in Anatidae older than the Pliocene. Taxa are listed by geological age and under the subfamily they are considered herein to belong to. A ‘?’ before the name indicates it is doubted that the taxon is correctly attributed to the subfamily it is listed under. A ‘?’ after the genus indicates that there is cause to question the generic attribution.

Anserinae Dendrocygninae Oxyurinae & Tadorninae Anatinae Merginae and Dendrocheninae Aythyinae Oligocene Cygnopterus affinis ?Telornis impressus ?Cygnopterus lambrechti Guguschia nailiae Cygnavus formosus Aminornis excavatus Late Oligocene- Cygnopterus alphonsi Mionetta blanchardi Early Miocene Cygnavus senckenbergi Mionetta consobrina Mionetta natator Dendrochen robusta Dendrochen integra Middle Anserobranta? robusta Aythya chauvirae cf Tadorna Nördlinger Miocene Ries & Calvert Fm Cygnus atavus Cygnus sp. Megalodytes morejohni Mergus? miscellus Presbychen abavus Megalodytes sp Japan Anser sp. Anser scaldi Late Miocene Cygnus csakvarensis Dendrocygna sp. Oxyura? doksana Anserobranta Dendronessa sp. Mergus sp. tarabukuni Cygnus mariae Oxyura cf dominica Anas sp. 1& 2 Florida Bucephala ossivallis Afrocygnus chauvirae Anas sp. 1, 2, 3 Aythya shihubas Arizona Anser thraceiensis Aythya sp. Florida Anser arenosus 16

Anser arizonae Anser? eldaricus Anser? udabnensis Branta woolfendeni Branta? thessaliensis Branta sp. Anabernicula sp.

17

1.3.4.3 Pliocene Record (5.3-1.8 Ma)

Globally most modern genera of waterfowl have their first appearance in the Pliocene and many modern species have a Pliocene record.

Anseriforms are globally widespread in Pliocene deposits (Howard 1964; Mlíkovský 2002). Howard’s (1964) comprehensive review reported 28 Pliocene species of anseriforms including three genera and 18 species that were extinct. A single, and the oldest record for Dendrocygninae, is Dendrocygna eversa Wetmore, 1924 from Arizona, placed in the Late Pliocene (Howard 1964), but in the Lower Pleistocene (Brodkorb 1964).

Pliocene anserines are diverse, with most in extant genera, and several species still extant (number of extinct species in brackets) in Cygnus 3(2), Anser 2(1), and Branta 3(3) (Miller, L. 1930, 1944; Miller, A. 1948; Howard 1964). Only one recorded Pliocene anserine is an extinct genus; Eremochen russelli Brodkorb, 1961 (Brodkorb 1964; Howard 1964). Cygnus hibbardi Brodkorb, 1958, described from the Early Pleistocene Hagerman Lake beds, Idaho (Brodkorb 1958a), was later listed in Olor (Brodkorb 1964), and the lake beds assigned a Pliocene age Howard (1964).

Tadornines were represented by a single extinct species Anabernicula minuscula (Wetmore, 1924). A number species in modern genera were listed by Howard (1964) as follows, (number of extinct species in brackets): Anas 9 (4), Aythya 4 (2), Nettapus 1 (1), Bucephala 2 (1). Howard (1964) did not accept Nettion as distinct from Anas and so listed Nettion bunkeri Wetmore, 1944 in Anas. Nettion bunkeri was reported from Pliocene sediments by Brodkorb (1958b). Two other teal, Nettion greeni Brodkorb, 1964 and N. ogallalae Brodkorb, 1962, were also listed from Pliocene sites by Brodkorb (1964). Nettion was used as a subgenus of Anas by Livezey (1997a). Since Brodkorb’s (1964) listing in Aythya of Fuligula aretina Portis, 1889 and Fuligula sepulta Portis, 1889, both of the Upper Pliocene of Italy, both taxa have been transferred to Anas by Cheneval (1987), and most recently synonymised with Anas platyrhynchos by Mlíkovský (2002: 120).

Since Brodkorb’s (1964) and Howard’s (1964) reviews, many Pliocene taxa have been named. From Europe, Boeuf & Mourer-Chauviré (1992) described a small fauna of Late Pliocene (MN 17) age in France, which included Anser sp., the recent shelduck Tadorna cf. T. tadorna, and the new species Bucephala cereti Boeuf & Mourer-Chauviré, 1992. They also reported that Tadorna tadorna is common in karst fillings of Pliocene age in Hungary. Boev (1998, 2002) reported Balcanas pliocaenica Boev, 1998, as ‘a medium 18

sized anatine’, and Anas sp. from the Early Pliocene (MN 14) of Bulgaria; Balcanas pliocaenica was subsequently synonymised with Tadorna tadorna by Mlíkovský (2002: 117). Anas submajor Janossy, 1979, based on an ulna from Hungary (MN 17), has also been included in the synonymy of Tadorna tadorna (Bocheski 1997; Mlíkovský 2002: 117). Boev (2000) described the swan Cygnus verae Boev, 2000 from the Early Pliocene (MN 14) of Bulgaria.

Several anseriforms from the Middle Pliocene of Western Mongolia, reported and figured in Kurochkin (1985), were reviewed by Mlíkovský & Švec (1986) as follows: the proximal coracoid named Anas soporata Kurochkin, 1968 was transferred to Dendrocygna, but inter-generic comparisons were limited; the distal humerus named Anser liskunae Kurochkin, 1976 was placed in Olor as a smaller relative of Olor bewickii (Yarrell, 1830); Cygnus pristinus Kurochkin, 1971, described from a distal radius, was synonymised with Cygnus olor (Gmelin, 1789); the distal tibiotarsus named Anser devjatkini Kurochkin, 1971 was accepted in Anser; the proximal tarsometatarsus named Heterochen vicinus Kurochkin, 1971 was transferred to Anser, with the proviso that it may be the same as the preceding species; and last, the generic assignment of Aythya spatiosa Kurochkin, 1976, based on a distal femur, was accepted. From the Caucasian region between Europe and Asia comes Anas apscheronica Burczak-Abramovicz, 1958 (fide Mlíkovský & Švec 1986).

Several more species have also been named from North America. An extinct goose Anser thompsoni Martin & Mengel, 1980, was described from the Late Pliocene of Nebraska (Martin & Mengel 1980). Short (1970) described a goose-sized tarsometatarsus from the Pliocene of Nebraska as Heterochen pratensis Short, 1970, but was unable to determine its relationships. Emslie (1992) reported a significant Late Blancan (2.5-2.0 Ma) fauna from Florida that included Dendrocygna sp., the extant Branta canadensis, the extinct Anabernicula gracilenta Ross, 1935, five extant species of Anas, two extant Aythya species, one extant Bucephala species and one extant Mergus species. In addition, he described the new genus and species Helonetta brodkorbi Emslie, 1992, which had similarities to Nettapus, and the new species Oxyura hulberti Emslie, 1992. Alvarez (1977) described Oxyura zapatanima Alvarez, 1977 from the Late Pliocene-Early Pleistocene of Mexico. Olson & Rasmussen (2001) reported at least 20 anseriform species from the Yorktown Formation, North Carolina, United States, of Pliocene age (c. 4.8-3.7 Ma). Of these, only one Anabernicula minuscula was extinct, and modern genera not listed by Howard included Somateria, Histrionicus, Melanitta and Mergus. 19

1.4 The New Zealand vertebrate fossil record

New Zealand is the presently emergent part of a continental fragment now known as Zealandia, which was once part of (Campbell & Hutching 2007). It has generally been considered to have existed as an archipelago since its separation from Australia and Antarctica around 82-60 Ma (Fleming 1979; Cooper & Millener 1993; Gaina et al. 1998a, b; Sutherland 1999), but see below for a contrary opinion. It lies on the margin of the Pacific and Australian Plates and reached its present position with respect to Antarctica by about 60 Ma. Australia broke free of Antarctica much later, c. 38 Ma, and continued its unimpeded northward drift into lower latitudes until 15 Ma (Brown et al. 2006).

The Recent indigenous New Zealand terrestrial flora and fauna is highly distinctive with Podocarpus conifers, beech trees, arthropods such as velvet worms (onychophorans) and giant weta (stenopelmatids: Orthoptera), and vertebrates such as primitive leiopelmatid frogs, tuatara (Sphenodon), (Aves: Dinornithiformes), and the New Zealand wrens (Acanthisittidae) often interpreted as relict Gondwanan vicariant taxa (e.g., Fleming 1979; Worthy & Holdaway 2002; Gibbs 2006). The acanthisittid wrens and the (Strigops habroptilus) are now understood as the sister groups to all other and , respectively, and are interpreted as vicariant taxa (Barker et al. 2004; Ericson et al. 2002; Tavares et al. 2006).

Recently, however, some geologists and palaeontologists have controversially suggested New Zealand was completely submerged during the latest Oligocene–earliest Miocene 25-22 Ma, requiring the modern fauna and flora to be derived entirely by long- distance dispersal, presumably mainly from Australia (Pole 1994; Heads 2006; Waters & Craw 2006; Campbell & Hutching 2007; Landis et al. 2008). This hypothesis, however, is not supported by an increasing array of genetic studies that show New Zealand taxa to have diverged from sister taxa elsewhere long before the Oligocene highstand, e.g. kauri (Knapp et al. 2007), hyriid molluscs (Graf & Foighil 2000), freshwater crayfish (Apte et al. 2007), leiopelmatid frogs (Roelants & Bossuyt 2005), tuatara (Rest et al. 2003), the wattlebirds and (Driskell et al. 2007; Shepherd & Lambert 2007), kakapo and kaka (Tavares et al. 2006) and acanthisittid wrens (Ericson et al. 2002). 20

New Zealand, whose main islands lie between 34 and 47ºS, some 1400 km east of Australia, had some 245 breeding bird species and c. 64 skinks and geckoes in the Recent fauna, with known land represented only by three bat species (Worthy & Holdaway 2002). New Zealand’s terrestrial vertebrate fossil record, while very rich in the Late Quaternary (<100,000 years BP), is one of the world’s poorest for the pre-Quaternary (>2 Ma). Except for fragmentary Late Cretaceous (80-71 Ma) (Molnar & Wiffen 1994) and isolated moa bones from marine sediments <2.5 Ma (Worthy et al. 1991; Worthy & Holdaway 2002), the record for terrestrial vertebrates >1 Ma has been non-existent. A small avifauna from 1 Ma shoreline deposits near Marton, , includes several species that are unknown from late Quaternary faunas and at least two Recent moa species (Worthy 1997). New Zealand has a relatively rich Cenozoic record of marine birds, especially (Turbott 1990; Fordyce & Jones 1990; Fordyce 1991), but until recently its Tertiary record of terrestrial and freshwater has been a complete mystery. While endemic taxa of Gondwanan origin noted above attest to a probable vicariant origin to some of the biota, most birds are assumed to have dispersed from Australia (e.g., Falla 1953; Fleming 1962, 1979; Millener 1991). The main questions are when did such lineages arrive in New Zealand and how long have they been there?

With discovery of what is now called the St Bathans Fauna (Worthy et al. 2007) from the Early Miocene Manuherikia Group sediments (19-16 Ma), a first window into the Tertiary origins of the Recent fauna was afforded. In 1978, avian eggshell and anatid bones were found in the basal Bannockburn Formation (Fm) of the Manuherikia Group sediments, near St Bathans in Central Otago, New Zealand (Douglas 1986; Douglas et al. 1981; Fordyce 1991). Other early discoveries, in 1980-1981 by Ewan Fordyce at Vinegar Hill, near St Bathans, included two anatids (Fordyce 1991), presumed to be a duck and a small goose (Fordyce 2003). Until 2000, the only other vertebrate specimen to have been reported was a crocodilian angular (Molnar & Pole 1997), also from the basal Bannockburn Fm in Mata Creek near St Bathans (B. Douglas pers. obs.). The Manuherikia Group sediments have since been the focus of much palaeobotanical work investigating both macrofloras and palynofloras (Pole 1989, 1992a-c, 1993a-g, 1997), culminating in summaries of vegetation and environment (Pole & Douglas 1998; Pole et al. 2003).

The Manuherikia Group sediments were deposited shortly after much of New Zealand re-emerged from an Oligocene marine transgression. Maximum submergence occurred in the Late Oligocene to earliest Miocene 25-22 Ma, when only c.18% of the 21

present land area remained (Cooper & Cooper 1995), but see Campbell & Hutching (2007) and Landis et al. (2008) for a complete submersion hypothesis. I follow Gibbs (2006) in not accepting total drowning of Zealandia, as it is more parsimonious to derive endemic taxa like the numerous discussed by Gibbs, and especially the vertebrates Leiopelma, Sphenodon and the mammal noticed by Worthy et al. (2006), by a vicariant origin in the , than by post-Oligocene dispersal from other landmasses (followed by of close relatives in the source landmass). These three vertebrate taxa have their nearest relatives in the Mesozoic of Gondwana and have no Tertiary relatives. Moreover, there are increasing genetic data that supports pre-drowning divergences of New Zealand taxa from nearest overseas counterparts (cited above). In addition, the fossil record is now being fleshed out for the Late Oligocene-Early Miocene, for example, a record of the extant kauri Agathis australis (Lee et al. 2007).

Given the persistence of some land, the Oligocene marine transgression has been postulated to be a time of evolutionary stress on the fauna, a bottleneck that would have whittled down the diversity of the terrestrial biota (Cooper & Cooper 1995). Mitochondrial DNA from moa, and wrens has provided evidence that there were significant post- Miocene radiations, which may reflect a response to the spread of drier, cooler environments from Middle to Late Miocene, as well as the increase in geographic area for re-colonisation (Baker et al. 1995; Cooper & Cooper 1995; Cooper et al. 2001; Baker et al. 2005). With the onset of renewed tectonism and lowered sea level in the Early Miocene, land area rapidly increased, but it was not until the Late Pliocene, c.2 Ma, that the North and South islands reached their present configuration.

A renewed search for vertebrate faunas began in 2000 and 2001 (Worthy et al. 2002a, b). Exploration was concentrated on sites along the Mata Creek and Manuherikia River where the lower Bannockburn Fm is exposed. The ages and the stratigraphy of the Manuherikia Group have been determined using palynological data (Mildenhall 1989; Mildenhall & Pocknall 1989), and the palynostratigraphic zonation of the terrestrial bone- bearing outcrops given by Pole & Douglas (1998). Terrestrial bones typically occur in the lowest 30 m of the Bannockburn Fm. These sediments were deposited in a large freshwater lake (Lake Manuherikia), which was >5,600 km2 in area (Douglas 1986). A broad fluvial plain with major channels and interchannel flood-basins surrounded the lake. Shallow wetland habitats were vegetated by grasslands, herbfields and peat-forming swamp- woodland, with relatively dry Casuarinaceae woodland occurring nearby (Pole et al. 2003). 22

The St Bathans fauna is now (2008), after eight years of investigation, known to include diverse terrestrial vertebrates in addition to ubiquitous fish (Chapter 1). It represents the only window through which to view the entire Cenozoic history of land vertebrates in the New Zealand Gondwanan fragment. Reptiles: apart from crocodilians, fossils include undetermined gekkonids and skincids, and the first pre-Holocene New Zealand record for tuataras (sphenodontids). There is no evidence for agamids, varanids, iguanids or snakes. Frogs: Three bones include a leiopelmatid vertebra and another vertebra of an unidentified group of frogs. Mammals. The most surprising discovery from the St Bathans Fauna is New Zealand’s first native non-volant terrestrial mammal: a mouse-sized that appears best interpreted as a stem therian more primitive than and placentals but more advanced than (Appendix 1). A minimum of five new bat species in four families with relationships to both South American and Australian faunas has also been identified (Hand et al. 2007). Birds: The St Bathans avifauna is already more diverse than any Tertiary bird fauna from Australia with at least 25 species represented by thousands of bones (Chapters 2-4). Ratites are represented by eggshell fragments, whose thickness attests to these moa ancestors being then large and flightless. Anatids dominate the fauna with six described species in four genera, and a further two undescribed anserines (data herein). This is the most diverse Early Miocene duck fauna known from anywhere in the world. A diving petrel (Pelecanoides) extends the geological range of the genus globally from the Pliocene to the Middle Miocene. The remaining 16 bird taxa include gulls and (charadriiforms), a large gruiform, terrestrial rails, an eagle, a pigeon, an owlet-nightjar (Aegotheles), a (Collocalia), three parrots and several passerines including one similar to a currawong (Cracticidae).

The continued excavation of this fauna will undoubtedly enlarge the species diversity and improve the species representation of water birds in particular. This is especially so when it is appreciated that the European and American faunas have taken often many decades of work to develop and 150 years in some instances, whereas the St Bathans Fauna derives essentially from eight seasons’ work.

1.5 The Australian avian fossil record

The Tertiary fossil record of non-marine Australian avifaunas is relatively rich with the earliest being the Tingamara LF of Eocene age in Queensland (Godthelp et al. 1992; Boles 1995a, 1997a). Most faunas however, derive from Late Oligocene and younger 23

deposits as reviewed by Vickers-Rich (1991). A significant source of Australian Tertiary birds is the Oligo-Pliocene record of inland South Australia (Rich 1976; Stirton et al. 1961, 1967; Woodburne et al. 1994), particularly as it is exposed about Lakes Palankarinna, Pinpa, Ngapakaldi, and Yanda (Rich & Van Tets 1982). The Pliocene is represented by faunas in the Tirari Formation derived from the sediments at Lake Palankarinna (Mampuwordu Member, Palankarinna Fauna) and Lake Kanunka (Kanunka Fauna) (Tedford et al. 1986; Tedford & Wells 1990; Tedford et al. 1992).

The mihirung birds (Aves: Dromornithidae) are basal neornithines related to anseriforms and are well-known and widespread members of the Tertiary avifaunas (e.g. Rich 1976, 1979; Murray & Vickers-Rich 2004). Other prominent components of these avifaunas are and palaelodids (Miller 1963; Rich 1976; Rich et al. 1987; Baird & Vickers-Rich 1998) and pelicans (Rich & van Tets 1981). Since Vickers-Rich’s (1991) review, there have been notable contributions from the South Australian deposits, for example, ratites (Boles 1992) and (Boles & Ivison 1999). But the greatest advances in Australian palaeornithology derive from the large faunas obtained from numerous deposits at Riversleigh in northwestern Queensland, which have produced a diverse array of fossil birds (Boles 1993a-c, 1995b, 1997b-d, 1998, 1999, 2001, 2005a-c).

Waterfowl (Aves: Anseriformes) are known from various Australian Tertiary fossil faunas (e.g., Tedford et al. 1977; Rich & van Tets 1982; Pledge 1984; Pledge & Tedford 1990; Tedford & Wells 1990; Vickers-Rich 1991; Rich et al. 1991; Tedford et al. 1992; Boles & Mackness 1994; Boles 1997c), but before this project, the identity and phylogenetic relationships of these fossils had not been assessed.

The oldest anseriform fossils in Australia are known from the Late Oligocene– Miocene lacustrine deposits of the Lake Eyre and Tarkarooloo basins of central Australia (Vickers-Rich 1991). Anseriform fossils are relatively common in Pliocene-Pleistocene deposits (Tedford & Wells 1990; Tedford et al. 1992), but all nine extinct taxa named by de Vis have been referred to modern taxa by Olson (1977b), and no other extinct anseriforms have been described from Australia. No extant taxa are known from pre-Pliocene deposits (Vickers-Rich 1991). This is despite taxa such as Anseranas (Anseranatidae), the dendrocygnines, the anserines Cereopsis and Cnemiornis, and Stictonetta, all being the most primitive members of the Recent fauna (Livezey 1986, 1989a, 1996a, 1997a; Worthy et al. 1997) and, therefore, presumed to have had a long history in the region. As noted above, globally most modern genera evolved between the Middle Miocene and the Pliocene, so 24

determination of the composition of Australasian anseriform faunas in the Oligocene- Pliocene period will reveal much about the evolution of the modern fauna. Moreover, analyses of Australasian taxa will test hypotheses that some taxa evolved in the south and spread northwards into Asia and Europe, as postulated by Heizmann & Hesse (1995) and Cracraft (2001).

1.6 Morphological character analysis

Other than for the most well preserved and very recent fossils, determination of phylogenetic relationships is necessarily reliant on morphological analyses. Taxonomists have long based avian on skeletal characters, at least in part (e.g., Eyton 1838; 1867, 1869, 1873-75; Huxley 1867; Meyer 1879-1897; Fürbringer 1888; Gadow 1892; Beddard 1898; Shufeldt 1909; Verheyen 1955a-d), and detailed osteological descriptions of anatids, both fossil and recent, are abundant (e.g. Owen 1866; Milne-Edwards 1863, 1867- 71; Shufeldt 1913, 1914; Raikow 1971). Woolfenden (1961), in a highly regarded and now classic osteological study, synthesised most available data and characterised all anseriform Recent genera osteologically. The characters Woolfenden described have often been used in the characterisation of fossil taxa (e.g. Short 1970; Alvarez & Olson 1978; Bickart 1990; Olson & James 1991; Emslie 1992; Olson 1999; Louchart 2005).

Following the development of cladistic phylogenetic methods of analysis (Hennig 1966; Wiley 1981), the characters identified by Woolfenden (1961) formed the basis for the skeletal characters used in the extensive studies of Livezey (1986, 1989ab, 1990, 1991, 1995ab, 1996a-c, 1997a-c), Ericson (1997), and some in Livezey & Zusi (2007). Characters are homologous morphological features that vary consistently between taxa among which several states may be observed. A character has a primitive state (plesiomorphic) and one or more derived (apomorphic) states, which may or may not form a linear sequence. The plesiomorphic state is determined as the state in the outgroup taxon and often (but not always) in the more basal taxa.

A major problem of cladistic phylogenetic analyses is that of homoplasy, where taxa acquire similar character states owing to convergence of form, usually related to a common functional requirement, rather than their being due to a shared derived origin; for example, large eyes in nocturnal birds is often a homoplasous character. It is well known that 25

homoplasy related to paedomorphosis can mislead interpretations of phylogeny (e.g. Wiens et al. 2005).

Problems of homoplasy are evident in diverse avian groups for which conclusions derived from phylogenetic analyses of morphological datasets using skeletal characters are compared with genetic analyses. For example, Livezey (1998) found that flightlessness- related homoplasy resulted in the improbable association of several large, flightless taxa of rails. McCracken & Sheldon (1998) analysed skeletal and genetic datasets for and concluded that use of characters based on highly adaptive morphology, particularly the bill, suffered from homoplasy. In studies of accipitrid relationships, Griffiths et al. (2007) found little congruence between those recovered in the morphological analysis of Holdaway (1994) and those from genetic data sets. Moreover, the studies by Griffith et al. (2007) and others (e.g. Helbig et al. 2005) have revealed extensive non-monophyly at the generic level in accipitrids, showing that homoplasy in morphology has widely misled previous phylogenetic interpretations.

Homoplasy has affected phylogenetic analyses of anatids. For example, Livezey (1996a) found the flightless, goose-sized moa-nalos of Hawaii to be anserines, contra their referral by Olson & James (1991) to Anatines based on the presence of a syringeal bulla. Genetic evidence clearly supported a phylogenetic relationship of moa-nalos with Anatinae (Sorenson et al. 1999), contra Callaghan & Harshman (2005), strongly suggesting that homoplasy related to large size affected the morphological analysis. Another example concerns the affinity of Oxyura and Biziura. Livezey’s (1995a, 1997a) analyses associated the diving taxa Aythyini, and in a linear sequence after Anas in a phylogenetic hypothesis at odds with those based on genetic data (e.g. Madsen et al. 1988; Sibley & Ahlquist 1990; Sraml et al. 1996; Johnson & Sorenson 1999; Sorenson et al. 1999; Donne-Goussé et al. 2002). In new combined morphological and genetic analysis, it was shown that the association of Oxyura with Aythyini and Mergini was related to problems of homoplasy arising from of diving adaptive structures (McCracken et al. 1999). Thus, it is clear that any analysis based on skeletal characters is dependent on objective definition of characters and clear identification of problems associated with homoplasy, particularly owing to flightlessness and diving.

In this thesis, I necessarily make extensive use of morphological characters to define taxa and to determine their phylogenetic relationships. I have used characters described by Livezey (1986, 1989a,b, 1990, 1991, 1995a,b, 1996a-c, 1997a-c) as a basis from which to 26

define those used here. Other characters have been sourced from those described in Raikow (1971), Ericson (1997), and in my own previous studies of waterfowl (Worthy 1988, 1995, 2002, 2004, 2005; Worthy & Gill 2002; Worthy & Olson 2002; Worthy et al. 1997). In an attempt to maximise detection of phylogenetic signal in datasets, all definitions have been checked and, where necessary, revised following extensive comparisons of taxa to determine the extent of variability and homology between taxa, and to ensure functional complexes are not being confused within a single character. Characters have been chosen and many new ones defined to maximise information recovery from the available fossils. All taxa are coded following original observation of actual or figured specimens. Further, methods of analysis are advanced in the relevant chapters to mitigate potential problems of homoplasy.

1.7 The aims of the thesis

The overall objective of this thesis is to determine the taxonomic diversity of the fossil waterfowl and the systematic relationships of the taxa with adequate fossil representation for phylogenetic analyses in the Tertiary faunas of New Zealand and Australia. The project is divided into a number of aims based on the source and the geological age of the available fossils.

Aim 1. Description and phylogenetic interpretation of the waterfowl from the Early Miocene St Bathans Fauna from Otago, New Zealand. The phylogenetic relationships of the three species/genera that have the most complete skeletal representation will be determined by analyses of morphological data, in comparisons including an extensive range of modern anseriforms and selected fossil taxa.

Aim 2. Description and phylogenetic interpretation of the waterfowl from the Oligocene-Miocene of Australia. The Australian fossil avifaunas can be conveniently divided into Oligocene-Miocene and Pliocene components on account of the major gap in fossil representation between the Middle Miocene and the Pliocene, noted above. Where taxa are adequately known, phylogenetic analyses will be undertaken to augment taxon descriptions and determine their relationships to Oligocene-Miocene taxa in New Zealand and elsewhere in the world.

Aim 3. Description and phylogenetic interpretation of the waterfowl from the Pliocene of Australia. 27

Aim 4. The identification of the extent and timing of faunal turnover and the origin of the modern fauna.

Aim 5. To test, using a morphological dataset, the phylogenetic hypothesis of anseriform generic interrelationships advanced by Livezey (1997a). There is as yet no comprehensive genetic-based analysis of anatid generic relationships, thus the conclusions of Livezey (1997a) have been widely adopted (e.g. Callaghan & Harshman 2005), despite the contradictory evidence noted in less taxonomically-sampled genetic analyses (e.g. Sraml et al. 1996; Johnson & Sorenson 1999; McCracken et al. 1999; Sorenson et al. 1999; Donne-Goussé et al. 2002; McCracken & Sorenson 2005). Chapters 3 and 5, while primarily to ascertain the phylogenetic relationships of fossil taxa, develop a new hypothesis of anatid generic relationships.

1.8 Thesis structure This thesis is presented as a series of seven papers that are either published, in press, or submitted for publication. As such, they are presented in journal format, preceded by a thesis title page and, where necessary, a statement of authorship. The thesis concludes with a discussion summarizing the outcomes from this project and their significance.

The appendices include two papers based on research done concomitantly with that for this thesis and which are related to the research presented herein. Appendix 1 is a paper published in Proceeding of the National Academy of Sciences that describes the first terrestrial mammal known from New Zealand, from fossils derived from the St Bathans Fauna. They represent a stem mammal ancestral to both therians and metatherians, but which is more derived than monotremes. These fossils have great significance as they reveal a previously unknown lineage of mammals, whose presence in New Zealand, along with frogs and sphenodontids, is best explained as by vicariance, rather than by dispersal since the Oligocene marine highstand.

Appendix 2 is a paper that is in press in the Bulletin of the British Ornithologists’ Club reassessing the taxonomic identity of Anser scaldii. This is significant as this poorly known fossil was formerly interpreted as a Middle Miocene form: in the event, it is a Pleistocene fossil of a Recent species. It, therefore, has no significance in interpreting anseriform evolution. 28

29 30 31 32 33 34

Journal of Systematic Palaeontology 5 (1): 1–39 Issued 1 March 2007 doi:10.1017/S1477201906001957 Printed in the United Kingdom C The Natural History Museum Miocene waterfowl and other birds from central Otago, New Zealand

T. H. Worthy∗ Darling Building, DP 418, Dept. of Earth and Environmental Sciences, The University of Adelaide, SA 5005, Australia

A. J. D. Tennyson Museum of New Zealand Te Papa Tongarewa, P.O. Box 467, , New Zealand

C. Jones Institute of Geological and Nuclear Sciences, P.O. Box 30368, Lower Hutt, New Zealand

J. A. McNamara South Australian Museum, North Terrace, Adelaide, Australia

B. J. Douglas 14 Jubilee Street, , New Zealand

SYNOPSIS Abundant fossil bird bones from the lower Bannockburn Formation, Manuherikia Group, an Early-Middle Miocene lacustrine deposit, 16–19 Ma, from Otago in New Zealand, reveal the “St Bathans Fauna” (new name), a first Tertiary avifauna of land and freshwater birds from New Zealand. At least 23 species of birds are represented by bones, and probable moa, Aves: Dinornithiformes, by eggshell. Anatids dominate the fauna with four genera and five species described as new: a sixth and largest anatid species is represented by just one bone. This is the most diverse Early-Middle Miocene duck fauna known worldwide. Among ducks, two species of dendrochenines are most numerous in the fauna, but a tadornine is common as well. A diving petrel (Pelecanoididae: Pelecanoides) is described, so extending the geological range of this genus worldwide from the Pliocene to the Middle Miocene, at least. The remaining 16 taxa are left undescribed but include: a large species of gull (Laridae); two small waders (Charadriiformes, genus indet.), the size of Charadrius bicinctus and Calidris ruficollis, respectively; a gruiform represented by one specimen similar to Aptornis; abundant (Rallidae) bones, including a common flightless rail and a rarer slightly larger taxon, about the size of Gallirallus philippensis; an ?eagle (Accipitridae); a pigeon (); three parrots (Psittacidae); an owlet nightjar (Aegothelidae: Aegotheles sp.); a swiftlet (Apodidae: Collocalia sp.); and three taxa, of which the largest is a member of the Cracticidae. The absence of some waterbirds, such as anserines (including swans), (Podicipedidae) and shags (Phalacrocoracidae), among the abundant bones, indicates their probable absence from New Zealand in the Early-Middle Miocene.

KEY WORDS Avifauna, fossils, new taxa, Anatidae, lacustrine, Early-Middle Miocene, New Zealand A Worthy, T.H., Tennyson, A.J.D., Jones, C., McNamara, J.A. & Douglas, B.J. (2007) Miocene waterfowl and other birds from Central Otago, New Zealand. Contents Journal of Systematic Palaeontology, v. 5(1), pp. 1-39 Introduction 3 Geological and palaeoenvironmental setting 4 A Methods 4 AbbreviationsNOTE: 4 ThisInstitutions publication is included on pages 34-72 in the print copy 4 Skeletalof the elementsthesis held and descriptivein the University terms used of Adelaide Library. 4 MeasurementsA 4 ComparativeIt is materialalso available online to authorised users at: 6 Taxonomic issues A 7 Taphonomy http://dx.doi.org/10.1017/S1477201906001957 7 * Corresponding author. E-mail address: [email protected]. A 73 74 75

[Palaeontology, Vol. 51, Part 3, 2008, pp. 677–708]

AFFINITIES OF MIOCENE WATERFOWL (ANATIDAE: MANUHERIKIA, DUNSTANETTA AND MIOTADORNA) FROM THE ST BATHANS FAUNA, NEW ZEALAND by TREVOR H. WORTHY* and MICHAEL S. Y. LEE* *Darling Building, DP 418, Department of Earth and Environmental Sciences, The University of Adelaide, Adelaide, SA 5005, Australia; e-mails: [email protected], [email protected] Earth Sciences Section, South Australian Museum, North Terrace, Adelaide, SA 5000

Typescript received 2 January 2007; accepted in revised form 19 June 2007

Abstract: The recently described St Bathans Fauna, from associated with this oxyurine clade or formed separate lin- the Manuherikia Group, Early–Middle Miocene, 19–16 Ma, eages of an approximately oxyurine , New Zealand, includes six anatid taxa. Here we present depending on whether diving characters were included or detailed morphological descriptions of all available skeletal excluded. Similarly, Biziura, Thalassornis, and a Stictonetta– elements of the three best represented anatids: Manuherikia Malacorhynchus clade either associated with oxyurines or lacustrina (551 specimens), Dunstanetta johnstoneorum (7 formed independent lineages of approximately oxyurine specimens), and Miotadorna sanctibathansi (115 specimens). grade. Above oxyurines, a well-supported clade groups Mio- The affinities of these taxa, and of the similar-aged European tadorna with Tadorna species exclusive of T. radjah. These taxon Mionetta blanchardi, are evaluated with phylogenetic results cast doubt on the distinctiveness of Dendrocheninae analyses using a dataset of 133 characters (128 osteological, 5 Livezey and Martin, 1988, and instead suggest oxyurine affin- integumental) and 57 terminal taxa. Representatives of all ities for the Miocene fossils Mionetta, Dendrochen, Manuheri- main anatid groups were included, with dense sampling of kia and Dunstanetta, and the modern Biziura, Thalassornis, Australasian taxa including the recently extinct New Zealand Oxyura, Nomonyx, Stictonetta and Malacorhynchus. The forms (Cnemiornis, Chenonetta finschi), and relatively primi- association of Mionetta with oxyurines indicates that the tive taxa (anserines, Dendrocygna, oxyurines, tadornines). divergence between oxyurines and higher anatids occurred Analyses were conducted with no constraints, and with cer- around 25 Ma, while the position of Miotadorna within tain taxa constrained to conform to arrangements supported Tadorna indicates that the basal divergence between living by multiple independent genetic studies. In the preferred Tadorna occurred by the Early–Middle Miocene; together, (constrained) analyses: (1) anserines were the most basal these two dates indicate that many basal splits within anatids anatids; (2) the four Tertiary fossil taxa were more derived occurred within a short interval during the Miocene. than anserines and Dendrocygna; and (3) the European Mionetta blanchardi was basal to an oxyurine clade. The Key words: New Zealand, Neogene waterfowl, Manuherikia, New Zealand fossils Manuherikia and Dunstanetta either Dunstanetta, Miotadorna, Mionetta, phylogenetic analysis. A AWorthy,nseriformes T.H. & Lee,comprise M.S.Y. the (2008) extant Affinities Anhimidae of Mioceneclassical study, waterfowl synthesised (Anatidae: most available Manuherikia, data and char- (SouthDunstanetta American and screamers), Miotadorna Anseranatidae) from the (Australian St Bathans acterisedFauna, New osteologically Zealand. all Recent genera of anseriforms. magpiePalaeontology, geese) and truev. 51(3), waterfowl pp. Anatidae677-708 (cosmopolitan The characters he described have often been used in the swans, geese and ducks), as listed in Delacour and Mayr diagnosis of fossil taxa (e.g. Short 1970; Alvarez and Ol- (1945), Dickinson (2003) and Kear (2005). Phylogenetic son 1978; Bickart 1990; Olson and James 1991; Emslie relationships withinA the order have been determined from 1992; Olson 1999; Louchart et al. 2005). Following the diverse data sets, but avian taxonomists have long reliedNOTE:development of cladistic methodology (Hennig 1966; heavily on skeletal charactersThis publication (e.g. Eyton is 1838,included 1867, on pagesWiley 1981)75-106 these in skeletalthe print characters copy formed the basis of 1869, 1873–75; Huxley 1867;of the Meyer thesis 1879–97; held Fu in¨rbringer the Universitythe extensive of Adelaide studies ofLibrary. Livezey (1986, 1989a, b, 1990, 1888; Gadow 1892; Beddard 1898; Shufeldt 1909; Verh- 1991, 1995a, b, 1996a–c, 1997a–c) and Ericson (1997). eyen 1955a–d), and detailed osteological descriptions of A Livezey’s (1997a) phylogenetic conclusions, based on fossil and recent anatids are abundantIt is also (e.g. available Owen 1866; onlineanalyses to authorised of both skeletalusers at: and integumental characters for Milne-Edwards 1863, 1867–71; Shufeldt 1913, 1914; Rai- A modern anatids, differ in many ways from traditional kow 1971). Woolfenden (1961),http://dx.doi.org/10.1111/j.1475-4983.2008.00778.x in a highly regarded, views and are also at odds with phylogenies based on A

ª The Palaeontological Association doi: 10.1111/j.1475-4983.2008.00778.x 677 107 1 SUPPLEMENTARY MATERIAL Affinities of Miocene waterfowl (Anatidae: Manuherikia, Dunstanetta and Miotadorna) from the St Bathans Fauna, New Zealand. Palaeontology (2008), doi: 10.1111/j.1475-4983.2008.00778.x.

Comparative material The following comparative material was consulted during this study. All are extant species unless stated otherwise; sex (M or F), is indicated after the catalogue number of the specimen. Anhima cornuta (Linnaeus, 1766), horned : MV B12574. Anseranas semipalmata (Latham, 1798), magpie goose: AM O59362; SAM B48035, M; B39824, F. Cereopsis novaehollandiae Latham, 1802, Cape Barren goose: MNZ 25217, 25143; SAM B39638, B32830. Cnemiornis calcitrans Owen, 1866, goose: fossil, notably MNZ S.35266; MV P160495. Cnemiornis gracilis Forbes, 1892, North Island goose: fossil, notably MNZ S.35683-706; MNZ S.35892. Branta canadensis (Linnaeus, 1758), Canada goose: MNZ 23745, 26738, 26739, 26740, 26741; SAM B31086, MV B6364, M. Anser brachyrhynchus Baillon, 1834, pink-footed goose: MV B25672. Anser caerulescens (Linnaeus, 1758), snow goose: SAM B36868 F. Anser anser (Linnaeus, 1758), greylag goose: MNZ 20812, 24519. Cygnus atratus (Latham, 1790), black swan: MNZ 15266, 15267, 17250; SAM B46110, M. Cygnus olor (Gmelin, 1789), mute swan: MNZ 16454. Dendrocygna arcuata (Horsfield, 1824), wandering : AM O64697, M, O64697, M; ANWC 22213, M; ANWC 22214, M. Dendrocygna eytoni (Eyton, 1838), plumed whistling duck; MNZ 27024; SAM B45769, M. Dendrocygna bicolor, (Vieillot, 1816), fulvous whistling duck: SAM B36869; MV B13722. Dendrocygna arborea, (Linnaeus, 1758), West Indian whistling duck: USNM 226455, F. USNM 344488, F. Dendrocygna autumnalis (Linnaeus, 1758), black-bellied whistling duck: USNM 345767, M; USNM 345768, F (both are D. a. discolor). Thalassornis leuconotus Eyton, 1838, white-backed duck: BMNH 1901.10.20.156, F. Oxyura australis Gould, 1836, blue- billed duck: CM Av31408; AM O65518; SAM B49214, M, B36390, F, B31910, F, B32842, ?. Nomonyx (Oxyura) dominicus, (Linnaeus, 1766), : USNM 430927, F; USNM 347324, M. Oxyura vittata, (Philippi, 1860), Argentine ruddy duck: USNM 614580, M. Oxyura maccoa, (Eyton, 1838), : USNM 558442, M. Oxyura jamaicensis (Gmelin, 1789), ruddy duck: ANWC 22640 (ANSS 386), ANWC 22641 (ANSS 391); MNZ 27335; SAM B36872, F. Oxyura vantetsi Worthy, 2005, New Zealand blue-billed duck: Specimens (fossil) listed in Worthy (2004, 2005). Biziura lobata 108 2 (Shaw, 1796), : MNZ 26190, 26191, CM Av7116; SAM B38592, M, B47221, M, B11405, M, B46027, F, B49170, F, B36393, F, B31087, F. Stictonetta naevosa (Gould, 1841), freckled duck: MNZ 25141; SAM B46743, M, B47950, M, B39659, M, B31585, F, B45045, F. Malacorhynchus membranaceus (Latham, 1802), Australian pink-eared duck: MNZ 23880, 23881; SAM B39384, M, B39385, F. Malacorhynchus scarletti Olson, 1977, Scarlett’s duck: fossil, MNZ, Lake Poukawa specimens listed in Worthy (2004); CM Av5855, Av14697, Av15044, Av36035, Av36285. Nettapus pulchellus (Gould, 1842), green pygmy-goose: MNZ 27025, 27026; SAM B45606; MV B14485, M, B14490, M. N. coromandelianus (Gmelin, 1789), cotton pygmy-goose: MV B20293, F; QM O.21080, QM O.21081. Tadorna ferriginea (Pallas, 1764), : SAM B38602. Tadorna tadornoides (Jardine & Selby, 1828), : MNZ 22921, 23888a, 27367; ANWC 22240; SAM B39873, F, B39591, M. Tadorna tadorna (Linnaeus, 1758) : MNZ 12280; MV B25679; ANWC 22408, M. Tadorna variegata (Gmelin, 1789), : MNZ 15146, 16471, 16472, 16473, 16501, 16590, 24559, 25139, 25669, 26562, 26563; MV B5103, F. Tadorna radjah (Lesson, 1828), : MNZ 26206, 26207; ANWC 22411, M. Alopochen aegyptiacus (Linnaeus, 1766), African : ANWC 22239 (ANSS 753), M; BMNH 1930.3.24.217, ?; MV B25678; MNZ 24283. Chloephaga poliocephala Sclater, 1857, ashy-headed sheldgoose: MV B13714, F. Chloephaga hybrida (Molina, 1782), sheldgoose: MV B13227, M. Chloephaga picta (Gmelin, 1789), upland sheldgoose: BMNH 1860.11.4.15. Tachyeres leucocephalus Humphrey and Thompson, 1981, white-headed flightless : MV B14144, M. Cairina moschata (Linnaeus, 1758), ; MNZ 19842; SAM B32415. Chenonetta jubata (Latham, 1802), maned duck: MNZ 1487, 23188a, 25142, 25400, 25194a; SAM B37195, M, B39011, F. Chenonetta finschi (Van Beneden, 1875), Finsch’s duck: fossil, MNZ specimens, notably S.35885, S.23794, S.35885, S.38931, S.39838, S.39840, S.44279; MV P204243. Hymenolaimus malacorhynchos (Gmelin, 1789), blue duck: MNZ 16699, 23924, 23963; 24586, 24587. Somateria mollissima (Linnaeus, 1758), common : MNZ 12277, 12278, 12279; SAM B48744, F. Melanitta fusca (Linnaeus, 1758), velvet : SAM B47234, M. Clangula hyemalis (Linnaeus, 1758), long-tailed duck: SAM B47233, M. Bucephala albeola (Linnaeus, 1758), : MNZ 12708, 12709. Bucephala clangula (Linnaeus, 1758), common : MV B13553, M. Lophydytes cucullatus (Linnaeus, 1758), hooded merganser: MNZ 12706; MV B13541, M, SAM B47750, M. Mergus merganser Linnaeus, 1758, common merganser: MV B13320, M. Mergus serrator Linnaeus, 1758, red-breasted merganser: MNZ 12707; MV B14546, M. Mergus australis Hombron & Jacquinot, 1841, 109 3 Island merganser: MNZ S30046, S31777, collections of fossils from Chatham Island. Anas superciliosa Gmelin, 1789, gray duck: MNZ 13686, 15030, 16586, 18132, 16476, 17341, 17261, 16698, 16584, 24607; SAM B49654, M. Anas rhynchotis variegata (Gould, 1856), Australasian : MNZ 17000, 18971, 16591, 24588, 24589; SAM B37429, M. Anas castanea (Eyton, 1838), chestnut teal: SAM B32512, M; AM O.67041, M. Anas chlorotis G. R. Gray, 1845, brown teal: MNZ 14978, 15628, 15935 (= CM Av31828), 18898, 21544, 22086, 22802, 22806, 24535, 24536, 24537, 25105, 25106, 26630, 26631, 26940a, 26941a, 26942a, 26943a, 26944a, 26945a, 26946, 26947, 26949, 26950a, 26951a, 26952a. Anas aucklandica (Gray, 1849), Auckland Island teal: MNZ 24367, 24052. Anas nesiotis (Fleming, 1935), Campbell Island teal: MNZ 25727, 26742. Anas gracilis Buller, 1869, Australasian gray teal: MNZ 19351, 19348, 19301, 19322, 13688, 19323, 26815 (ex 24545), 18099, 19324, CM Av36764. galericulata (Linnaeus, 1758), Mandarin duck: MNZ 27368; SAM B47841, M. Aythya fuligula (Linnaeus, 1758), tufted scaup: MV B12458. Aythya australis (Eyton, 1838), Australian white-eyed duck: AM O65772; SAM B32513. Aythya novaeseelandiae (Gmelin, 1789), : CM Av22382, Av22413; MNZ 8726, 13685, 16588, 16589, 17001, 17002, 17003, 23144, 24245; MV B19895, M. Aythya affinis (Eyton, 1838), : MNZ 24041; MV B25696, M. rufina (Pallas, 1773), red-crested pochard: MV B13723, M.

Mionetta blanchardi (Milne-Edwards, 1863) fossil: Université Claude Bernard - Lyon, ex St. Gérand-le-Puy, France, all prefixed with FSL: 442.807 pmx 442.800 cran L hum 332.274, R hum 332.277, R ulna 331.783, R ulna 331.788, L cmc 331.497, R cmc 331.492, L cor 331.364, R cor 331.381, L fem 331.176, R fem 331.185, L tib 331.104, R tib 331.109, L tmt 331.005, R tmt 331.007; CM Av11394, 1 ant stern, pt cran, fur, LR scap, pt 2 pel, LR MII.1, 2L1R cmc, R rad, LdR ulna, 1L2R cor, 1L2R hum, LR tmt, 1L3R tt, 1L2R fem; CM Av13902, 2L6R cor, 1R3L hum, 2L1R cmc, pt fur.

110 4 Appendix 1. Osteological characters used in the analysis. Characters marked with a star were treated as ordered in the analysis.

SKULL 1. Facies arcus suborbitalis, fused os lacrimale and proc. postorbitalis below orbit: 0, absent; 1, present, typical. (=Character 1, Livezey (1996a)). Livezey (1995a) stated that fusion is a synapomorphy of Dendrocygna, however, fusion was not observed in D. eytoni, nor in D. arcuata, in any of several examples of each taxon. Cnemiornis calcitrans MNZ S.35266 has near fusion of the post-orbital process and lacrimal, but it is effected by an ossified ligamental extension of the lacrimal and is a feature likely associated with old age in the individual.

2*. Facies, pila supranasalis (dorsal border of nares) dorsal convexity: 0, absent; 1, present, not pronounced; 2, present pronounced. (=Livezey (1996a): character 2; Livezey (1986): character 19, revised).

3. Facies apertura nasalis: 0, length approximately equal to width craniofacial hinge; 1, long, length greater than width craniofacial hinge; 2, short, length much less than width craniofacial hinge. (Not, Livezey (1996a): character 3, which related to highly autapomorphic taxa).

4. Os prefrontale (=lacrimals), synostosis with os frontale: 0, lacking; 1, present. (=Livezey (1996a): character 6; = Livezey (1986): character 10). In the Chloephaga specimens examined only the posterior part of joint was fused; in Tadorna this synostosis is the last cranial joint to fuse. The Bucephala examined (MV B13553) was subadult as the gonads were undeveloped and the frontals were not fully synostosed with each other or the parietals, thus lack of synostosis of the lacrimals and frontals was assumed to be a result of immaturity.

5. Os prefrontale, proc. supraorbitalis (supraorbital process): 0, process absent or small, variably laterally oriented; 1, large, flat (dorsoventrally thin), laterocaudally directed with notch caudally (e.g. Malacorhynchus membranaceus); 2, large, thick, dorsolaterally directed with notch caudally (e.g. Branta); 3, long, slender, dorsally directed with notch caudally (e.g. Clangula). Modified from Livezey (1996a): character 7; Livezey (1986): character 11 part.

The juvenile structure of Cereopsis, e.g. SAM B32830, shows that the lacrimal forms the solid preorbital process laterad of the salt gland impressions, so this homology allows Cereopsis to be coded as ‘2’. In Netta MV B13723, the salt gland impressions are excavated at the posterior margin of the supraorbital process accentuating their size: given this allowance it is considered the 111 5 supraorbital processes would be of similar size to those in Anas superciliosa and so are coded as ‘0’.

6. Ossa nasale et frontale, immediately caudad to zona elastica craniofacialis, rounded pneumatic dorsal swellings; Anhimidae and Anseranas treated as missing data because of unique ornament. 0, absent; 1, present, variably prominent, especially medially (Cereopsis, Anser cygnoides, Cygnus olor extreme, Branta less pronounced). =Livezey (1996a: character 8); Livezey (1986: character 16), revised).

Livezey (1996a) identifies this character as intraspecifically variable, sometimes manifested only by bilateral swellings with a shallow medial sulcus, but which may be particularly deep in adult males, and extreme in domestic varieties of Anser.

7. Os frontale, facies dorsalis, sulcus glandulae nasalis (salt gland impressions): 0, absent; 1, present, typically lateral, (Branta sandvicensis and B. hylobadistes vestigial. =Livezey (1996a: character 9 part).

Livezey (1996a) described a separate character state, where the gland is present but enclosed dorsally by an osseous lamina, for the extinct Hawaiian anatids Thambetochen chauliodous and T. xanion and coded Cnemiornis similarly. This is incorrect: the gland is situated dorsally on the cranium as in Cereopsis.

8. Occipital region, fontanelles: 0, absent; 1, present; 2, present or secondarily closed. =Livezey (1986: character 9).

Some individuals, especially males tend to be more heavily ossified and fontanelles closed over, e.g. in Cygnus atratus SAM B48352, juvenile, has fontanelles but all adults examined (SAM) have none, so taxon coded ‘2’. Anhimidae typically loose fontanelles in adulthood (Ericson 1997). Cnemiornis calcitrans normally has no evidence of fontanelles but MNZ S.33863 has small fontanelles and most specimens of C. gracilis have small, partially occluded, fontanelles, so both taxa were coded ‘2’.

9*. Facies interorbitalis, depression frontalis: 0, absent; 1, shallow, elongate groove, restricted to area between lacrimals; 2, shallow elongate groove between lacrimals extending through interorbital area; 3, marked concavity extending from between lacrimals through interorbital area.

Anseranas treated as missing data because of unique ornament. Similarly the combination of deep salt gland impressions and prominent nasal swellings do not allow coding for Melanitta fusca and 112 6 Bucephala clangula. Clangula was coded as ‘1’ as the inter-lacrimal depression had markedly shallowed in the caudal direction before meeting the salt gland impressions, although these obscure the interorbital zone. Aix is coded as ‘1’ because, while the elevated rims of the orbits create a concave frontal surface, this is not homologous to a medial groove extending from the inter- lacrimal region.

10. Os prefrontale (lacrimal), ventrocaudal process: 0, short (in dorsoventral plane), tapering markedly ventrally (e.g. Anseranas); 1, elongate, narrow (e.g. Aythya australis, including here Dendrocygna sp. with lacrimals fused to postorbital process, as lacrimal process narrow before linking to postorbital process); 2, elongate, very broad over much of craniocaudal length, thin (lateromedially), tapering ventrally (e.g. Oxyura, Stictonetta, Malacorhynchus); 3, elongate, narrow, marked distal expansion; 4, elongate and robust, with distal expansion (e.g. Branta, Tadorna).

11*. Os prefrontale (lacrimal) – os ectethmoidale (ectethmoid) complex: 0, Ectethmoid not ossified, or just an incipient ridge dorsally below the nasals; 1, Ectethmoid ossified, small, not fused to lacrimal; 2, Ectethmoid ossified, large, fused to or abuts lacrimal forming lacrimal- ectethmoid complex (e.g. Malacorhynchus).

12*. Recessus tympanicus (=tympanic cavity) bound ventrolaterally by distinct bony flange formed from the ala parasphenoidalis, which is fused medially with the lamina parasphenoidalis and joined caudally to the proc. paroccipitalis of os exoccipitale: 0, not so; 1, yes and dorsal surface of ala parasphenoidalis forms a latero-ventrally sloping floor to cavity (e.g. Dendrocygna); 2, yes, forming flat or concave floor in mediolateral plane (e.g. Anas).

Note: Dendrocygna autumnalis discolor (USNM 345768), uniquely among all Dendrocygna examined, has the dorsal surface of the ala parasphenoidalis forming a concave floor, so this taxon was coded ‘1&2’.

13. Facies articularis frontonasalis (= lacrimal articulation), length relative to anterocaudal orbit diameter: 0, shorter than or equal to; 1, longer than.

14. Os premaxillare (= rostrum maxillare): 0, sides convergent; 1, broad to spathulate usually with sub-parallel sides or slightly divergent;

15. Os premaxillare (= rostrum maxillare) when with sub-parallel sides or slightly divergent: 0, broad to spathulate; 1, narrow (typified by Mergus).

113 7 16. Os premaxillare, anterior median terminus or nail (only ossified structures considered here): 0, pronounced ventral curvature of nail or narrow medial section of tip, below adjacent ventral margins (e.g. Malacorhynchus, Oxyura); 1, pronounced ventral extension of wide section of tip below rest of ventral margin (e.g. Biziura, some Tadorna); 2, no ventral curvature of tip. =Livezey (1996a: character 10), modified; Livezey (1986: character 12), modified.

Aythya has a bill that, in dorsal view, widens towards the tip, and in lateral view has a ventral margin which is straight over the posterior ¾ of its length and which rises anteriorly to level out at the tip. Alternatively, the premaxilla could be described as having a tip with a flat ventral margin, immediately posterad of which are ventrally down-curved flanges, a consequence of the extra width of the premaxilla. Either way it has no ventral curvature of the osseus tip.

17. Proc. zygomaticus links the os squamosum with an outgrowth of the ala parasphenoidalis anteriorly enclosing the recessus tympanicus rostralis: 0, absent; 1, present. =Worthy et al. (1997: character 124).

18. Os premaxillare (rostrum maxillare), palatal surface: 0, central groove opens caudally via a slot or ill-defined foramen to narial cavity anterior to maxillo-palatines (defined in Howard 1929) (e.g. Malacorhynchus); 1, broad, elongate, fossa with margins thin and variable in shape opening into nares; 2, broad, solidly emarginate, elongate, fossa opening into nares (e.g. Anas).

Anhima is coded as missing data as its palatal surface has a large foramen, a non-homologous structure to the flat osseous surface in anatids.

19*. Lamina parasphenoidalis: 0, flat or concave across width; 1, with low medial elevation; 2, with marked elevation forming a rounded medial ridge.

Quadrate (Os quadratum) 20. Proc. mandibularis in lateral view, profile above cotyla quadratojugalis: 0, subparallel to plane across ventral margins of condyli lateralis et medialis, meeting ascending ramus of proc. oticus at about right angles (e.g. Malacorhynchus membranaceus); 1, ascends to meet ascending ramus of proc. oticus in wide angle (e.g. Anas).

21. The capitulum squamosum of proc. oticus overhangs lateral surface of proc. oticus: 0, yes (e.g. Anseranas); 1, no.

114 8 22. The dorsolateral margin extending from proc. oticus to lateral tuberosity on the proc. orbitalis: 0, forms straight line (e.g. Dendrocygna); 1, markedly concave. =Livezey (1986: character 15); also see Raikow (1971).

Mandible (mandibula) 23. Mandible, ventral curvature: 0, essentially lacking, < ½ depth os dentale (dentary) extends below middle of line linking tip with point of inflection at cranial end of dentary; 1, pronounced - > ½ depth below such line. =Livezey (1996a: character 4).

Anhima is non-comparable so treated as missing data.

24*. Mandible, regio coronoidei: 0, not deeper than distal end of dentary; 1, not markedly deep (c. 2x depth of the adjacent posterior dentary); 2, markedly deeper, >2x depth of the adjacent posterior dentary. Modified from Livezey (1996a: character 5), to better capture variation, because as defined by Livezey (1996a) it only applied to Chelychelynechen quassus Olson & James, 1991.

25. Proc. retroarticularis (retroarticular process), where present strongly lateromedially compressed, but with much variation in profile: 0, long, depth tapers markedly caudally, caudal profile a gentle curve with tip directed posterodorsally or dorsally, not recurved; 1, long, dorsoventrally broad (deep), caudal end rounded and tip little elevated above dorsal margin; 2, long, deep, caudal profile rounded or flattened, tip recurved and terminates high, c. x2 depth before upturn (e.g. Tadorna). Much modified from Livezey (1986: character 14).

26. Recessus conicalis: 0, absent or very shallow (e.g. Anseranas); 1, present, deep.

27. Foramen pneumaticum articulare in proc. medialis mandibulae: 0, present (e.g. Anseranas); 1, absent.

28. Os dentale essentially coplanar with os articulare: 0, yes; 1, no, os articulare down-curved posterior to proc. coronoideus (e.g. Malacorhynchus).

STERNUM 29*. Corpus sterni, presence of pneumatic foramina on sulcus medianus sterni (midline): 0, area of pneumatic foramina along midline (e.g. Anseranas); 1, zone of pneumatic foramina anteriorly on midline (e.g. Dendrocygna bicolor); 2, single well defined foramen anteriorly; 3, no foramen. =Livezey (1996a: character 24), modified; Livezey (1986: character 78), modified.

115 9 Note: The Lophodytes examined had a deep closed fossa in place of a pneumatic foramen so loss of pneumaticity is assumed to be secondary and the character coded as ‘2’. Most Cnemiornis sterna have no midline foramen, but MNZ S.35266 has a small pneumatic region near the midline cranially located dorsal to the left side of the carina suggesting a former state of paired pneumatic regions either side of a central bridge as seen in Anseranas; thus the absence is a reversal from state ‘0’.

30. Corpus sterni, pars cardiaca, pori pneumatici: 0, widely scattered on it (e.g. Anseranas); 1, limited to caudal margin of pila coracoidea (e.g. Cereopsis); 2, essentially absent. =Livezey (1996a: character 23); Livezey (1986: character 89).

31*. Corpus sterni, margo costalis, number of proc. costalis: 0, eight; 1, seven; 2, six or less. =Livezey (1996a: character 25).

Note: for this character, and for character 29, states may differ for congeneric species, especially between northern and southern hemisphere taxa.

32. Corpus sterni, margo caudalis with thickened ridge: 0, present; 1, absent (e.g. Cereopsis, coincident with carina sterni not extending to caudal margin). Modified from Livezey (1996a: character 26).

Note: Clangula has an autapomorphic condition where cartilage extending from the caudal margin of the sternum is ossified, but the caudal margin of the sternum anterior to this ossification has thickened laterally directed ridges.

33. Rostrum sterni, dorsal manubrial region, spina interna (dorsal manubrial spine): 0, absent, or a broad shallow medial notch bound by prominences on the labrum internum; 1, forms single, median rectanguloid flange (e.g. Malacorhynchus); 2, absent, replaced by a broad notch with a medial prominence (e.g. Anas). Modified from Livezey (1996a: character 28); Livezey (1986: character 82).

The notch is shallow in Oxyura and formed in a thin flange, and in Chloephaga hybrida, lateral thickenings are connected by a thin flange, so there is no notch: both taxa are here coded ‘0’.

34. Rostrum sterni, dorsal manubrial region, pila coracoidea: 0, thickened, robust ridge; 1, thin, not at all thickened (e.g. Oxyura).

116 10 35. Rostrum sterni, ventral manubrial region, spina externa (ventral manubrial spine): 0, lacking (e.g. Dendrocygna); 1, present.

36. Rostrum sterni, ventral manubrial region, spina externa (ventral manubrial spine): 0, short peg-like prominence, sometimes paired; 1, large (longer than wide) peg-like prominence; 2, dorso-ventrally flattened flange (e.g. Oxyura). Modified from Livezey (1996a: character 29); Livezey (1986: character 79).

Note: dorso-ventrally flattening of the flange is correlated with diving as Somateria, Oxyura, Aythya and Clangula all have flattened spina externa. Woolfenden (1961) stated that Aythya has a forked spina externa, but we note that in A. australis it is not forked. Oxyura vittata (USNM 614580) with a flattened ventral manubrial spine cleft by a v-shaped notch is coded ‘2’.

37*. Margo costalis, distal-most costal process: 0, located in distal half of corpus sterni (basin) measured from dorsal lip coracoidal sulcus and costal length >1/2 basin length (e.g. Anseranas); 1, located in distal half of basin but costal length <1/2 basin length (e.g. Dendrocygna); 2, located < or = midpoint of basin length. Modified from Livezey (1986: character 86).

SCAPULA 38. Scapus scapulae (blade): 0, uniform height or decreases over first 2/3 of length; 1, height increases distally of collum scapulae to a maximum at about ½ to ¾ blade length. Modified from Livezey (1996a: character 40); Livezey (1986: character 108).

Biziura has an unusual form, with the blade with parallel or slightly convergent dorsal and ventral margins, except in the distal 20% of length, where height expands markedly. Here it is coded ‘0’.

39. Extremitas cranialis scapulae, acromion: 0, equal cranial extent to tuberculum coracoideum (e.g. Anseranas); 1, extends distinctly craniad to tuberculum coracoideum. =Livezey (1996a: character 38); Livezey (1986: character 109).

40. Extremitas cranialis scapulae, foramen pneumaticum: 0, present laterally, variable in shape; 1, absent. =Livezey (1996a: character 37); Livezey (1986: character 111).

Oxyura jamaicensis MV B14125 has a small foramen, as does Tadorna tadorna MV B25679. In both species of Cnemiornis a pneumatic foramen is variably present, but its occurrence is rare in C. calcitrans, e.g. in MNZ S38934 one scapula has a small foramen, while in C. gracilis pneumatic foramina occur in most specimens and are large. 117 11

41. Collum scapulae, facies lateralis: 0, a single prominent attachment scars; 1, two prominent attachment scars, cranially by tuber. retinaculi, and caudally by tuber. scapulare (sensu Livezey (1996a)). This state arises as the caudal attachment point is enlarged and shifted ventrally from its homologous position on margo dorsalis in character state ‘0’. (e.g. Cereopsis). =Livezey (1996a: character 39); Livezey (1986: character 112).

CORACOIDEUM 42. Extremitas omalis coracoidei, processus procoracoideus (procoracoid): 0, foramen present in procoracoid and is pneumatic, opens into corpus; 1, foramen in procoracoid, but not opening into corpus; 2, foramen formed by ossified ligaments present (Cereopsis) or a notch formed by partial ossification of ligaments, especially cranially (Cygnus); 3, no foramen. =Livezey (1996a: character 43); Livezey (1986: character 92).

In Branta and some Anser the notch is small but clearly present with small ossified ligaments, so these taxa are coded as ‘2’. In Tadorna, the base of the procoracoid has a very shallow notch which may be a homologous structure, but it is clearly less developed than in Branta or Cygnus so is coded ‘3’ here.

43. Extremitas omalis coracoidei, processus acrocoracoideus (acrocoracoid) with pneumatic foramina under facies artic. clavicularis ( facet): 0, lacking; 1, present, in well defined fossa below dorsal part of clavicle facet; 2, present, in a broad area under clavicle facet (Cygnus a narrow area, Cereopsis a broad area). Modified from Livezey (1996a: character 42); Livezey (1986: character 95).

Stictonetta lacks pneumatic foramina under the head but overhanging lobes of the clavicle facet create a deep pocket.

44. Extremitas omalis coracoidei, acrocoracoid, clavicle facet, dorsal and ventral lobes: 0, not pronounced over sulcus supracoracoidei; 1, not pronounced equally, only dorsal lobe overhangs sulcus; 2, both lobes slightly overhang supracoracoidal sulcus; 3, pronounced overhang of both lobes over supracoracoidal sulcus (e.g. M. membranaceus). Modified from Livezey (1986: character 97).

In Malacorhynchus, the sulcus variably forms a pocket or a groove under the clavicle facet. For both Cnemiornis taxa, this character is coded as missing data as reduction of the element coincident with flightlessness precludes determining homology of the character state. 118 12

45. Extremitas omalis coracoidei, acrocoracoid, orientation, dorsal view: 0, plane through depth of acrocoracoid predominantly directed ventrally in range 60-90° to plane of sternal articulation so cranial end of acrocoracoid does not significantly overhang shaft (e.g. Anseranas, Tadorna); 1, plane through depth of acrocoracoid with distinct ventro-medial inclination (c. 45°) so often the cranial end of acrocoracoid significantly overhangs shaft (e.g. Anas).

In Lophodytes, the acrocoracoid has a marked medial inflection (1) despite the clavicle facet not significantly overhanging the shaft medial margin, thus characters 44 and 45 are not necessarily correlated. For both Cnemiornis taxa, this character is coded as missing data as reduction of the element coincident with flightlessness precludes determining homology of the character state.

46. Extremitas omalis coracoidei, supracoracoidal sulcus, excavated below facies artic. humeralis: 0, absent; 1, present.

For both Cnemiornis taxa, this character is coded as missing data as reduction of the element coincident with flightlessness precludes determining homology of the character state.

47. Corpus coracoidei, facies dorsalis, foramen pneumaticum cranial to facies artic. sternalis: 0, present; 1, absent. =Livezey (1996a: character 44); Livezey (1986: character 93).

48*. Corpus coracoidei, facies ventralis, impressio m. supracoracoideus a hollow bound laterally by a linea muscularis and caudally by the facies artic. sternalis: 0, absent, facies flat or convex ventrally; 1, present, distinct but shallow (e.g. Oxyura, Malacorhynchus); 2, present deep, typified by Stictonetta and Dendrocygna. Modified from Livezey (1996a: character 45); Livezey (1986: character 96).

49. Extremitas sternalis coracoidei, ventral facies artic. sternalis: 0, forms distinct buttressed facet with elevated rounded cranial margin (e.g. Dendrocygna); 1, facet not prominent over ventral surface (although a distinct facet may be present): allows the angulus medialis to fit into an acute angled socket of the sulcus artic. coracoideus in sternum; 2, broad flat articular facet for sternal articulation, no distinct ventral facies (unique to Anhima). Modified from Livezey (1986: character 100).

Oxyura jamaicensis SAM B36872 and MV B14125 have a flat ventral sternal facet not elevated above the ventral facies, but in SAM B36872, the facet is bound above by a narrow osseous lip. This probable pathology is unlike the typical buttress as seen in e.g. Dendrocygna, so the species is coded ‘1’. 119 13

50. Corpus coracoidei, orientation, line linking acrocoracoid with medial angle forms angle with line across lateral and medial extremes of sternal facet: 0, markedly greater than 90-100º; 1, approximates 90-100º.

This character is intrinsically related to the perception of the extent of the sternal lateral process, i.e. Livezey (1986: character 99), but as defined here the character better captures the form of the bone. For both Cnemiornis taxa this character is coded as missing data due to the reduced form of the coracoid coincident with flightlessness precluding determination of the homology of the character state.

HUMERUS 51. Margo caudalis, capital shaft ridge: 0, present; 1, absent.

In flightless taxa, or those with significant reduced flight ability, the presence of a compressed margo caudalis may be a non-homologous situation, and so for them this character is treated as missing data, e.g. Tachyeres. Livezey (1986) considered Chenonetta jubata to be problematic re determination of this state. Here we found a weak ridge to be consistently present, and one that is more developed than in T. tadorna. While Chenonetta finschi had reduced flight ability compared to C. jubata, it was probably not flightless (Worthy & Olson 2002) so is coded here with a capital ridge. Livezey (1996b) coded Nettapus with no ridge, however, we found a short distinct ridge extending from the base of the external tuberosity that merged into a broad rounded shaft, which could be argued is a vestigial capital ridge, but as this is not clear the taxon is coded ‘1’.

52*. Margo caudalis, capital shaft ridge: 0, prominent, directed towards caput (head) (e.g. Anseranas); 1, prominent, directed towards the zone between the head and the tuber. dorsale (external tuberosity); 2, prominent, directed/extends to external tuberosity. Modified from Livezey (1996a: character 51); Livezey (1986: character 22).

Coding the presence of a capital ridge as either directed towards the head or towards the external tuberosity can be problematic and has led to different determinations in taxa e.g. Thalassornis Ericson (1997) vs Livezey (1989), Livezey & Martin (1988). Therefore, we have added an intermediate state (1) for those taxa where the ridge is directed towards the ventral side of the external tuberosity. In Tadorna tadorna and T. tadornoides the capital ridge is weak, but as in other Tadorna species is directed to the ventral side of the external tuberosity so are coded ‘1’.

120 14 53*. Proximal end, fossa pneumotricipitalis dorsalis (dorsal pneumotricipital fossa) between incisura capitis (capital groove) and tuberculum dorsale (external tuberosity): 0, obsolete (e.g. Anseranas, Cereopsis); 1, forms narrow, shallow fossa extending to base of head: < width fossa pneumotricipitalis ventralis (ventral pneumotricipital fossa); 2, wide, shallow fossa: width ventral pneumotricipital fossa. Modified from Livezey (1986: characters 23 and 24).

54. Proximal end, crista deltopectoralis (deltoid crest): 0, anconally concave, dorsal margin rounded over length; 1, anconally flat or convex, dorsal margin angular. =Livezey (1986: character 25).

Note: all Oxyura are here considered to have a flat deltoid crest so coded ‘1’.

55*. Proximal end, crista deltopectoralis (deltoid crest), in cranial/palmar view, relative to junction of crista bicipitalis (bicipital crest) with shaft: 0, about 50% of length of deltoid crest extends distad of bicipital crest; 1, about 30-40% of length of deltoid crest extends distad of bicipital crest; 2, significantly less than 30% of deltoid crest extends distad of bicipital crest.

56. Proximal end, tuberculum dorsale (external tuberosity): 0, prominent, buttressed, elevated above surface of shaft; 1, essentially coplanar with shaft. Modified from Livezey (1986: character 32).

57. Proximal end, tuber. ventrale, in caudal/anconal view: 0, directed proximally so does not overhang ventral pneumotricipital fossa at all; 1, directed caudo-cranially so distal margin either is directed at right angles to the fossorial plane or overhangs the ventral pneumotricipital fossa. Modified from Livezey (1986: character 27).

58. Proximal end, ventral pneumotricipital fossa (= pneumatic foramen, Howard (1929)): 0, open, highly pneumatic, cavity with bony struts extends under margo caudalis (shallow in Anseranas, deep in e.g. Tadorna); 1, closed by bony wall internally, forming a conical-shaped pocket or fossa with the apex extending under the head and in anconal view the medial margin of this fossa forming a planar surface extending from the apex of the fossa to the junction of the bicipital crest and shaft. Modified from Livezey (1986: character 28).

The fossa wall may be perforated by macro- or microscopic foramina: this is especially evident in some Oxyura jamacensis individuals e.g. MV B14125. In osteologically immature specimens, the fossa is infilled with spongy bone, which is resorbed with maturity progressively deepening the fossa (e.g., Malacorhynchus, Oxyura, Clangula, Aythya novaeseelandiae). The pocket is variably deep in M. membranaceus, e.g. SAM B39385 is very deep, extending well under head. A closed 121 15 fossa has been related to diving (McCracken et al. 1999), but Malacorhynchus is not a diver (Frith 1977, Kear 2005). Both it and Mionetta lack diving such as elongated cnemial crests and flattened anterior shafts on tibiotarsi, or deepened popliteal fossa or curved shafts of femora (Worthy et al. 2007), and divers such as Mergus, Lophodytes and Aythya australis lack a closed fossa. Anseranas and Anhima have a shallow, though pneumatic, fossa, which could be coded as a separate state.

59*. Proximal end, incisura capitis, either anconal (caudal) or cranial view: 0, proximal profile not or barely interrupted by incisura; 1. proximal profile with very shallow notch, e.g., tadornids; 2, proximal profile with distinct notch created by incisura.

60*. Proximal end, crista bicipitalis (bicipital crest), shape in anconal/caudal view: 0, width across pneumatic foramen below tuberculum ventrale from crus dorsale fossae to crus dorsale ventrale distinctly less than length from ventral tubercle to where bicipital crest joins shaft (e.g. Anseranas); 1, width approximately equals length; 2, width distinctly greater than length e.g. Anas.

For both Cnemiornis taxa this character is coded as missing data as reduction of the humerus coincident with flightlessness precludes determining the homologous extent of bicipital crest.

61. Shaft: 0, essentially parallel sides in caudal or cranial views; 1, narrows distally (at least 10% reduction on mid-length width), narrowest point in distal third.

62. Proximal end, attachment scar of m. latissimus dorsi anterioris, (as distinct from attachment for m. latissimus dorsi posterioris) caudal view: 0, commences proximal to and links to distal end of deltoid crest before extending down shaft (e.g. Anseranas); 1, commences about level with and does not link to end of deltoid crest, and extends distally past end of deltoid crest; 2, commences proximal to distal end of deltoid crest and extends distad of deltoid crest without connection. (e.g. Tadorna).

63. Distal end, proc. flexorius (= entepicondyle): 0, short, ends proximad to condylus ventralis (in caudal view line across distal extreme of ventral condyle at right angles to long axis of shaft passes well distad of ventral condyle) (e.g. Anseranas, Cereopsis); 1, long, distal extent roughly equal to that of the entepicondyle.

64. Distal end, tuber. supracondylare dorsale (ectepicondylar prominence), in cranial view: 0, present, a distinct caudo-cranially thickened prominence in dorsal margin at proximal end dorsal condyle (e.g. Anseranas, Dendrocygna); 1, No prominence distinct from epicondylus dorsalis, which usually forms a low short proximally directed ridge. 122 16

Stictonetta and Malacorhynchus were coded ‘0’, as both have a distinct though weak prominence.

65. Distal end, tuber. supracondylare ventrale (attachment of the anterior artic. lig.): 0, facet parallel to shaft, not buttressed anteriorly (e.g. Anseranas); 1, facet buttressed anteriorly, tilted distally and/or medially (e.g. Anas).

A medial rotation of the facet is common to all divers (McCracken et al. 1999, pers. obs.) so is not factored into this character. Presence or absence of a buttress is considered the primary feature here, so for Anser in which the tubercle has little distal tilt but is clearly buttressed, is assigned state ‘1’. Modified from Livezey (1986: character 26).

66*. Distal end, sulcus scapulotricipitalis (external tricipital groove): 0, absent or barely defined (e.g. Anseranas); 1, present on anconal/caudal face, but not extending around distal end of ectepicondyle (e.g. Dendrocygna); 2, present, extends distally around ectepicondyle, forms distal notch in caudal (anconal) view.

67. Distal end, attachment of pronator brevis, sensu (Howard 1929), = attachment M. flexor carpi ulnaris (Livezey 1986): 0, pit on ventral facies, separate from facet for attachment of anterior ligament; 1, pit incorporated into ventral margin of facet for attachment of anterior ligament.

68. Distal end, fossa m. brachialis (brachial fossa): 0, well defined by margins, forms marked depression, often deepest distomedially; 1, poorly defined, flat, hardly shallower than surrounding facies (e.g. Somateria, Clangula).

In Dendrocygna, the brachial fossa is well defined but unusually small.

ULNA 69. Proximal end, palmar view, tuber. bicipitale ulnae (for insertion of M. biceps brachii): 0, forms prominent elongate tuberculum separated from cotylar margins and extending from below (=distad) proc. cotylaris dorsalis (dorsal cotylar process) diagonally and proximally towards cotylaris ventralis (ventral or internal cotyla) (e.g. Anseranas); 1, forms two distinct attachment points; one abutting the ventral cotyla in the incisura radialis and bound dorsally by the dorsal cotylar process; the second distal to the dorsal cotylar process (e.g. Malacorhynchus, Anas); 2, forms a single area of attachment abutting the ventral cotyla and bound dorsally by the dorsal cotylar process (e.g. Nettapus pulchellus).

123 17 One Dendrocygna arcuata specimen (ANWC 22214) had a divided tuber. bicipitale ulnae so the species was coded ‘0&1’.

70. Proximal end, fossa or pneumatic foramen under proc. cotylaris dorsalis (dorsal cotylar process): 0, absent; 1, present (e.g. Manuherikia, Malacorhynchus).

71. Length: 0, approximately equal to or greater than length humerus; 1, significantly (>5%) shorter than humerus.

Flightless taxa, or those with reduced flight ability, scored as missing data, e.g. Tachyeres; Chenonetta finschi is scored as its ulna is not reduced in length compared to highly volant Anas species and it probably was still volant, even though wing elements did shorten during the Holocene indicating some loss of flight ability (Worthy 1988, Worthy & Olson 2002).

CARPOMETACARPUS 72*. Proximal end, dorsal aspect, external rim trochlea carpalis (carpal trochlea): 0, forms even caudally convex curve to shaft, uninterrupted by notch (e.g. Anseranas); 1, with marked notch caudally; 2, interrupted by notch caudally but rim absent distal to notch (e.g. Biziura). Modified from Livezey (1986: characters 37+38), Livezey (1996a: character 55).

No hint of a notch was visible in the studied specimens of Dendrocygna bicolor and only a very indistinct notch was seen in D. eytoni and D. arcuata, one which was no more pronounced than in Anseranas, so both taxa were coded as ‘0’, contra Ericson (1997: 472) but as per Livezey (1986, 1996a) and Livezey & Martin (1988). Both Dendrocygna arborea examined in this study had a distinct notch, as did D. autumnalis, though it was shallower. The D. autumnalis carpometacarpus figured by Livezey & Martin (1988: Fig. 5) had a more developed notch than seen in D. eytoni, and D. autumnalis is the Dendrocygna Ericson (1997) examined. As for several other characters, there is variation among the species of Dendrocygna, precluding the use of the state in one species as representative of the genus. Specimens of Chenonetta finschi of Late Holocene age had shallower notches than Late Pleistocene specimens, and since over this period the species significantly progressed towards flightlessness (Worthy 1988), depth of the notch may be affected by flightlessness. Mionetta had a distinct shallow notch in all eight specimens examined, so was coded ‘1’.

73. Proximal end, ventral (internal) rim carpal trochlea, distal part: 0, uniform thickness with proximal part; 1, distinctly thickened. =Livezey (1986: character 47).

124 18 74. Proximal end, fovea carpalis cranialis (anterior carpal fossa): 0, absent, cranial margin of carpal trochlea flat – slightly convex (e.g. Anseranas); 1, present, cranial margin of carpal trochlea concave; 2, present, contains pneumatic foramen.

75. Proximal end, fovea carpalis caudalis (cuniform fossa): 0, shallow to moderately deep; 1, deep, high dorsal margin bounding it, fossa extending markedly below plane of metacarpal III; 2, present, shallow pneumatic foramen in it. Modified from Livezey (1986: character 46).

A deep fossa appears to be correlated with diving, as it is uniformly present in diving taxa.

76. Proximal end, ventral facies, fossa infratrochlearis (internal ligamental fossa): 0, shallow, (e.g. Anseranas); 1, deep, extends to at or below level of facies extensor process (e.g. Dendrocygna).

In taxa where the ventral surface of carpal trochlea is roughly coplanar with the ventral surface of the extensor process, e.g. anserines, depth is coded as shallow if depth is c. < 25% width of fossa.

77*. Proximal end, ventral facies, how ridge linking internal rim of carpal trochlea with proc. pisiformis (pisiform process) connects to facies of proc. extensorius (extensor process): 0, rounded profile (e.g. Dendrocygna); 1, sharp drop-off; 2, overhangs facies of extensor process with a fovea formed under ridge (e.g. Malacorhynchus).

Anhima is not comparable so treated as missing data.

78. Proximal end, proc. extensorius (process metacarpal I, extensor process): 0, perpendicular to, or proximally directed relative to shaft; 1, with distinct distal slope to proximal side of process (e.g. Cygnus). =Livezey (1986: character 41).

79. Proximal end, extensor process in ventral view: 0, not elongate, craniocaudal length less than craniocaudal width carpal trochlea; 1, elongate, length equal to or greater than width carpal trochlea. Modified from Livezey (1986: character 42); Livezey (1996a: character 56).

The presence of a rugosity on extensor process is related to fighting behaviour so is not characterised. The spur on Anhima is an autapomorphy not related to primary length of metacarpal 1 so Anhinga is coded as missing data for this character.

80*. Proximal end, caudal facies, os metacarpale minus (metacarpal III): 0, rounded or flattened adjacent to fornix with metacarpal II; 1, weakly or very shallowly grooved; 2, distinctly grooved. 125 19 Modified from Livezey (1986: character 44); Livezey (1996a: character 59). Contra Livezey & Martin (1988), metacarpal III in Mionetta was either weakly or strongly grooved, not ungrooved.

81. Proximal end, length of metacarpal II distal of proc. alularis (pollical facet) to start of intermetacarpal space relative to width (ventral view) of the fused metacarpals II and III in this section: 0, long, than width; 1, short, < than width.

82. Proximal end, facies dorsalis, ligament attachments near carpal rim: 0, with a single distinct scar proximally for the insertion of lig. ulnocarpo-metacarpale dorsale (external ligamental) (e.g. Anseranas); 1, with a distinct scar for the external ligament and a much smaller indistinct scar for the external scapholunar ligament (Woolfenden 1961: 25) located level with the top of the cuniform fossa on the ridge distad of the external ligament scar (e.g. Malacorhynchus); 2, with an external ligament scar and a slightly smaller distinct scar for the external scapholunar ligament, that is usually at least partly elevated off the surrounding facies, located as in ‘1’ (e.g. Anas); 3, with an external ligamental scar and a large distinct scar for the external scapholunar ligament, sometimes elongate, located further distad than in state ‘2’, level with the middle of the cuniform fossa, (e.g. Tadorna).

This character needs to be assessed with use of a microscope and is difficult or impossible to assess on greasy specimens. Note in Cygnus, which has state ‘3’, the more distally placed extensor process results in the scar for the external scapholunar ligament being level with the tip of the extensor process, which is the same spatial relationship as in ‘2’.

83*. Proximal end, dorsal view, M. extensor carpi ulnaris or flexor attachment (sensu Howard 1929): 0, two distinct rugosities, one adjacent to fornix os metacarpale minus et majus (metacarpal II and III), the other more proximad (e.g. Anseranas, Biziura); 1, one rugosity, approximately adjacent to fornix metacarpals II and III; 2, one rugosity, distal to fornix. Modified from Livezey (1986: character 43); Livezey (1996a: character 57).

Oxyura jamaicensis and O. australis have two abutting scars, herein coded as ‘0&1’. The intermetacarpal process of Gallus is treated as ‘2’.

84. Distal end, synostosis metacarpals II and III, maximum length, measured from distal end of intermetacarpal space to facies artic. digitalis minoris (facet for digit III): 0, length of synostosis less than width measured just distad of spatium intermetacarpale (intermetacarpal space) ie short synostosis (e.g. Anseranas, Malacorhynchus, Oxyura); 1, length width of synostosis, ie long synostosis, (e.g. Dendrocygna).

126 20 85*. Distal end, facies artic. digitalis minoris et major (facets for digits II and III): 0, facet for digit III extends farther distad than that for digit II; 1, facets for digits II and III of equal distal extent; 2, facet for digit III ends proximad to facet for digit II. =Livezey (1986: character 45); Livezey (1996a: character 61).

Contra Ericson (1997) but as per Livezey (1986, 1996a) we found Anseranas to be state ‘0’.

FEMORA Notes: Modifications associated with diving among various taxa result in marked similarity of features or homoplasy: Characters 89 (in part), 90, 92-96 tend to be homoplastic in diving taxa. Other characters clearly related to diving, such as marked narrowing of the femur shaft and marked distal expansion of the fibular condyle are not considered here.

86. Proximal end, caudal view, linea intermuscularis caudali (caudal intermuscular lines): 0, two lines, one laterally and one medially placed, oriented proximally; 1, one line, beginning at tubercle at mid-length, directed laterally towards trochanter, and a distinct ligamental tuberosity medially below head (e.g. Anseranas); 2, no lines, two distinct tuberosities, one near mid length, one as in ‘1’ below head; 3, one distinct line oriented towards caput (head) connecting tubercles at mid-length and below head. Modified from Livezey (1986: character 58).

87. Proximal end, facies articularis antitrochanterica, lateromedial plane: 0, surface concave (e.g. Anseranas); 1, surface convex.

88. Proximal end, cranial facies, crista trochanteris (trochanter), bounds or overhangs cranial surface to form: 0, flat or shallow fossa medially; 1, deep fossa medially (e.g. Stictonetta).

Note: taxa with a low trochanter (diving taxa) may be incomparable, code as missing data (?). Anhima has low ridge bounding pneumatic foramen.

89*. Distal end, tuber. m. gastrocnemialis lateralis forms ridge that is: 0, short, prominent, ovate, not extending proximally beyond sulcus patellaris; 1, elongate ridge with distinct medial bend, or oriented medially, but NOT extending proximad of patella sulcus; 2, long, extending well proximad of patella sulcus, narrow. In some taxa the ridge is straight e.g. Oxyura, whereas in others e.g. Anas, it is long but also has a distinct medial bend.

127 21 90. proximal end, anterior extent of trochanter compared to depth head: 0, relatively great, depth greatly exceeds depth head; 1, reduced, depth of trochanter approximately equal or slightly greater than depth head. =Livezey (1986: character 52); Livezey (1996a: character 68).

91. Distal end, distal extent in posterior view of condylus medialis (internal condyle): 0, approximately equal to that of condylus lateralis (external condyle); 1, distinctly less than that of external condyle. =Livezey (1986: character 53).

92. Distal end, sulcus patellaris (rotular groove) dorsal view: 0, short and broad, shallow - length of groove from proximal end of medial condyle (taken medially between the condyles) is approximately equal to or shorter than width across the condyles (taken from centre of their maximum elevations at mid-length in dorsal view); 1, elongate – length of groove much longer than width (as defined in ‘0’), usually deep. Modified from Livezey (1986: character 54).

Gallus has length approximately =width, but rotular groove is deep. Here coded ‘0’ as length prioritised. Livezey (1986) differentiated the variation seen in rotular groove structure by depth and relative distinction of the proximal margin, but we perceive length and relative narrowness as most significant.

93*. Curvature of shaft, lateral view: 0, straight or slight; 1, moderate ventrocaudal bending of distal third; 2, strongly bent. =Livezey (1986: character 55).

94. Distal end, fossa poplitea (popliteal fossa): 0, shallow; 1, deep. =Livezey (1986: character 56).

95. Posterior facies, ligamental attachment at mid-shaft (that is distal end of posterior intermuscular line): 0, not prominent; 1, prominent or markedly enlarged. Modified from Livezey (1986: character 57).

Note: Biziura is extreme.

96. Posterior facies, medial view, internal edge of distal end of shaft: 0, smoothly curving, continuous to condyle; 1, crista supracondylaris medialis expanded and or long, interrupts line of shaft leading to condyle (e.g. Biziura); 2, medial supracondylar ridge short, so medial profile notched. Modified from Livezey (1986: character 59).

128 22 TIBIOTARSUS Note. Height crista cnemialis cranialis (inner or procnemial crest), flattened anterior facies, and increased medial inflexion of condylus medialis (medial condyle) are all related to diving so are not employed as characters here. Further, characters 103 and 104 below are flagged as correlated with diving and so subject to homoplasy.

97*. Proximal end, impressio lig. collateralis medialis (ligamental attachment): 0, low, not prominent (e.g. Anseranas); 1, prominent relative to facies on which it is based (e.g. Tadorna, Anas); 2, very prominent (e.g. Malacorhynchus; see Woolfenden (1961).

98. Proximal end, medial facies, linea extensoria (intermuscular line) relative to ligamental attachment: 0, ligamental attachment separated from intermuscular line and medial facies approximately at right angles to adjacent cranial/anterior facies at distal end of ligamental attachment (e.g. Anseranas); 1, ligamental attachment separated from intermuscular line, which has moved laterally from medial margin (anterior view) so intervening section of medial shaft faces anteriomedially (e.g. Branta); 2, ligamental attachment abuts intermuscular line - resulting in anterior facies at top of fibular crest being markedly convex.

In specialist divers, where the inner cnemial ridge is extended down the shaft margin and the shaft flattened anteriorly, the intermuscular line cannot be distinguished from the extended cnemial crest so the character is coded as missing.

99. Distal end, epicondylus medialis with internal ligamental prominence: 0, pronounced, visible in anterior view (e.g. Anseranas); 1, present, occluded by rim of medial condyle in anterior view; 2, absent.

100. Distal end, medial condyle, distal aspect of medial facies of rim: 0, lacking notch (e.g. Anseranas); 1, shallow notch present, deep in some taxa e.g. Clangula. =Livezey (1986: character 62).

101. Distal end, sulcus m. fibularis (groove for peroneus profundus): 0, bridged cranially by bone (e.g., Dendrocygna); 1, bound laterally by sharp well defined crest and faces craniad (e.g. Anseranas); 2, poorly defined, bound laterally by rounded external ligamental prominence; 3, groove not discernable, external ligamental prominence elongate proximally as crest on craniomedial margin. Very much redefined from Livezey (1986: character 67).

Note: the groove is placed on lateral facies in Gallus so bounding crests are not homologous, so coded ‘-‘. 129 23

102. Proximal end, crista cnemialis cranialis (inner cnemial crest): 0, not deflected laterally (e.g. ancestor); 1, deflected laterally. =Livezey (1986: character 63); Livezey (1996a: character 70).

103. Distal end, anterior extent (depth) of condyles: 0, condylus medialis (internal condyle) distinctly greater than external; 1, approximately equal. =Livezey (1986: character 64).

104. Proximal end, inner cnemial crest: 0, lacking distinct anteriorly-raised ridge extending distally along anterior facies; 1, continued by a distinct ridge distally along antero-medial surface of shaft to a point well distad of the proximal end of the fibular crest. =Livezey (1986: character 65).

In Nettapus, the inner cnemial crest does not continue down the shaft, but the merging of the ligamental attachment with the intermuscular line and flattening of the anterior facies gives a superficially similar condition.

TARSOMETATARSUS

Several features of the tarsometatarsus are especially prone to homoplasy because of convergence related to either cursorality or diving. The relative lateral deflection of trochlea metatarsi IV (TIV) is greater in more terrestrial taxa, so extent of it is not coded herein. All divers are marked by narrowing and caudal rotation of TII so extent of this is not coded here.

105*. Trochlea metatarsi II (trochlea for digit II), distal extent: 0, approximately equal to TIV (e.g. Anseranas); 1, proximal to TIV, but extends to or distad of lateral inter-trochlear notch; 2, proximal to TIV, but distal extreme proximal to lateral inter-trochlear notch. Modified from Livezey (1986: character 68); Livezey (1996a: character 75); see also Worthy et al. (1997).

106. Proximal end, crista medialis hypotarsi (medial calcaneal ridge) posterior extent: 0, greatly elevated off posterior surface relative to other crista, or depth of medial side of crista depth of cotyla adjacent to this point (e.g. Anseranas, Biziura); 1, Not considerably greater than adjacent cristae and depth

107*. Proximal end, hypotarsus, width of base adjacent to cotylae, relative to proximal width: 0, much <½ proximal width; 1, about ½ of proximal width; 2, much >½ proximal width.

Note: Livezey’s (1986) character 72 combined number of hypotarsal ridges and position of the hypotarsus relative to the midline. Anseranas has four ridges, although the middle two are very poorly developed and defined merely by shallow sulci (not absent, contra Livezey 1986) and all 130 24 other anatids have four ridges with the middle cristae well-developed. Therefore, here we have restricted definition of this character to assessing the width of the hypotarsus only.

108*. Proximal end, fossa parahypotarsalis medialis: 0, very large deep, (e.g. Anseranas); 1, shallow, surface from medial calcaneal ridge to anterior margin of medial shaft concave (e.g. Dendrocygna); 2, absent, surface from medial calcaneal ridge to anterior margin of medial facies shaft flat or convex.

109. Distal end, trochlea metatarsi II (trochlea for digit II): 0, no medial groove, but a notch on medial surface (e.g. Anseranas); 1, medial groove present so distal margin notched. =Livezey (1986: character 74); Livezey (1996a: character 76).

Note: In most Dendrocygna species there is a notch on the medial surface of TII centred at mid- dorsoplantar depth, which is the precursor of the medial groove in other taxa. In Dendrocygna autumnalis (USNM 345768) this notch is more expanded than in other Dendrocygna and forms a shallow notch in the distal profile of the trochlea. In Oxyura the medial groove has its origin at about mid-dorsoplantar depth of the trochlea but in Anas the origin has moved dorsally to be near the dorsal surface of the trochlea. Livezey & Martin (1988) coded Mionetta as ‘0’ but our observations of several specimens indicate that there is a groove present.

110. Proximal end, groove extending distally from sulcus extensorius (extensor sulcus): 0, relatively shallow, not extending beyond 50% of total length; 1, deeper, may have high ridge medially, extends well beyond 50% total length (Stictonetta extreme).

111. Lateral margin, anterior view: 0, trochlea metatarsi IV (TIV) deflected from line of distal half of shaft (most taxa); 1, TIV with no deflection laterally relative to distal half of shaft (e.g. Biziura, Oxyura).

Note: as Clangula, Aythya, Lophodytes, Somateria, and Melanitta lack the derived astate (1) and all are divers yet Biziura/Oxyura, also divers, are state (0), this character is not directly related to diving.

112. Anterior end of two ligamental canals between trochlea for digits III and IV, in distal wall of foramen vasculare distale (distal foramen): 0, roofed over entirely in bone, not visible in anterior view; 1, largely or completely exposed anteriorly by reduction in bony covering. =Livezey (1986: character 69).

131 25 113. Fossa metatarsi I (metatarsal facet): 0, present, well marked; 1, absent or obsolete. =Livezey (1986: character 71).

114. Distal end, trochlea metatarsi II (trochlea for digit II): 0, expanded medially and caudally as a ‘wing’ (e.g. Anseranas); 1, flattened medial facies, not expanded medially. Modified from Livezey (1986: character 73).

115. Distal end, posterior opening of foramen vasculare distale (distal foramen): 0, opens flush onto plantar surface; 1, directed distoplantarly, so partially recessed into incisura intertrochlearis lateralis. =Livezey (1986: character 77).

116*. Shaft, width at mid-length: 0, wider than deep; 1, width approximately equals depth; 2, depth exceeds width.

117. Hypotarsus, medial hypotarsal ridge, distal end: 0, markedly hooked posterodistally forming notch; 1, ridge terminates abruptly, drops steeply to shaft; 2, ridge terminates by gradually lowering to shaft.

118*. Tarsometatarsus length, relative to femur: 0, longer than femur; 1, approximately equal to (+- 5%) femur; 2, shorter than femur.

TRACHEA 119. Syrinx, bulla syringealis, of males: 0, absent e.g. Oxyura; 1, present, small symmetrical enlargement (e.g. Dendrocygna); 2, present, large, asymmetrical osseus bulla. Modified from Livezey (1996a: character 81).

Notes: Delacour & Mayr (1945) reported the presence/absence of an osseus bulla for most taxa in their review of Anatidae. Livezey (1995a: 91) described Dendrocygna as having symmetrically enlarged bulla, which as reported by Delacour & Mayr (1945) are very different to other anatids. Livezey (1996b: 78, 79) stated Malacorhynchus has a vestigial bulla, and his fig 16F showed a syrinx in dorsal view revealed a tympanum comprised of several fused tracheosyringeal cartilages, but no enlarged bulla. SAM B39384 (male) shows a similar structure with a tympanum comprised of 9 fused tracheosyringeal cartilages and only a very slight dilation of the tympanum dorsally on the right side such that the opening to the right bronchia is more circular and slightly larger than that to the left, which is slightly ovoid, resulting in the tympanum being slightly asymmetrical. The syrinx of SAM B39384 is larger and more osseous than that of the similar-sized female SAM B39385, but we do not consider it a vestigial bulla as this implies a presence now nearly lost: at 132 26 most it could be considered rudimentary, but as it is much less developed than in Dendrocygna, assessment of such homology is unwarranted. Here Malacorhynchus is coded as not having a bulla. Similarly, Nettapus, e.g. MV B14490, lacks an enlarged bulla on its tympanum and is here coded as lacking a bulla, contra Livezey (1996b: 85), but consistent with observations reported in Marchant & Higgins (1990: 1223) and Kear (2005: 475, 477).

FURCULUM 120*. Extremitas omalis claviculae, facies artic. acrocoracoidea (coracoidal tuberosities or facets): 0, absent; 1, present, low facet; 2, present, prominent tuberosity. Modified from Livezey (1986: character 101), Livezey (1996a: character 36).

This character has been split to capture the low facets typical of anserines and tadornids as different from the prominent tuberosities of most anatids. Only Anhima is considered to be primarily without facets and Cnemiornis is considered to have lost them secondarily.

121*. Extremitas sternalis claviculae, apophysis furculae, (furcular process or hypocleideum): 0, obsolete, no structure visible; 1, present as single (e.g. Tadorna) or paired (e.g. Cereopsis) low ridges; 2, present as a prominent ridge or lobe. Modified from Livezey (1986: character 102); Livezey (1996a: character 33).

Anseranas males have a low ridge dorsally (1) and a unique projection ventrally.

122. Scapus claviculae, facies lateralis: 0, lacking foramina pneumatica; 1, with foramina pneumatica. =Livezey (1986: character 105); Livezey (1996a: character 35).

123. Furculum, overall form: 0, robust, apophysis furculae tending towards V-shaped with furculae divergent dorsally; 1, robust, apophysis furculae U-shaped, furculae diverging dorsally; 2, robust, apophysis furculae broadly U-shaped, furculae not diverging further dorsally; 3, slender, apophysis furculae broadly U-shaped, furculae not diverging further dorsally.

For both Cnemiornis taxa this character is coded as missing data as reduction of the coincident with flightlessness precludes determining homologous shape.

MISCELLANEOUS SKELETAL CHARACTERS

124*. Vertebrae – total number of cervical and thoracic vertebrae: 0, 20; 1, 21; 2, 22; 3, 23; 4, 24- 25; 5, 26-30.

133 27 Data is mostly after Verheyen (1955a-d) and Woolfenden (1961): data for Cnemiornis is from Worthy et al. (1997); for Chenonetta finschi is from Worthy & Olson (2002) and specimens examined.

125. Femora caudal aspect, trochlea fibularis with distinct depression immediately proximal of the articular surface: 0, Not so; 1, Yes.

It seems that loss of this feature is related to diving as most divers lack it, but not all.

126. Femora, caudal aspect, impressio ansae m. iliofibularis forms distinct facet at proximal end of the lateral edge of the trochlear fibularis: 0, Yes, at junction of caudal and lateral facies and directed caudally-caudolaterally; 1, Yes, entirely on lateral facies (not visible in caudal view) and faces laterally; 2, Yes, on caudal facies (not visible in lateral view), and directed caudoproximally; 3, No facet discernible.

Note: in Gallus, the distal location of a round tuberculum m. gastrocnemialis lateralis results in the impressio ansae m. iliofibularis being displaced distally and laterally, but while it is on the lateral surface it is visible in caudal view and directed caudolaterally so is coded ‘0’.

127. At least 2 thoracic vertebrae fused/ossified forming a notarium: 0, yes eg, Anseranas; 1, no.

Note: Tadorna may have vertebrae held together by ossified tendons but corpus of vertebrae not fused.

128. Mandibular rami, lateral compression defining long, very narrow inter-ramal region: 0, absent; 1, present. = Livezey (1996b: character 72), though here Malacorhynchus is coded 'present' as very similar to Stictonetta in its inter-ramal conformation.

INTEGUMENT CHARACTERS 129*. Modal number of pairs of retrices: 0, five; 1, six; 2, seven; 3, eight; 4, nine; 5, 10-12.

Derived from Livezey (1996b: character 78) and Livezey (1996a: character 124). Coded following data in Cramp & Simmons (1977) and Marchant & Higgins (1990). The two Australasian Dendrocygna species have 6 pairs retrices (Marchant & Higgins 1990), whereas Livezey (1996a) coded ‘Dendrocygna’ with 7 pairs so we have left as ? those taxa for which we did not obtain direct data. Similarly, Livezey (1996b) coded ‘oxyurines’ with 7 pairs, whereas O. australis and O. jamaicensis have 9 pairs and Biziura 10-12 pairs (Cramp & Simmons 1977, Marchant & Higgins 134 28 1990): both taxa were included as oxyurines by Livezey (1995b), thus other Oxyura species and Nomonyx are left as ?. Tadorna is a third genus with interspecific variation, as T. tadornoides and T. variegata each have 6 pairs of retrices, but T. radjah, T. tadorna and T. ferriginea have 7 pairs (Cramp & Simmons 1977, Marchant & Higgins 1990).

130. Leg colour in adults: 0, black or grey; 1, yellow; 2, red; 3, flesh or pink. =Livezey (1996b: character 81) modified with the addition of ‘3’, and so relates to Livezey (1995a: character 24). Coded following data in Marchant & Higgins (1990) and Kear (2005).

131. Bill colour in breeding males: 0, black or gray; 1, pinkish white; 2, orange, pink, or red; 3, yellow; 4, blue. =Livezey (1996b: character 82), modified with the addition of ‘4’ for Oxyura. Coded following data in Marchant & Higgins (1990) and Kear (2005).

132. Tarsus, centre of dorsal surface: 0, reticulate; 1, scutellate. =Livezey (1996b: character 87) and Livezey (1996a: character 112).

133. Pedal unguis: 0, not strongly developed or curved; 1, strongly developed, curved, claw-like. =Livezey (1996b: character 100).

135 29 Appendix 2. Character matrix, 57 taxa, 133 characters.

Gallus gallus 0 - 1 0 1 0 0 0 1 0 0 1 0 0 - - 0 - 0 1 1 - - 0 0 [01] 0 0 3 2 2 - - - 1 1 2 1 1 1 - 3 0 3 - 0 1 0 2 1 - - 2 1 2 1 0 0 2 2 - - 1 0 0 0 0 0 0 0 0 [12] 0 1 0 0 2 0 0 0 1 0 2 0 0 0 - 0 0 0 0 0 0 0 ? 0 0 1 [12] 0 - 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 2 0 0 0 1 0 0 0 2 [01] 0 1 1 Anhima cornuta 0 0 1 0 0 - 0 2 0 0 0 0 0 0 - - 0 - 0 0 1 0 - 0 1 0 0 0 0 0 0 1 0 0 0 - 1 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 2 0 2 0 0 1 0 0 0 0 1 0 0 0 2 2 0 - 0 - 0 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 2 1 1 0 0 0 0 1 0 0 0 1 0 0 0 1 0 0 ? 0 0 0 0 1 2 3 ? ? 0 0 2 0 0 0 1 Anseranas semipalmata 0 0 0 0 0 - 0 1 - 0 0 0 0 1 0 - 0 0 1 0 0 0 0 1 2 0 0 0 0 0 [12] 0 0 0 0 - 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 [01] 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 4 1 0 0 0 1 1 0 0 1 Thalassornis leuconotus 0 0 1 1 0 0 0 1 ? 1 ? ? 0 1 0 ? 0 ? ? ? ? 0 0 1 1 1 1 0 3 2 1 ? 0 1 0 - ? 0 1 1 0 3 0 3 1 1 1 2 0 1 0 1 1 0 0 0 1 1 2 2 1 0 1 0 0 1 1 0 ? ? 0 0 ? 1 ? 1 0 0 0 2 0 ? 0 0 1 3 1 - 2 1 1 ? 1 1 0 0 1 - 1 ? 1 1 1 1 2 1 2 1 1 0 1 0 1 1 1 0 0 ? 0 1 ? 0 ? ? ? ? ? 0 2 0 0 0 0 Dendrocygna arborea 1 0 1 1 0 0 0 1 3 1 0 1 0 1 0 2 0 1 2 0 1 0 0 1 2 1 0 0 3 1 1 1 0 0 0 - 1 1 1 1 0 3 0 1 0 1 1 2 0 1 0 1 1 0 1 0 0 0 0 0 0 2 1 0 1 1 0 0 0 0 0 1 0 1 1 1 0 0 0 0 0 0 0 1 1 3 1 0 1 0 1 1 0 0 0 0 1 0 1 0 1 1 0 0 1 1 2 1 0 0 0 0 1 1 1 0 [01] 0 1 1 0 0 2 2 1 0 1 0 ? 0 0 0 1 Dendrocygna autumnalis 1 0 0 1 0 0 0 1 2 1 0 [12] 0 1 0 2 0 1 2 0 1 0 0 1 2 1 1 0 [01] 1 [12] 0 0 0 0 - 1 0 1 1 0 3 0 1 0 1 1 2 0 1 0 1 1 0 1 0 0 0 0 0 0 [02] 1 0 1 1 0 0 0 0 0 1 0 1 1 1 0 0 0 1 0 0 0 1 1 3 1 0 1 0 1 1 0 0 0 0 1 2 1 0 1 1 0 0 2 1 2 1 [01] 0 0 0 1 1 1 0 [01] 0 1 1 1 0 2 2 1 0 1 0 ? 3 2 0 1 Dendrocygna bicolor 1 0 0 1 0 0 0 1 2 1 0 1 0 1 0 1 0 1 1 0 1 0 0 1 2 1 0 0 2 1 1 0 0 0 0 - 1 0 1 1 0 3 0 1 0 1 1 2 0 1 0 1 1 0 2 0 0 0 0 0 0 0 1 0 1 1 0 0 0 0 0 0 0 1 0 1 0 0 0 1 0 0 0 1 1 3 1 0 1 0 1 0 0 0 0 0 1 2 [01] 1 [01] 1 0 0 1 1 2 1 0 1 0 0 1 1 1 0 1 0 1 1 1 0 2 2 1 0 1 0 ? 0 0 0 1 Dendrocygna eytoni 0 0 0 1 1 0 0 1 2 4 0 1 0 1 0 2 0 1 1 0 1 0 0 1 2 1 0 0 1 1 2 0 0 0 0 - 1 1 1 [01] 0 3 0 1 0 1 1 2 0 1 0 1 1 0 1 0 0 0 1 0 0 2 1 0 1 1 0 0 0 0 0 0 0 1 0 1 0 0 0 1 0 0 0 1 1 3 1 0 1 0 1 0 0 0 0 0 1 0 1 1 [01] 1 0 0 1 1 2 1 0 1 0 0 1 1 1 0 0 0 1 1 1 0 1 3 1 0 0 0 1 3 1 0 1 Dendrocygna arcuata 0 0 0 1 0 0 0 1 2 4 0 1 0 1 0 1 0 1 2 [01] 1 0 0 1 2 1 1 0 1 1 2 0 0 0 0 - 1 1 1 1 0 3 0 1 0 1 1 2 0 1 0 1 1 0 2 0 0 0 1 0 0 0 1 0 1 1 0 0 [01] 0 0 0 0 1 1 1 0 0 0 0 0 1 1 1 1 3 1 0 1 0 1 1 0 0 0 1 1 2 1 1 1 1 0 0 1 1 2 1 0 0 0 0 1 1 1 0 0 1 1 2 1 0 2 3 1 0 1 0 1 0 0 0 1 Oxyura vittata 0 0 1 1 1 0 0 1 3 2 2 2 0 1 0 0 0 0 2 0 1 [01] 0 2 2 1 1 1 3 2 1 0 0 1 1 2 2 1 1 1 0 3 0 1 0 1 1 0 1 1 1 - 2 1 1 0 1 1 2 1 1 0 1 1 0 2 0 0 1 0 1 1 1 1 1 0 0 0 0 0 1 1 0 0 2 3 1 - 2 1 1 1 1 1 1 1 1 - 1 1 1 1 0 1 2 1 2 2 1 1 1 0 1 1 1 0 0 2 0 2 0 0 3 1 0 0 1 0 ? 0 4 1 0 Oxyura jamaicensis 0 0 1 1 0 0 0 1 3 2 2 2 0 1 0 0 0 0 0 0 1 [01] 0 2 2 1 1 1 3 2 1 0 0 1 1 2 2 0 1 1 0 3 0 2 0 1 1 1 1 1 0 2 2 1 2 0 1 1 2 2 1 0 1 1 0 2 1 0 1 1 1 1 1 1 1 0 1 0 0 0 1 0 [01] 0 1 3 1 - 2 1 1 1 1 1 1 1 1 - 1 1 2 1 1 1 2 1 2 2 1 [01] 1 0 1 1 1 0 1 2 0 2 0 0 3 [12] 0 0 1 0 4 0 4 1 0 Oxyura maccoa 0 0 1 1 0 1 0 1 3 2 2 2 1 1 0 0 0 0 1 0 1 [01] 0 1 2 1 1 0 3 2 2 0 0 1 1 2 2 1 1 1 0 3 0 1 0 1 1 1 1 1 0 2 2 1 2 0 1 1 2 2 1 0 1 1 0 2 1 0 0 0 1 1 0 1 1 1 1 0 0 1 0 0 0 0 2 3 1 - 2 1 1 1 1 1 1 1 1 - 1 1 1 1 1 1 2 1 2 2 1 0 1 0 1 1 1 0 0 2 0 2 0 0 2 1 0 0 1 0 ? 0 4 1 0 Oxyura australis 0 0 1 1 0 0 0 1 3 2 2 2 0 1 0 0 0 0 0 0 1 [01] 0 1 2 1 1 0 3 2 2 0 0 1 1 2 2 0 1 1 0 3 0 2 0 1 1 1 1 1 0 2 2 1 2 0 1 1 2 2 1 0 1 1 0 2 1 0 1 0 1 1 1 1 1 0 0 0 0 1 1 0 [01] 0 1 3 1 - 2 1 1 1 1 1 1 1 1 - 1 1 2 1 1 1 2 1 2 2 1 [01] 1 0 1 1 1 0 0 2 0 2 0 0 3 1 1 0 1 0 4 0 4 1 0 Nomonyx dominicus 0 0 2 1 0 0 0 1 2 1 2 2 0 1 0 0 0 1 1 0 1 0 0 2 2 1 1 0 3 2 2 0 0 0 1 1 2 0 1 1 0 3 0 1 0 1 1 0 1 1 1 - 2 1 2 0 1 0 2 2 0 0 1 1 1 2 1 0 0 0 1 1 0 1 1 0 [12] 0 0 1 0 0 0 0 1 3 1 - 2 1 1 1 0 1 1 1 1 - 1 1 1 1 0 1 2 1 2 2 1 0 1 0 1 1 1 0 0 2 0 2 0 0 2 1 0 0 1 0 ? 0 4 1 0 Stictonetta naevosa 0 0 2 1 0 1 0 1 2 2 2 1 1 1 0 0 0 0 0 0 1 0 1 1 2 1 1 0 2 2 2 0 0 0 1 1 2 0 1 1 0 3 0 3 1 1 1 2 1 1 0 2 1 0 2 0 0 0 2 1 0 1 1 0 1 2 0 1 1 0 1 1 0 1 0 0 0 0 0 2 1 2 2 1 2 3 1 1 1 0 1 0 0 0 0 0 1 2 1 1 2 1 0 0 1 1 2 2 1 1 0 0 1 1 1 0 2 2 0 2 1 1 2 3 1 0 1 1 2 0 0 0 0 Biziura lobata 0 0 1 0 0 0 0 2 2 1 0 2 0 1 0 1 0 1 1 0 0 1 0 1 1 1 1 0 3 2 1 1 0 0 0 - 1 0 1 1 0 3 0 0 0 0 1 0 0 1 0 1 1 0 1 0 1 1 2 1 0 2 1 1 0 0 1 0 2 0 1 2 1 0 1 1 0 0 0 0 1 1 0 [01] 1 2 1 - 2 1 1 1 2 1 1 1 0 - 1 1 3 1 1 1 2 0 2 2 1 0 1 0 1 1 1 0 [01] 2 0 2 0 0 3 2 [01] 2 1 0 5 0 0 1 0 Malacorhynchus membranaceus 0 1 2 1 1 0 0 1 3 2 2 2 1 1 0 0 0 0 0 0 1 1 0 1 1 1 1 1 2 [12] 1 0 1 0 1 1 2 0 1 [01] 0 3 0 3 1 0 1 1 1 1 0 2 1 0 2 0 1 1 2 2 0 1 1 0 1 2 0 0 1 1 1 1 0 1 0 0 2 0 0 2 1 1 2 0 2 3 1 0 2 0 1 0 0 1 1 0 2 2 1 1 [23] 1 0 0 2 1 2 2 1 0 0 0 1 1 1 1 0 1 0 2 1 0 2 1 1 0 1 1 1 0 0 1 0 Nettapus pulchellus 0 0 2 1 1 0 0 1 3 4 2 1 0 1 0 2 0 0 1 1 0 1 0 1 0 1 1 1 2 2 2 0 2 0 1 1 2 1 1 1 0 3 0 2 1 0 1 1 0 1 1 - 1 1 2 1 1 0 2 2 0 1 1 1 1 2 0 1 2 0 1 1 0 1 0 0 1 0 0 0 1 2 2 [01] 2 3 1 0 2 0 1 1 1 0 1 0 2 2 1 1 2 1 0 0 2 1 2 2 1 0 1 0 1 1 1 0 0 2 0 2 2 0 1 1 1 0 1 0 2 0 0 1 0 N. coromandelianus 0 0 0 1 1 0 0 1 3 4 2 1 0 1 0 2 0 0 1 1 0 1 0 1 0 1 1 1 2 2 2 0 2 0 1 1 2 1 1 1 0 3 0 3 1 0 1 1 1 1 1 - 1 1 2 1 1 0 2 2 0 1 1 1 1 2 0 0 [12] 0 1 1 0 1 0 [01] 0 0 [01] 0 1 2 2 1 2 3 1 1 2 0 1 1 1 0 0 0 2 2 1 1 2 1 0 0 2 1 2 2 1 0 [01] 0 1 1 [01] 0 [01] 2 0 2 2 0 1 1 1 0 1 0 2 0 0 1 0 Cereopsis novaehollandiae 0 2 0 1 2 1 1 2 1 4 2 1 0 1 0 2 1 0 0 0 1 0 0 1 0 1 1 0 1 1 2 1 0 0 [01] 0 1 0 1 0 1 2 2 3 0 1 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 1 0 [01] 0 0 0 0 1 [01] 0 3 1 1 1 3 1 0 1 0 1 0 0 0 1 1 1 2 0 0 1 1 0 0 1 1 1 1 1 0 0 0 1 1 1 0 2 0 0 1 1 1 2 [34] 1 0 1 0 2 2 0 0 1 136 30 Cnemiornis calcitrans 0 0 0 1 0 0 1 2 0 4 1 1 0 1 0 2 1 1 0 0 1 0 0 1 0 1 1 0 3 1 1 1 0 0 0 - 1 0 0 [01] - 2 0 - - - 1 0 1 - - - 0 0 0 0 0 0 0 - 0 1 0 0 0 0 0 0 0 0 - 0 0 0 0 0 0 0 0 0 1 0 1 1 1 0 1 0 1 0 1 0 0 0 1 0 2 1 0 0 1 1 0 0 0 1 1 1 1 0 0 1 1 1 1 0 2 2 ? 0 0 1 - 3 0 0 1 0 ? ? ? ? ? Cnemiornis gracilis 0 0 0 1 0 0 1 2 0 ? 1 1 0 1 0 2 1 1 0 0 1 0 0 1 0 1 1 0 3 1 1 1 0 0 0 - 1 0 0 [01] - 2 [01] - - - 1 0 1 - - - 0 0 0 0 0 0 0 - 0 1 0 0 0 0 1 0 ? 0 - 0 0 0 0 0 ? ? ? 0 1 0 1 0 [12] 0 1 0 1 0 1 0 0 0 1 0 2 1 0 0 3 1 0 0 0 1 1 1 1 0 0 1 1 1 1 0 2 2 ? 0 0 1 - ? 0 0 1 0 ? ? ? ? ? Branta canadensis 0 0 1 1 2 1 1 1 1 4 1 1 0 1 0 2 0 2 2 1 1 1 0 1 0 1 1 0 [01] [01] 1 0 1 0 1 1 1 1 1 0 0 2 2 1 0 1 1 [01] 0 0 0 1 1 0 1 0 0 0 0 0 0 2 0 1 1 1 0 0 0 0 1 1 0 0 0 0 0 0 1 [12] 0 3 1 1 [12] 3 1 0 1 0 1 0 0 0 1 1 1 1 1 0 2 1 0 0 [12] 1 2 2 1 0 0 0 1 1 1 1 2 0 0 1 1 0 2 4 0 1 1 0 [34] 0 0 0 0 Anser brachyrhynchus 0 0 0 1 0 0 0 1 0 4 1 1 0 1 0 2 0 2 2 1 1 0 0 1 0 1 1 0 1 1 1 0 1 0 1 0 1 0 1 0 0 2 2 1 0 1 1 0 0 0 0 0 1 0 1 0 0 0 0 1 0 2 0 1 1 1 0 0 0 0 1 1 0 1 0 0 0 0 0 2 1 3 1 1 2 3 1 0 1 0 1 0 0 0 1 1 1 1 1 0 3 1 0 0 1 1 2 2 1 0 0 0 1 1 1 1 2 1 0 1 1 0 2 4 0 1 1 0 4 3 [02] 0 0 Anser caerulescens 0 0 0 1 0 [01] 0 1 0 4 1 1 0 1 0 2 0 2 2 1 1 0 1 1 0 1 1 0 2 1 1 1 1 0 1 1 1 1 1 0 0 3 2 1 0 1 1 0 0 0 0 0 1 0 1 0 0 0 0 0 0 2 0 1 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 3 0 0 2 3 1 0 1 0 1 1 0 0 1 1 1 0 0 0 3 1 0 0 1 1 2 2 1 0 0 0 1 1 1 1 2 0 0 1 1 1 2 3 1 1 1 0 3 3 1 0 0 Cygnus atratus 0 0 0 1 0 1 1 2 0 4 1 1 0 1 0 2 0 2 2 1 0 1 0 1 2 1 1 0 0 1 1 1 0 0 1 1 1 0 1 1 0 2 2 1 0 0 1 0 0 1 0 0 1 0 1 0 0 0 0 0 0 2 0 0 0 2 0 0 0 0 1 1 0 0 0 0 0 1 0 0 0 3 1 1 1 1 1 0 1 0 1 0 0 0 1 0 0 2 1 0 1 1 0 0 2 1 2 2 1 0 0 0 1 1 1 0 2 0 0 1 0 1 1 5 1 1 1 0 3 0 2 0 0 Tadorna ferriginea 0 0 0 1 0 0 0 1 2 4 1 2 1 1 0 1 0 1 1 1 1 1 1 1 2 1 1 0 2 2 2 0 0 0 1 1 2 0 1 1 0 3 0 1 1 1 1 0 1 1 0 1 1 0 2 0 0 0 1 0 0 2 1 1 1 2 0 1 1 0 0 1 0 1 0 0 0 0 0 2 0 3 1 0 2 3 1 0 1 0 1 0 0 0 0 0 1 2 0 1 3 1 0 0 2 1 2 2 1 0 0 1 1 1 1 1 2 0 2 1 1 0 2 1 0 1 1 0 2 2 0 1 0 Tadorna tadornoides 0 0 0 1 0 0 0 1 1 4 1 2 1 1 0 2 0 1 0 1 1 1 1 1 2 1 1 0 2 2 1 0 0 0 1 1 2 0 1 1 0 3 0 1 0 1 1 0 1 1 0 1 1 0 1 0 0 0 1 1 0 2 1 1 1 2 0 0 1 0 1 1 0 1 0 0 0 0 0 2 0 3 1 0 2 3 1 0 1 0 1 0 0 0 0 0 1 2 1 1 3 1 0 0 1 1 2 2 1 0 0 0 1 1 1 1 1 1 2 1 1 0 2 2 1 1 1 0 1 2 0 1 0 Tadorna tadorna 0 0 0 1 2 0 1 1 1 4 1 2 1 1 0 1 0 1 1 1 1 1 1 1 2 1 1 0 2 2 1 0 0 0 1 1 1 0 1 1 0 3 0 1 0 1 1 0 1 1 0 1 1 0 1 0 0 0 1 0 0 2 1 1 1 2 0 0 1 0 1 1 0 1 0 0 0 0 0 2 1 3 2 0 2 3 1 0 1 0 1 0 0 0 0 0 1 2 1 1 1 1 0 0 1 1 2 2 1 0 0 0 1 1 1 1 0 1 2 1 0 0 2 2 1 1 1 0 2 2 0 1 0 Tadorna variegata 0 0 0 1 0 0 0 1 1 4 1 2 1 1 0 2 0 1 0 1 1 1 1 1 2 1 1 0 2 2 2 0 0 0 1 1 2 0 1 1 0 3 0 1 0 1 1 0 1 1 0 1 1 0 1 0 0 0 1 1 0 2 1 1 1 2 0 0 1 0 1 1 0 1 0 0 0 0 1 2 1 3 2 1 2 3 1 0 1 0 1 0 0 0 0 0 1 [12] 1 1 2 1 0 0 1 1 2 2 1 0 0 0 1 1 1 1 2 0 2 1 0 0 2 [12] 1 1 1 0 1 2 0 1 0 Tadorna radjah 0 0 2 1 2 0 0 1 1 4 1 2 1 1 0 2 0 1 1 1 1 1 1 1 2 1 1 1 1 1 2 0 0 0 1 1 2 1 1 1 0 2 0 1 0 0 1 0 1 1 0 2 1 0 1 1 0 0 1 1 0 2 1 1 1 2 0 0 1 0 1 1 0 1 0 0 0 0 0 2 0 3 2 1 2 3 1 0 1 0 1 0 0 0 0 0 1 2 1 1 1 1 0 0 1 1 2 2 1 0 0 0 1 1 1 0 0 0 2 1 1 0 2 1 1 1 1 0 2 2 1 1 0 Alopochen aegyptiacus 0 0 0 1 2 1 0 2 3 4 1 2 1 1 0 2 0 1 2 1 1 1 0 1 0 1 1 0 2 1 2 0 0 0 0 - 1 0 1 1 0 3 1 1 0 1 1 0 0 1 0 2 1 0 1 0 0 0 0 2 0 2 1 1 1 2 0 0 1 0 1 1 0 1 0 0 0 0 1 2 0 3 2 1 1 3 1 0 1 0 1 0 0 0 0 0 1 0 [01] 1 1 1 0 0 1 1 2 2 1 0 0 0 1 1 1 1 2 0 2 1 1 0 2 1 1 1 1 0 2 3 1 1 0 Chloephaga hybrida 0 0 1 [01] 2 0 1 1 1 4 0 2 0 1 0 2 0 2 0 0 1 1 0 2 ? 1 1 0 1 1 2 0 0 0 0 - 2 1 1 1 0 3 0 1 0 1 1 0 0 1 0 1 2 0 1 0 0 0 0 ? 0 2 1 1 1 2 0 0 0 0 1 1 0 1 0 0 0 0 1 2 1 3 1 1 2 3 1 0 1 0 1 0 0 0 0 2 1 2 1 1 1 1 0 0 1 1 2 1 1 0 0 0 1 1 1 0 0 0 2 1 0 0 2 1 ? ? ? 0 2 1 0 1 0 Chloephaga picta 0 0 1 1 2 1 1 1 0 4 1 2 1 1 0 2 0 1 ? 0 1 1 0 2 0 1 1 1 2 1 2 1 0 0 0 - 2 1 1 1 0 3 0 1 1 1 0 0 1 1 1 - 1 0 1 0 0 0 0 1 0 2 1 1 1 2 0 0 1 0 0 1 0 1 0 0 0 0 1 2 1 3 1 1 2 3 1 0 1 0 1 0 0 0 1 0 1 1 1 1 1 1 0 0 1 1 2 2 1 0 0 0 1 1 1 0 2 0 2 1 1 0 2 1 ? ? ? 0 2 0 0 1 0 Somateria mollisima 0 0 1 1 3 0 1 1 1 3 0 2 1 1 0 2 0 1 1 1 1 1 0 2 2 1 1 0 3 2 2 1 0 0 1 0 2 0 1 0 0 3 0 1 0 0 1 0 1 1 0 2 1 1 2 1 1 1 2 1 0 2 1 1 1 2 0 1 1 0 1 1 0 1 1 1 0 0 0 1 0 2 1 1 2 3 1 - 2 1 1 1 1 1 0 0 1 - 1 1 1 1 1 1 2 1 2 2 1 0 0 1 1 1 1 1 0 2 2 2 1 0 3 1 0 1 1 0 2 0 [03] 1 0 Melanitta fusca 0 2 0 1 0 1 1 1 - 4 0 2 0 1 0 2 0 1 0 1 1 1 1 2 2 1 1 0 3 2 2 0 0 0 0 - 2 1 1 1 0 3 0 1 0 1 1 0 1 1 0 2 2 1 2 1 1 1 2 0 0 2 1 1 1 2 0 1 1 0 1 1 1 0 1 0 0 0 0 1 1 2 1 0 2 3 1 - 2 1 1 1 2 1 1 1 2 - 1 1 1 1 1 1 2 1 2 2 1 0 0 1 1 1 1 1 0 2 2 2 0 0 3 [12] 1 1 1 0 2 0 [03] 1 0 Clangula hyemalis 0 0 1 1 3 0 1 1 1 3 0 2 0 1 0 2 0 1 1 1 1 1 1 2 2 1 1 0 3 2 1 0 0 0 1 0 2 0 1 1 0 3 0 1 0 1 1 0 1 1 1 - 1 1 2 1 1 1 2 2 0 1 1 1 1 2 0 1 2 0 1 1 1 1 1 0 0 0 0 1 1 2 2 0 1 3 1 - 2 1 1 1 2 1 0 0 0 - 1 1 1 1 1 1 2 1 2 2 1 0 0 0 1 1 1 1 0 2 2 2 1 0 3 1 0 1 1 0 2 0 [01] 1 0 Bucephala clangula 0 0 0 1 0 1 1 1 - 4 0 2 0 1 0 2 0 1 2 1 1 1 0 1 2 1 1 0 2 2 2 0 0 0 0 - 2 1 1 1 0 3 0 1 0 1 1 0 1 1 0 2 2 1 2 0 1 1 2 2 1 2 1 1 1 2 0 0 2 0 1 1 0 1 0 1 0 0 0 2 0 2 1 0 2 3 1 - 2 1 1 1 2 1 1 0 1 - 1 1 3 1 1 1 2 1 2 2 1 0 0 0 1 1 1 0 0 2 2 2 0 0 2 1 0 1 1 0 3 0 0 1 0 Chenonetta jubata 0 0 1 1 0 0 0 1 2 1 0 2 0 1 0 2 0 1 2 1 1 1 0 2 0 1 1 1 2 2 2 0 2 0 1 1 2 1 1 1 0 3 0 [13] 0 1 1 0 1 1 0 2 1 0 2 0 0 0 2 0 0 2 1 1 1 2 0 0 1 0 1 1 0 1 0 0 0 0 0 1 0 2 1 1 2 3 1 0 1 0 1 0 0 0 0 2 1 0 1 1 1 1 0 0 1 1 2 2 1 0 0 0 1 1 1 1 [02] 1 2 2 0 0 2 1 1 0 1 0 2 0 0 1 1 Chenonetta finschi 0 0 1 1 2 0 1 1 2 1 0 2 0 1 0 2 0 1 0 1 1 1 0 2 0 1 1 1 2 2 2 0 2 0 1 0 2 1 1 1 0 3 0 [13] 0 1 1 0 1 1 0 2 1 0 1 0 1 0 2 1 0 2 1 1 1 2 0 0 1 0 1 1 0 1 0 1 0 0 0 2 1 1 2 0 2 3 1 0 1 0 1 0 0 0 0 2 1 0 1 1 1 1 0 0 1 1 2 2 1 0 0 0 1 1 1 0 2 2 2 2 1 0 2 1 0 1 1 0 ? ? ? ? ? Hymenolaimus malacorhynchos 0 0 2 1 0 0 0 1 2 1 1 2 0 1 0 2 0 1 1 1 1 1 0 1 1 1 1 0 2 2 2 0 0 0 1 0 2 1 1 1 0 3 0 1 0 1 1 0 1 1 0 2 1 1 1 1 1 0 2 1 0 2 1 1 1 2 0 0 2 0 1 1 1 1 0 1 0 0 0 0 1 3 2 1 2 3 1 0 2 0 1 0 0 1 0 1 1 2 1 1 1 1 0 0 [12] 1 2 2 1 0 0 0 1 1 1 1 2 1 2 2 1 0 2 1 1 1 1 0 1 0 1 1 0 137 31 Lophodytes cucullatus 0 0 1 1 0 0 0 1 2 1 1 2 0 1 1 1 0 1 1 0 1 1 0 1 2 1 1 0 2 2 [12] 0 0 0 0 - 2 1 1 1 0 3 0 1 1 1 1 0 1 1 1 - 2 1 2 1 1 0 2 2 0 1 1 1 1 2 0 [01] 1 0 1 1 0 1 1 0 [01] 0 0 1 1 2 2 0 2 3 1 - 2 1 1 1 2 1 1 0 1 - 1 1 1 1 1 1 2 1 2 2 1 0 0 0 1 1 1 0 [01] 2 2 2 1 0 1 1 0 1 1 0 4 0 0 1 0 Mergus merganser 0 0 1 1 0 0 1 2 2 1 1 2 0 1 1 1 0 1 1 0 1 1 1 1 2 1 1 0 2 2 2 0 0 0 0 - 2 1 1 1 0 3 0 1 1 1 1 0 1 1 1 - 1 1 2 1 1 0 2 2 0 1 1 1 1 2 0 0 1 0 1 1 0 1 1 0 0 0 0 1 0 0 2 0 2 3 1 - 2 1 1 1 1 1 1 1 1 - 1 1 2 1 1 1 1 1 2 2 1 0 0 0 1 1 1 2 0 2 2 2 0 0 1 1 0 1 1 0 4 1 2 1 0 Mergus serrator 0 0 1 1 0 0 0 2 2 1 1 2 0 1 1 1 0 1 2 0 0 1 0 1 2 1 1 0 2 2 2 0 0 0 0 - 2 1 1 1 0 3 0 1 1 1 1 0 1 1 0 1 1 1 2 1 1 0 2 2 1 1 1 1 1 2 0 0 1 0 1 1 0 1 0 0 0 0 0 2 1 1 2 1 2 3 1 - 2 1 1 1 1 1 1 1 1 - 1 1 2 1 1 1 1 1 2 2 1 0 0 0 1 1 1 2 0 1 2 2 0 0 2 [12] 0 1 1 0 4 0 2 1 0 Anas superciliosa 0 0 0 1 0 0 0 1 2 4 0 2 1 1 0 2 0 2 1 1 1 1 0 1 2 1 1 1 2 2 1 0 2 0 1 1 2 1 1 1 0 3 0 2 1 1 1 0 1 1 1 - 1 1 2 1 1 0 2 1 0 1 1 1 1 2 0 0 1 0 1 1 0 1 0 0 0 0 0 2 1 2 2 1 2 3 1 0 2 0 1 1 0 0 0 0 1 2 1 1 2 1 0 0 1 1 2 2 1 0 0 0 1 1 1 0 0 2 2 2 1 0 2 1 1 0 1 0 4 0 [02] 1 0 Anas rhynchotis 0 0 1 1 0 0 0 1 2 4 0 2 1 1 0 2 0 2 2 1 1 1 1 1 2 1 1 1 2 2 1 0 1 0 1 1 2 1 1 1 0 3 0 2 1 1 1 0 1 1 1 - 1 1 2 1 1 0 2 1 0 2 1 1 1 2 0 0 1 0 1 1 0 1 0 0 0 0 0 2 1 2 2 1 2 3 1 0 2 0 1 1 0 0 0 0 1 2 1 1 3 1 0 0 1 1 2 2 1 0 0 0 1 1 1 1 0 2 2 2 1 0 2 1 1 0 1 0 2 0 [02] 1 0 Anas castanea 0 0 1 1 [02] 1 0 1 2 4 2 2 1 1 0 2 0 2 2 0 1 1 1 1 2 1 1 1 2 2 1 0 2 0 1 1 2 1 1 1 0 3 0 [12] 1 0 1 0 1 1 1 - 1 1 2 1 1 0 2 0 0 1 1 1 1 2 0 0 1 0 1 1 0 1 0 0 0 0 0 2 1 2 2 1 2 3 1 0 2 0 1 1 0 0 0 0 1 2 1 1 2 1 0 0 1 1 2 2 1 0 0 0 1 1 1 0 1 2 2 2 1 0 2 1 1 0 1 0 3 0 [02] 1 0 Aix galericulata 0 0 0 1 0 0 0 1 1 4 0 2 0 1 0 2 0 1 2 1 1 1 0 1 0 1 1 1 2 1 2 0 0 0 0 - 2 1 1 1 0 3 0 2 1 1 1 0 1 1 1 - 1 1 2 1 1 0 2 1 0 1 1 1 1 2 0 0 1 0 1 1 0 1 0 1 0 0 0 2 1 1 2 1 2 3 1 0 2 0 1 1 0 1 0 0 1 2 1 1 2 1 0 0 1 1 2 2 1 0 0 0 1 1 1 0 1 2 2 2 2 0 2 1 1 0 1 0 3 1 2 1 0 Aythya fuligula 0 0 0 1 0 0 1 1 3 1 0 2 0 1 0 2 0 1 0 1 1 1 0 2 2 1 1 0 2 2 1 0 0 0 0 - 2 0 1 0 0 3 0 1 1 0 1 0 1 1 1 - 2 1 2 1 1 1 2 1 1 1 1 1 1 2 0 0 1 0 1 1 0 1 1 1 0 0 0 2 1 2 1 1 2 3 1 - 2 1 1 1 1 1 0 0 1 - 1 1 2 1 1 1 2 1 2 2 1 0 0 0 1 1 1 0 1 2 2 2 1 0 1 2 0 1 1 0 2 0 0 1 0 Aythya australis 0 0 0 1 0 0 1 1 3 1 1 2 0 1 0 2 0 1 2 1 1 1 0 2 2 1 1 0 2 2 1 0 0 0 1 0 2 1 1 1 0 3 0 1 1 1 1 0 1 1 1 - 2 1 2 1 1 0 2 1 0 2 1 1 1 2 0 0 1 0 1 1 0 1 1 1 0 0 0 2 0 2 1 1 2 3 1 - 2 0 1 1 0 0 0 0 0 - 1 1 2 1 0 1 2 1 2 2 1 0 0 0 1 1 1 0 1 2 2 2 1 0 2 2 1 1 1 0 2 0 0 1 0 Aythya novaeseelandiae 0 0 0 1 0 0 0 1 3 1 0 2 1 1 0 2 0 1 1 1 1 1 1 2 2 1 1 0 2 2 1 0 0 0 1 0 2 1 1 1 0 3 0 1 1 1 1 0 1 1 1 - 2 1 2 1 1 1 2 1 1 2 1 1 1 2 0 0 1 0 1 1 0 1 1 1 0 0 0 2 0 2 1 1 2 3 1 - 2 1 1 1 2 1 1 0 1 - 1 1 2 1 1 1 2 1 2 2 1 0 0 0 1 1 1 0 0 2 2 2 0 0 2 2 0 1 1 0 2 0 0 1 0 Aythya affinus 0 0 1 1 3 0 1 1 3 1 0 2 0 1 0 2 0 1 1 1 1 1 0 2 2 1 1 0 2 2 1 0 0 0 1 0 2 0 1 1 0 3 0 1 1 1 1 0 1 1 1 - 2 1 2 1 1 1 2 1 0 2 1 1 1 2 0 0 1 0 1 1 0 1 1 0 0 0 0 2 0 2 1 1 2 3 1 - 2 1 1 1 1 1 1 0 1 - 1 1 1 1 1 1 2 1 2 2 1 0 0 0 1 1 1 1 1 2 2 2 0 0 1 2 0 1 1 0 2 0 0 1 0 Netta rufina 0 0 1 1 0 0 1 1 3 1 0 2 0 1 0 1 0 1 1 1 1 1 0 2 2 1 1 1 2 2 1 0 0 0 0 - 2 1 1 1 0 3 0 1 0 1 1 0 1 1 0 2 2 1 2 1 0 0 2 0 0 2 1 1 1 2 0 0 1 0 1 1 0 1 0 0 0 0 0 2 0 2 1 1 2 3 1 - 2 1 1 1 0 1 1 0 2 2 1 1 [01] 1 0 0 2 1 2 2 1 1 0 0 1 1 1 1 0 2 2 2 0 0 1 2 1 1 1 0 2 0 2 1 0 Mionetta blanchardi 0 ? ? ? 0 0 0 1 3 ? ? 2 0 1 0 2 0 0 [12] ? ? ? 0 1 ? ? ? ? 3 2 1 ? 0 0 1 1 0 1 1 1 0 3 0 1 0 1 1 1 1 1 0 2 1 0 0 0 0 1 2 1 0 0 1 1 1 2 0 0 1 0 1 1 0 1 0 1 0 0 0 [12] 0 1 1 1 2 3 1 0 2 0 1 1 0 0 0 0 1 2 1 1 2 1 0 0 2 1 2 2 1 0 0 0 1 1 1 0 0 1 ? ? 1 ? ? ? 1 0 ? ? ? ? ? ? ? Manuherikia lacustrina ? ? ? 1 2 0 1 ? 2 ? ? ? 0 ? ? ? ? ? ? 1 1 1 ? ? ? 1 ? ? [23] 2 ? ? ? ? ? ? ? 0 1 1 0 3 0 1 1 0 1 1 1 1 0 2 1 0 1 0 1 1 2 1 1 2 1 1 1 2 0 0 2 1 1 1 0 1 [01] 1 1 0 0 [12] 1 1 1 1 1 3 1 - 2 1 1 1 1 1 0 0 1 - 1 1 3 1 0 1 2 1 2 2 1 0 0 0 1 1 1 0 [01] 2 ? ? ? ? ? ? 0 2 ? ? ? ? ? ? ? Dunstanetta johnstoneorum ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 3 1 2 0 0 1 0 1 1 0 2 2 0 ? 0 ? 1 ? ? ? ? 1 0 1 2 0 0 1 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 3 ? - 2 ? 1 1 1 1 1 0 ? ? ? ? ? ? ? ? 2 1 2 2 1 0 0 0 1 1 1 0 2 2 ? ? ? ? ? ? 0 2 ? ? ? ? ? ? ? Miotadorna sanctibathansi ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 1 1 ? ? ? ? ? ? ? ? ? ? 0 ? 1 0 ? ? 1 1 0 3 2 2 0 [01] 1 0 0 1 0 1 1 0 2 0 0 0 1 1 0 2 1 0 1 2 0 0 1 0 1 1 0 1 0 0 0 0 0 1 1 1 2 0 2 3 1 0 1 0 1 1 0 0 0 1 ? ? 1 ? 3 ? 0 ? 1 ? ? 2 1 0 0 0 1 1 1 0 ? ? ? ? ? ? ? ? 1 1 ? ? ? ? ? ? ? 138 32 REFERENCES

BAILLON, L. A. F. 1834. Catalogue des Mammiferes, Oiseaux, Reptiles, Poissons et Mollusques testaces marins, observés dans l’arrondissement d’Abbeville. Mémoirs de la Socièté royale d’émulation d’Abbeville, sér. 2, no. 1 (1833), 49-80. BENEDEN, P. J. van 1875. Un oiseau fossile nouveau des caverns de la Nouvelle Zélande. Annales de la Société Géologique de Belgique 2: 123-130, Plate 3. BULLER, W. L. 1869. On some new species of New Zealand birds. Ibis 2 (5): 37-43. CRAMP, S. and SIMMONS, K. E. L. (eds) 1977. Handbook of the Birds of Europe, the Middle East and North Africa: The birds of the western Palearctic. Vol. 1, ostrich to ducks. Oxford University Press, Oxford. Vii + 722 pp + 40 pls. DELACOUR, J. and MAYR, E. 1945. The family Anatidae. Wilson Bulletin, 57, 3-55. ERICSON, P. G. P. 1997. Systematic relationships of the Palaeogene family Presbyornithidae (Aves: Anseriformes). Zoological Journal of the Linnean Society, 121, 429-483. EYTON, T. C. 1838. A monograph on the Anatidae, or Duck Tribe. Longmans, London, x + 178 pp. FLEMING, J. H. 1935. A new genus and species of flightless duck. Occasional Papers of the Royal Ontario Museum, Zoology 1: 1-3. FORBES, H. O. 1892 (March). On a recent discovery of the remains of extinct birds in New Zealand. Nature 45(1166): 416-418. FRITH, H. J. 1977. Waterfowl in Australia. A.H. & A.W. Reed, Sydney, 328 pp. GMELIN, J. F. 1789. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synomymis, locis. Tomus I. Linné, editio decima tertia, aucta, reformata. Regnum Animalium. G. E. Beer, Leipzig (Lipsiae), Pt. 2, 501-1032. GOULD, J. 1836. [Untitled, Oxyura australis] Proceedings of the Zoological Society of London IV [for 27 September 1836]: 85-86. GOULD, J. 1841. Untitled [= ‘Mr. Gould completed the exhibition of his fifty new species of Australian birds…. ’] Proceedings of the Zoological Society of London VIII [for 8 December 1840]: 169-179. GOULD, J. 1842. The Birds of Australia, pt. 6, Author, London,. GOULD, J. 1856. On two new species of birds ( notabilis and variegata) from the collection of Walter Mantell, Esq. Proceedings of the Zoological Society of London XXIV [for 22 April 1856]: 94-95. GRAY, G. R. 1845. Birds. In RICHARDSON, J. and GRAY, J. E. (eds). The Zoology of the Voyage of H.M.S. Erebus and Terror, under the Command of Captain Sir James Clark Ross, R.N., F.R.S., during the years 1839 to 1843. 2 vol. London, E.W. Janson. GRAY, G. R. 1849. The genera of Birds: comprising their generic characters, a notice of the habits of each genus, and an extensive list of species referred to their several genera. Longman, Brown, Green and Longmans, London, 3 vols, 669 pp, CLXXXV pls. HOMBRON, J. B. & JACQUINOT, H. 1841. Description de plusieurs oiseaux nouveaux ou peu connus, provenant de l’expédition autour de monde faite sur les corvettes l’Astrolabe et la Zéleé. Annales des Sciences Naturelles, séries 2, 16, Zoologie: 312-320. HORSFIELD, T. 1824. Zoological Researches in Java and the neighbouring islands, pt 8. Parbuny and Allen, Kingsbury, London, 328 pp, 72 pl. [Issued in 8 pts., 1821-24]. 139 33 HOWARD, H. 1929. The avifauna of Emeryville Shellmound. University of California Publications in Zoology, 32(2), 301-394. HUMPHREY, P. S., THOMPSON, M. C. 1981. A new species of steamer-duck (Tachyeres) from Argentina. University of Kansas Museum of Natural History, Occasional Papers 95: 1-12. JARDINE, W. & SELBY, P. J. 1828. Illustrations of , Volume 1, Pt IV, W. H. Lizars, Edinburgh, pp H-L (2), pls 51-65. KEAR, J. (ed.) 2005. Ducks, geese and swans. Oxford University Press, Oxford, UK. 2 vols, 908 pp. LATHAM, J. 1802. Index Ornithologicus, sive systema ornithologiæ: complectens Avium divisionem in classes, ordines, genera, species, ipsarumque varietates. Supplementum, &c. Leigh & Sotherby, London, lxxiv pp. LATHAM, J. 1790. Index Ornithologicus, sive systema ornithologiæ : complectens Avium divisionem in classes, ordines, genera, species, ipsarumque varietates. Vol. 1. Leigh & Sotherby, London, xviii + 920 pp. LATHAM, J. 1798. An essay on the tracheae or windpipes of various kinds of birds. Transactions of the Linnean Society, London 4: 90-128, pls. 9-16. LESSON, R. P. 1828. Manuel d’Ornithologie, ou Description des Genres et des Principales Espèces d’Oiseaux. Vol 2. Paris, Roret. LINNAEUS, C. 1758. Systema Naturae per Regna Tria Naturae, 10th Edition, revised, Vol 1: Regnum Animale. Salvii, L. Stockholm, Sweden, Holmiae. iv + 824 pp. LINNAEUS, C. 1766. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio duodecima, reformata. Holmiae, Laurentii Salvii, Vol. 1, 1327 & 36 pp. LIVEZEY, B. C. 1986. A phylogenetic analysis of Recent anseriform genera using morphological characters. The Auk, 103, 737-754. — 1995a. A phylogenetic analysis of the whistling and white-backed ducks (Anatidae: Dendrocygninae) using morphological characters. Annals of the Carnegie Museum, 64, 65-97. — 1995b. Phylogeny and comparative ecology of stiff-tailed ducks (Anatidae: Oxyurini). Wilson Bulletin, 107, 214-234. — 1996a. A phylogenetic analysis of geese and swans (Anseriformes: Anserinae), including selected fossil species. Systematic Biology, 45, 415-450. — 1996b. A phylogenetic reassessment of the tadornine-anatine divergence (Aves: Anseriformes: Anatidae). Annals of the Carnegie Museum, 65, 27-88. LIVEZEY, B. C. and MARTIN, L. D. 1988. The systematic position of the Miocene anatid Anas [?] blanchardi Milne- Edwards. Journal of Vertebrate Paleontology, 8(2), 196-211. MARCHANT, S. and HIGGINS, P. J. (co-ordinators) 1990. Handbook of Australian, New Zealand & Antarctic birds, vol. 1, Ratites to Ducks. Oxford University Press, , 1400 pp. MCCRACKEN, K. G.; HARSHMAN, J.; MCCLELLAN, D. A. and AFTON, A. D. 1999. Data set incongruence and correlated character evolution: an example of functional convergence in the hind-limbs of stiff-tail ducks. Systematic Biology, 48, 683- 714. MILNE-EDWARDS, A. 1863. Mémoire sur la distribution géologique des oiseaux fossils et description de quelques especies nouvelles. Annales des Sciences Naturelles 4(20): 132- 176. [Abbreviated versions in Bulletin de la Sociétié Philomatique 1863: 61-64; Comptes Rendus des Séances Hebdomadaires de l’Académie des Sciences (Paris) 56: 1219-1222; Revue des Sociétés Savantes (1)4: 1-5.] MOLINA, G. I. 1782. Saggio sulla Storia naturale del Chili. Fratelli Masi e Comp., Bologna, 8 vols, 367 pp, 1 map. 140 34 OLSON, S. L. 1977. Notes on the subfossil Anatidae from New Zealand, including a new species of pink-eared duck Malacorhynchus. The Emu 77: 123-135. OLSON, S. L. and JAMES, H. F. 1991. Descriptions of thirty-two new species of birds from the Hawaiian Islands: Part 1. Non-Passeriformes. Ornithological Monographs, 45, 1-88. OWEN, R. 1866. On Dinornis (Part X): containing a description of part of the skeleton of a indicative of a new genus and species (Cnemiornis calcitrans, Ow.). Transactions of the Zoological Society of London V(5): 395-404, pls. LXIII-LXVII. PALLAS, P. S. 1764. Adumbratiunculae avium variarum praecedenti Elencho insertarum, sed quae in Systemate Naturae Illustr. Linnaei nondum extant. 1-7. In VOSMAER, A. (ed.), Catalogue raisonné, d’une collection supérieurement belle d’Oiseaux, tant exotiques qu’Européens, de Quadrupedes et d’Insectes. Empaillés, & arrangés avec beaucoup d’art en situations & attitudes extrèmement naturelles, & garantis de la corruption d’une façon particulière. Le tout rassemblé & arrangé, pendant une longue suite d’années, avec beaucoup de peines & à grand fraix, par A. Vroeg. Collection qu’on offre aux amateurs entiére & à un prix raisonnable, jusqu’au 22 Septembre de cette année; après l’Echéance duquel terme, elle sera vendue aux plus offrands, le 6 Octobre 1764 à la Haye, dans la Maison de Mr. Coster, au coin du Veenestraat sur le petit Marché aux Herbes, par Pierre van Os, Libraire demeurant sur la Place à la Haye. Pp. i-xvi, 1-49, 1-7. PALLAS, P.S., 1773. Reise durch verschiedene Provinzen des Russischen Reichs. Zweyter Theil. St. Petersburg, pp. 371-750, 19 tt. PHILIPPI, R. A. 1860. Ueber zwei vermuthlich neue Chilenische Enten (Anas iopareia, Erismatura vitta), und über Fringlla barbata Mol. Wiegmann. Archiv für Naturgeschichte, 26 (1), 24-28. SCLATER, P. L. 1857. Note on the . Proceedings of the Zoological Society of London XXV [for 9 June 1857]: 128. SHAW, G. 1796. In SHAW, G and NODDER, F. P. eds. The Naturalist’s Miscellany; or coloured figures of natural objects drawn and described immediately from nature. Nodder, London, Vol. 8, fascicle LXXXV, pls 255-257. VERHEYEN, R. 1955a. La systématique des Ansériformes basée sur l’ostéologie comparée. Institut royal des Sciences natuelles de Belgique Bulletin, 31(35), 1-18. — 1955b. La systématique des Ansériformes basée sur l’ostéologie comparée. Institut royal des Sciences natuelles de Belgique Bulletin, 31(36), 1-16. — 1955c. La systématique des Ansériformes basée sur l’ostéologie comparée. Institut royal des Sciences natuelles de Belgique Bulletin, 31(37), 1-22. — 1955d. La systématique des Ansériformes basée sur l’ostéologie comparée. Institut royal des Sciences natuelles de Belgique Bulletin, 31(38), 1-16. VIEILLOT, L. J. P. 1816. Analyse d’une nouvelle ornithology elementaire. Paris, Déterville, 70 pp. WOOLFENDEN, G. E. 1961. Postcranial osteology of the waterfowl. Bulletin of the Florida State Museum. (Biol. Sci.), 6, 1-129. WORTHY, T. H. 1988. Loss of flight ability in the extinct New Zealand duck Euryanas finschi. Journal of Zoology, London, 215, 619-628. WORTHY, T. H. 2004. The Holocene fossil waterfowl fauna of Lake Poukawa, North Island, New Zealand. Tuhinga 15: 77-120. WORTHY, T. H. 2005. A new species of Oxyura (Aves: Anatidae) from the New Zealand Holocene. Longmann Symposium. Memoirs of the Queensland Museum 51: 255-272. WORTHY, T. H. and OLSON S. L. 2002. Relationships, adaptations, and habits of the extinct duck ‘Euryanas’ finschi. Notornis, 49, 1-17. 141 35 WORTHY, T. H.; HOLDAWAY, R. N.; SORENSON, M. D. and COOPER, A. C. 1997. Description of the first complete skeleton of the extinct New Zealand goose Cnemiornis calcitrans Owen, (Aves: Anatidae), and a reassessment of the relationships of Cnemiornis. Journal of Zoology, London, 243, 695-723. WORTHY, T. H.; TENNYSON, A. J. D.; JONES, C.; MCNAMARA, J. A. and DOUGLAS, B. J. 2007. Miocene waterfowl and other birds from Central Otago, New Zealand. Journal of Systematic Palaeontology, 5(1), 1-39.

142

143 144 145 146 147

A new species of the Manuherikia and evidence for geese (Aves: Anatidae: Anserinae) in the St Bathans Fauna (Early Miocene), New Zealand Darling Building, DP 418, Department of Earth and Environmental Sciences, The University of Adelaide, Adelaide, SA 5005, Australia. ‹ Museum of New Zealand Te Papa Tongarewa, P.O. Box 467, Wellington, New Zealand ‹ School of Biological, Earth and Environmental Sciences, University of New South Wales, New South Wales 2052, Australia

‹ Abstract Manuherikia Cereopsis CnemiornisCnemiornis Cereopsis

A Worthy, T.H., Tennyson, A.J.D., Hand, S.J. & Scofield, R.P. (2008) A new species of the diving duck Manuherikia and evidence for geese (Aves: Anatidae: Anserinae) in the St Bathans Fauna (Early Miocene), New Zealand. Journal of Royal Society of New Zealand, v. 38(2), pp. 97-114

A NOTE: This publication is included on pages 148-173 in the print copy of the thesis held in the University of Adelaide Library. 174

175

CHAPTER 5

Descriptions and phylogenetic relationships of two new genera and four new species of Oligo-Miocene waterfowl (Aves: Anatidae) from Australia.

Trevor H. Worthy

Department of Earth and Environmental Sciences, The University of Adelaide, Adelaide, SA 5005, AUSTRALIA

Zoological Journal of the Linnean Society (2008), accepted paper. 176

STATEMENT OF AUTHORSHIP

Descriptions and phylogenetic relationships of two new genera and four new species of Oligo-Miocene waterfowl (Aves: Anatidae) from Australia

Trevor H. Worthy

Zoological Journal of the Linnean Society (2008), accepted paper.

Worthy, T. H. (Candidate) Designed the research, did the research, interpreted the data, wrote the manuscript, and acted as corresponding author.

I hereby certify that the statement of contribution is accurate

Signed …………………………………………………………. Date ……………….

177

Abstract (177) The Tertiary anatid fossils (Aves: Anatidae) from Oligocene and Miocene deposits in Australia are described. Most fossils derive from the Late Oligocene – Early Miocene (26- 24 Ma) Etadunna and Namba Formations, respectively in the Lake Eyre and Lake Frome Basins of South Australia. The local faunas from these two formations contain the same suite of anatid species. Two new genera, the oxyurine Pinpanetta, with three new species (P. tedfordi, 18 specimens; P. vickersrichae, 15 specimens; P. fromensis, 20 specimens), and the tadornine Australotadorna, for a large new species known from eight specimens, are established. Three anatid bones from the Waite Formation (c. 8 Ma) at Alcoota, Northern Territory reveal the presence of a tadornine that is neither Australotadorna nor an extant Tadorna species, and an indeterminate duck about the size of Malacorhynchus. Phylogenetic analyses establish Pinpanetta as a basal member of an oxyurine (stiff-tailed duck) radiation. Oxyurines are found to include the Recent Stictonetta and Malacorhynchus as basal members, along with the fossil taxa Mionetta, Manuherikia and Dunstanetta, and the traditionally included Recent Oxyura, Biziura, Thalassornis and Nomonyx.

ADDITIONAL KEYWORDS: Oxyurinae – Pinpanetta – Australotadorna – phylogeny Running Title: Australian Tertiary anatids

A Worthy, T.H. (2008) Descriptions and phylogenetic relationships of two new genera and four new species of Oligo-Miocene waterfowl (Aves: Anatidae) from Australia Zoological Journal of the Linnean Society, v. 156(2), pp. 411-454

A NOTE: This publication is included on pages 177-257 in the print copy of the thesis held in the University of Adelaide Library. A It is also available online to authorised users at: A http://dx.doi.org/10.1111/j.1096-3642.2008.00483.x 254

SUPPLEMENTARY MATERIAL

COMPARATIVE MATERIAL The following comparative material was consulted during this study. All are extant species unless stated otherwise; sex (M or F), is indicated after the catalogue number of the specimen. Gallus gallus (Linnaeus, 1758, red : SAM B46451. Anhima cornuta (Linnaeus, 1766), : MV B12574. Anseranas semipalmata (Latham, 1798), magpie goose: AM O59362; SAM B48035, M; B39824, F. Cereopsis novaehollandiae Latham, 1802, Cape Barren goose: MNZ 25217, 25143; SAM B39638, B32830. Cnemiornis calcitrans Owen, 1866, South Island goose: fossil, notably MNZ S.35266; MV P160495. Cnemiornis gracilis Forbes, 1892, North Island goose: fossil, notably MNZ S.35683-706; MNZ S.35892. Branta canadensis (Linnaeus, 1758), Canada goose: MNZ 23745, 26738, 26739, 26740, 26741; SAM B31086, MV B6364, M. Anser brachyrhynchus Baillon, 1834, pink-footed goose: MV B25672. Anser caerulescens (Linnaeus, 1758), snow goose: SAM B36868 F. Anser anser (Linnaeus, 1758), greylag goose: MNZ 20812, 24519. Anser rossii (Cassin, 1861), Ross’s goose: MV B13340; Cygnus atratus (Latham, 1790), black swan: MNZ 15266, 15267, 17250; SAM B46110, M. Cygnus olor (Gmelin, 1789), mute swan: MNZ 16454. Dendrocygna arcuata (Horsfield, 1824), wandering whistling duck: AM O64697, M, O64697, M; ANWC 22213, M; ANWC 22214, M. Dendrocygna eytoni (Eyton, 1838), plumed whistling duck; MNZ 27024; SAM B45769, M. Dendrocygna bicolor, (Vieillot, 1816), fulvous whistling duck: SAM B36869; MV B13722. Dendrocygna arborea, (Linnaeus, 1758), West Indian whistling duck: USNM 226455, F. USNM 344488, F. Dendrocygna autumnalis (Linnaeus, 1758), black-bellied whistling duck: USNM 345767, M; USNM 345768, F (both are subspecies D. a. discolor). Thalassornis leuconotus Eyton, 1838, white-backed duck: BMNH 1901.10.20.156, F. Plectropterus gambensis (Linnaeus, 1766), spur-winged goose, CM Av33451. Oxyura australis Gould, 1836, blue- billed duck: CM Av31408; AM O65518; SAM B49214, M, B36390, F, B31910, F, B32842, ?. Nomonyx (Oxyura) dominicus, (Linnaeus, 1766), masked duck: USNM 430927, F; USNM 347324, M. Oxyura vittata, (Philippi, 1860), Argentine ruddy duck: USNM 614580, M. Oxyura maccoa, (Eyton, 1838), Maccoa duck: USNM 558442, M. Oxyura jamaicensis (Gmelin, 1789), ruddy duck: ANWC 22640 (ANSS 386), ANWC 22641 (ANSS 391); MNZ 27335; SAM B36872, F. Oxyura vantetsi Worthy, 2005, New Zealand blue-billed duck: Specimens (fossil) listed in Worthy (2004, 2005). Biziura lobata (Shaw, 255

1796), musk duck: MNZ 26190, 26191, CM Av7116; SAM B38592, M, B47221, M, B11405, M, B46027, F, B49170, F, B36393, F, B31087, F. Stictonetta naevosa (Gould, 1841), freckled duck: MNZ 25141; SAM B46743, M, B47950, M, B39659, M, B31585, F, B45045, F. Malacorhynchus membranaceus (Latham, 1802), Australian pink-eared duck: MNZ 23880, 23881; SAM B39384, M, B39385, F. Malacorhynchus scarletti Olson, 1977, Scarlett’s duck: fossil, MNZ, Lake Poukawa specimens listed in Worthy (2004); CM Av5855, Av14697, Av15044, Av36035, Av36285. Nettapus pulchellus (Gould, 1842), green pygmy-goose: MNZ 27025, 27026; SAM B45606; MV B14485, M, B14490, M. N. coromandelianus (Gmelin, 1789), cotton pygmy-goose: MV B20293, F; QM O.21080, QM O.21081. Tadorna ferriginea (Pallas, 1764), ruddy shelduck: SAM B38602. Tadorna tadornoides (Jardine & Selby, 1828), Australian shelduck: MNZ 22921, 23888a, 27367; ANWC 22240; SAM B39873, F, B39591, M. Tadorna tadorna (Linnaeus, 1758) common shelduck: MNZ 12280; MV B25679; ANWC 22408, M. Tadorna variegata (Gmelin, 1789), paradise shelduck: MNZ 15146, 16471, 16472, 16473, 16501, 16590, 24559, 25139, 25669, 26562, 26563; MV B5103, F. Tadorna radjah (Lesson, 1828), radjah shelduck: MNZ 26206, 26207; ANWC 22411, M. Alopochen aegyptiacus (Linnaeus, 1766), African sheldgoose: ANWC 22239 (ANSS 753), M; BMNH 1930.3.24.217, ?; MV B25678; MNZ 24283. Chloephaga poliocephala Sclater, 1857, ashy-headed sheldgoose: MV B13714, F. Chloephaga hybrida (Molina, 1782), kelp sheldgoose: MV B13227, M. Chloephaga picta (Gmelin, 1789), upland sheldgoose: BMNH 1860.11.4.15. Tachyeres leucocephalus Humphrey and Thompson, 1981, white-headed flightless steamer duck: MV B14144, M. Cairina moschata (Linnaeus, 1758), muscovy duck; MNZ 19842; SAM B32415. Chenonetta jubata (Latham, 1802), maned duck: MNZ 1487, 23188a, 25142, 25400, 25194a; SAM B37195, M, B39011, F. Chenonetta finschi (Van Beneden, 1875), Finsch’s duck: fossil, MNZ specimens, notably S.35885, S.23794, S.35885, S.38931, S.39838, S.39840, S.44279; MV P204243. Hymenolaimus malacorhynchos (Gmelin, 1789), blue duck: MNZ 16699, 23924, 23963; 24586, 24587. Somateria mollissima (Linnaeus, 1758), : MNZ 12277, 12278, 12279; SAM B48744, F. Melanitta fusca (Linnaeus, 1758), velvet scoter: SAM B47234, M. Clangula hyemalis (Linnaeus, 1758), long-tailed duck: SAM B47233, M. Bucephala albeola (Linnaeus, 1758), bufflehead: MNZ 12708, 12709. Bucephala clangula (Linnaeus, 1758), common goldeneye: MV B13553, M. Lophydytes cucullatus (Linnaeus, 1758), hooded merganser: MNZ 12706; MV B13541, M, SAM B47750, M. Mergus merganser Linnaeus, 1758, common merganser: MV B13320, M. Mergus serrator Linnaeus, 1758, red-breasted merganser: MNZ 12707; MV B14546, M. 256

Mergus australis Hombron & Jacquinot, 1841, Auckland Island merganser: MNZ S30046, S31777, collections of fossils from Chatham Island. Anas superciliosa Gmelin, 1789, gray duck: MNZ 13686, 15030, 16586, 18132, 16476, 17341, 17261, 16698, 16584, 24607; SAM B49654, M. Anas rhynchotis variegata (Gould, 1856), : MNZ 17000, 18971, 16591, 24588, 24589; SAM B37429, M. Anas castanea (Eyton, 1838), chestnut teal: SAM B32512, M; AM O.67041, M. Anas chlorotis G. R. Gray, 1845, brown teal: MNZ 14978, 15628, 15935 (= CM Av31828), 18898, 21544, 22086, 22802, 22806, 24535, 24536, 24537, 25105, 25106, 26630, 26631, 26940a, 26941a, 26942a, 26943a, 26944a, 26945a, 26946, 26947, 26949, 26950a, 26951a, 26952a. Anas aucklandica (Gray, 1849), Auckland Island teal: MNZ 24367, 24052. Anas nesiotis (Fleming, 1935), Campbell Island teal: MNZ 25727, 26742. Anas gracilis Buller, 1869, Australasian gray teal: MNZ 19351, 19348, 19301, 19322, 13688, 19323, 26815 (ex 24545), 18099, 19324, CM Av36764. Aix galericulata (Linnaeus, 1758), Mandarin duck: MNZ 27368; SAM B47841, M. Aythya fuligula (Linnaeus, 1758), tufted scaup: MV B12458. Aythya australis (Eyton, 1838), Australian white-eyed duck: AM O65772; SAM B32513. Aythya novaeseelandiae (Gmelin, 1789), New Zealand scaup: CM Av22382, Av22413; MNZ 8726, 13685, 16588, 16589, 17001, 17002, 17003, 23144, 24245; MV B19895, M. Aythya affinis (Eyton, 1838), lesser scaup: MNZ 24041; MV B25696, M. Netta rufina (Pallas, 1773), red-crested pochard: MV B13723, M.

Tertiary fossil taxa: Mionetta blanchardi (Milne-Edwards, 1863) fossil: Université Claude Bernard - Lyon, ex St. Gérand-le-Puy, France, all prefixed with FSL: 442.807 pmx 442.800 cran L hum 332.274, R hum 332.277, R ulna 331.783, R ulna 331.788, L cmc 331.497, R cmc 331.492, L cor 331.364, R cor 331.381, L fem 331.176, R fem 331.185, L tib 331.104, R tib 331.109, L tmt 331.005, R tmt 331.007; CM Av11394, 1 ant stern, pt cran, fur, LR scap, pt 2 pel, LR MII.1, 2L1R cmc, R rad, LdR ulna, 1L2R cor, 1L2R hum, LR tmt, 1L3R tt, 1L2R fem; CM Av13902, 2L6R cor, 1R3L hum, 2L1R cmc, pt fur. Manuherikia lacustrina Worthy et al., 2007: material as listed in Worthy et al., (2007). Dunstanetta johnstoneorum Worthy et al., 2007: material as listed in Worthy et al., (2007).

257

CONSTRAINT MATRIX USED IN BAYESIAN ANALYSES The constraint data matrix employed in MrBayes. Non-constrained taxa were interpolated in this matrix in the same order as in the real dataset, with all characters coded as ?

Gallus_gallus 000000000000000000000000000000000000000000000000000000000000 Anseranas 000000000000000000000000000000000000000000000000000000000000 Dendrocygna_arborea 000000000000000000001111111111000000000000000000001111111111 Dendrocygna_autumnalis 000000000000000000001111111111000000000000000000001111111111 Dendrocygna_bicolor 000000000000000000001111111111000000000000000000001111111111 Dendrocygna_eytoni 000000000000000000001111111111000000000000000000001111111111 Dendrocygna_arcuata 000000000000000000001111111111000000000000000000001111111111 Oxyura_vittata 000000000000000000000000000000111111111100000000001111111111 Oxyura_jamaicensis 000000000000000000000000000000111111111100000000001111111111 Oxyura_australis 000000000000000000000000000000111111111100000000001111111111 Oxyura_maccoa 000000000000000000000000000000111111111100000000001111111111 Stictonetta 000000000000000000000000000000000000000000000000001111111111 Branta_canadensis 000000000000000000000000000000000000000011111111111111111111 Anser_brachyrhynchus 000000000000000000000000000000000000000011111111111111111111 Cygnus_atratus 000000000000000000000000000000000000000011111111111111111111 Tadorna_ferriginea 000000000011111111110000000000000000000000000000001111111111 Tadorna_tadornoides 000000000011111111110000000000000000000000000000001111111111 Tadorna_tadorna 000000000011111111110000000000000000000000000000001111111111 Alopochen 000000000011111111110000000000000000000000000000001111111111 Somateria_mollisima 111111111111111111110000000000000000000000000000001111111111 Lophodytes_cucullatus 111111111111111111110000000000000000000000000000001111111111 Anas_superciliosa 111111111111111111110000000000000000000000000000001111111111 Aix_galericulata 111111111111111111110000000000000000000000000000001111111111 Aythya_australis 111111111111111111110000000000000000000000000000001111111111 Aythya_affinus 111111111111111111110000000000000000000000000000001111111111 Aythya_novaeseelandiae 111111111111111111110000000000000000000000000000001111111111

258

259

An Oligo-Miocene magpie goose Aves: Anseranatidae from Riversleigh, north western Queensland, Australia 

Journal of Vertebrate Paleontology (2008), submitted paper 260 261

 ‹

Running Title — — Anseranas semipalmata

A Worthy, T.H. & Scanlon, J.D. (2009) An Oligo-Miocene magpie goose Aves: Anseranatidae from Riversleigh, north western Queensland, Australia Journal of Vertebrate Paleontology, v. 29(1), pp. 205-211

A NOTE: This publication is included on pages 261-279 in the print copy of the thesis held in the University of Adelaide Library. 280

281

CHAPTER 7

Pliocene waterfowl (Aves: Anseriformes) from South Australia and a new genus and species

Trevor H. Worthy

School of Earth and Environmental Sciences, Darling Building DP 418, The University of Adelaide, North Terrace, AUSTRALIA 5005

The Emu (2008), accepted paper.

282

STATEMENT OF AUTHORSHIP

Pliocene waterfowl (Aves: Anseriformes) from South Australia and a new genus and species

Trevor H. Worthy

The Emu (2008), accepted paper.

Worthy, T. H. (Candidate) Was instrumental in research design, did the research, wrote the manuscript, and acted as corresponding author.

I hereby certify that the statement of contribution is accurate

Signed …………………………………………………………. Date ……………….

283

Pliocene waterfowl (Aves: Anseriformes) from South Australia and a new genus and species

Trevor H. Worthy Darling Building, DP 418, Department of Earth and Environmental Sciences, The University of Adelaide, Adelaide, SA 5005, Australia. E-mail: [email protected];

Abstract. The occurrences of fossil bones of waterfowl (Aves, Anseriformes) in Pliocene faunas from the Lake Eyre Basin are detailed. Nine modern taxa are present in either the Kanunka or the Toolapinna Faunas from the Tirari Formation as follows: Anseranas semipalmata, Cereopsis novaehollandiae, Cygnus atratus, Tadorna tadornoides, Biziura lobata, Oxyura australis, Anas cf A. castanea, A. cf A. gracilis and Aythya australis. A new genus and species of oxyurine Tirarinetta kanunka is described from the Kanunka Fauna on the basis of a humerus.

A Worthy, T.H. (2008) Pliocene waterfowl (Aves: Anseriformes) from South Australia and a new genus and species. Emu, v. 108(2), pp. 153-165

A NOTE: This publication is included on pages 283-310 in the print copy of the thesis held in the University of Adelaide Library. A It is also available online to authorised users at: A http://dx.doi.org/10.1071/MU07063 311 312 Transactions of the Royal Society of S. Aust. 131

A SHELDUCK (ANATIDAE: TADORNA) FROM THE PLIOCENE OF SOUTH AUSTRALIA ‹ ‹ Summary Tadorna Q KeywordsTadorna Running Head

A Worthy, T.H. & Pledge, N.S. (2007) A Shelduck (Anatidae: Tadorna) from the Pliocene of South Australia. Transactions of the Royal Society of South Australia, v. 131(1), pp. 107-115

A NOTE: This publication is included on pages 313-321 in the print copy of the thesis held in the University of Adelaide Library.

322

323

CHAPTER 9 Concluding Discussion

This thesis is submitted as a portfolio of publications, so the format of the discussion differs from that expected of a traditional thesis because detailed discussion of each piece of research is presented in each paper. Here the work presented in the preceding chapters is drawn together to show how each piece has contributed to the overall aim of the research programme. Its significance is identified and general conclusions are formed.

9.1 Summary of aims of the thesis

The overall objective of this thesis has been to determine the taxonomic diversity and phylogenetic relationships of the fossil waterfowl (Aves: Anseriformes) in the Oligo- Miocene and Pliocene faunas of New Zealand and Australia. This will allow patterns of evolutionary change in the region to be determined, in particular, when there were periods of major faunal turnover, and when the modern fauna evolved. Further, it would be instructive on the origin of modern taxa in a global context and may address the suggestion of a southern origin for Anseriformes (Olson 1989). Simultaneously, this research provides an objective test, based on new morphological and taxonomic information, of the hypothesis of anseriform phylogenetic relationships advanced by Livezey (1997a).

9.2 Description and phylogenetic interpretation of the waterfowl from the Early Miocene St Bathans Fauna from Otago, New Zealand.

In a series of papers, given here as Chapters 2-4, the St Bathans Fauna of Early Miocene age (19-16 Ma) is initially described and shown to have the richest Tertiary fauna of anseriforms in the world. Chapter 2 (Worthy et al. 2007) provides the first description of the St Bathans Fauna and the taxonomic description of four anatid genera with five species (Manuherikia, 2 sp.; Dunstanetta, 1 sp.; Matanas, 1 sp.; and Miotadorna, 1 sp.) based on humeri. In addition, a sixth undetermined anseriform taxon is noted based on a coracoid, and a further 18 taxa of non-anseriforms are reported, but only one, a Pelecanoides species, is named. In this initial study, Manuherikia is interpreted as a probable Dendrochenine, sensu Livezey & Martin (1988), based on limited characters. The vertebrate diversity of the St Bathans Fauna other than fish is now known to include frogs, skinks, geckoes, crocodilians, and mammals. Associated work has found the first terrestrial mammal known 324

from New Zealand, and one that is a stem taxon more derived than monotremes but more primitive than metatherians and therians (Appendix 1). Other work is in progress describing the bats (Hand et al. 2007, in prep).

Chapter 3 (Worthy & Lee 2008) provides comprehensive skeletal descriptions for, and investigates the phylogenetic relationships of, Manuherikia lacustrina, Dunstanetta johnstoneorum and Miotadorna sanctibathansi: taxa with the most complete skeletal representation (551, 7, and 115 specimens, respectively). The analyses were based on 133 morphological characters (128 skeletal, 5 integumental), chosen primarily to allow maximal representation in the fossil material.

Initial analyses of this dataset revealed homoplasy issues similar to those identified by McCracken et al. (1999) as having confounded the analyses leading to Livezey’s (1997a) phylogeny. Thus, all diving taxa were associated in a single clade, deeply nested in the tree, which is inconsistent with a range of genetic data, as Oxyura and Biziura are both now considered primitive anatids very divergent from aythyines and mergines (Frith 1964; Madsen et al. 1988; Sibley & Ahlquist 1990; Christidis & Boles 1994; Sraml et al. 1996; Livezey 1997a; Johnson & Sorenson 1999; Sorenson et al. 1999; Donne-Goussé et al. 2002; Dickinson 2003; McCracken & Sorenson 2005; Callaghan & Harshman 2005). Attempts to mitigate this homoplasy issue by exclusion or weighting of characters observed to be correlated with diving habit failed to overcome the problem. Homoplasy was thus more subtle, and not obvious to exclude, as was found to be the case in a study of paedomorphosis (Wiens et al. 2005). Therefore, in this study a simple phylogenetic constraint was employed based on strongly supported genetic data. This had the primary effect of constraining Oxyura to an unspecified position below tadornines, and selected aythyines and mergines to an unspecified place above tadornines with anatines. Notably, taxa like Biziura, Thalassornis, other mergines and aythyines, and none of the fossils were constrained. They were therefore free to associate in clades dependent on their phylogenetic signal. In parsimony analyses of the data so constrained, there was reasonable support for the inclusion of Manuherikia and Dunstanetta in a clade of oxyurine taxa, together with the European taxon Mionetta (e.g. Chapter 3, fig. 9). In analyses with 21 characters associated with diving adaptations removed, Manuherikia and Dunstanetta form lineages of approximate oxyurine grade either side of the oxyurine clade (Chapter 3, fig. 10). No support was found for the taxon Dendrocheninae erected by Livezey & Martin (1988) for 325

Mionetta, and this subfamily was synonymised in Oxyurinae. In these analyses, Miotadorna was verified as a tadornine, closely related to extant Tadorna species.

Chapter 4 (Worthy et al. in press) describes more recently found fossils that reveal a third species of Manuherikia and confirmed the presence of anserines in the St Bathans Fauna. This established a total of eight anseriforms in the St Bathans Fauna, making it more diverse than in any other Oligo-Miocene fauna. The very rich Late Oligocene–Early Miocene lacustrine fauna from St Gérand Le Puy, France, is probably the richest European fauna and is of similar lacustrine nature, but it has just three species of Mionetta and a probable anserine Cygnopterus alphonsi (Cheneval 1983, 1984). In Australia, the Late Oligocene anseriform faunas from the Etadunna and Namba Formations, c. 26-24 Ma (Woodburne et al. 1994), contain three oxyurines and a tadornine (Chapter 5). However, the sample size of birds from these faunas is much smaller than that for the St Bathans Fauna, and given the relative rarity of anserines, Matanas and Dunstanetta in the St Bathans Fauna, the known diversity in the Australian faunas is probably under-estimated. Only in Pliocene faunas are there found more diverse anseriform assemblages, such as that from Lee Creek Mine, North Carolina, that is derived from >10,000 fossils and is dominated by extant taxa (Olson & Rasmussen 2001).

While anserines were confirmed to be present in the St Bathans Fauna (Chapter 4), in the absence of humeral fossils, which individually are the most informative element, no description of a taxon was made. However, some of the fossils are more similar to Cereopsis than to other anserines, suggesting that they represent a probable Cereopsis/Cnemiornis ancestor. Cnemiornis, a giant flightless form that became extinct in New Zealand in the Holocene, has been argued to be the sister taxon to Cereopsis, the Cape Barren goose of Australia (Owen 1866; Worthy et al. 1997), although Livezey (1989a, 1997a) considered it to be of a separate and more basal lineage than anserines. These new fossils, therefore, suggest that the Cereopsis/Cnemiornis lineage has been present in Australasia for 19-16 Ma, and as such predates the emergence of true geese (Anser and Branta) in the Northern Hemisphere. A close relationship between Cereopsis and Coscoroba to the exclusion of other anserines (Donne-Goussé et al. 2002; St. John et al. 2005), together with these fossil data, thus support a radiation of geese in the Southern Hemisphere separate from that in the north.

326

9.3 Description and phylogenetic interpretation of the waterfowl from the Oligo- Miocene of Australia.

Chapters 5 and 6 establish the taxonomic diversity and relationships of anseriforms in the latest Oligocene and Miocene of Australia. Avifaunas are limited for this period in Australia, with the best known being from the Etadunna Formation at Lake Palankarinna in the Lake Eyre Basin and its temporal equivalent at Lake Pinpa, the Namba Formation of the Lake Frome Basin; these are about 26-24 Ma (Rich & Van Tets 1982; Pledge & Tedford 1990; Rich et al. 1991; Vickers-Rich 1991; Woodburne et al. 1994). These lacustrine South Australian faunas overlap in age those from the Carl Creek Limestone at Riversleigh in northwestern Queensland (Archer et al. 1997, 1999, 2006), but there birds are relatively few, and waterbirds poorly represented (Boles 1993a-c, 1995b, 1997b-d, 1998, 1999, 2001, 2005a-c).

Only two significant faunas for birds are known from the Middle–Late Miocene of Australia: the 12-10 Ma Bullock Creek LF from the Camfield Beds at Bullock Creek, and the c. 8 Ma Alcoota LF from the Waite Formation at Alcoota, both in the Northern Territory (Murray & Megirian 1992; Archer et al. 1999; Megirian et al. 2004).

In the Oligo-Miocene faunas, a radiation of three species of oxyurines in one genus (Pinpanetta) and a large tadornine (Australotadorna) are described (Chapter 5). The Riversleigh faunas contain very few anseriform bones, but a single humerus from Ringtail Site, Faunal Zone C (Travoullion et al. 2006), is referred to one of the species otherwise found in the Etadunna and Namba formations. Faunal Zone C sites are considered to be somewhat younger than the Etadunna so if the same species is represented in both places then it existed for several million years. Alternatively, Ringtail Site may contain a local fauna that is older than most attributed to this Faunal Zone.

Phylogenetic analyses in Chapter 5 assess the relationships of the Pinpanetta species using an expanded dataset (seven more humeral and ten pelvic characters, for a total of 150), and an enlarged set of ingroup taxa (61) from that used in Chapter 3. Initial parsimony analyses, employing the same phylogenetic constraints as in Chapter 3, were followed by Bayesian analyses on the same dataset. These analyses confirmed Pinpanetta was a member of the oxyurine lineage despite all three species being relatively poorly represented compared to the New Zealand fossil taxa – none of the species had more than 28 specimens and cranial material was lacking entirely. Pinpanetta was found to take a position in the 327

oxyurine lineage more derived than Stictonetta, Malacorhynchus and Mionetta, but basal to the New Zealand Tertiary fossils Manuherikia and Dunstanetta and to the extant oxyurines (Oxyura, Nomonyx, Biziura and Thalassornis). This more basal position than the New Zealand fossils is consistent with the older age of the Australian fossils. The larger dataset used in these analyses provided stronger support for the clade of oxyurines, which was recovered both when all characters were unordered and when a subset of characters were ordered as morphoclines. Low bootstrap and Bayesian support for this clade is due in part to the inclusion of fossil taxa with much missing data (e.g. Dunstanetta), so as these fossil taxa become better known it is likely support for the clade will markedly improve.

A tadornine, described in Chapter 5 from Lake Pinpa, is probably also represented by fragmentary material from Lake Palankarinna. The Pinpa material includes a near complete humerus allowing its unequivocal referral to Tadorninae and reveals that it is quite distinct from Miotadorna known from the slightly younger St Bathans Fauna of New Zealand. It is thus the oldest tadornine known, some 5-10 Ma older than Miotadorna and 10 Ma older than the undescribed possible tadornines from Nördlinger Ries in Germany. Its presence strongly supports the suggestion that tadornines evolved in the Southern Hemisphere (Chapter 3; Kear & Murton 1976). Anser scaldii, originally described as of Miocene age from Belgium, is determined to be a Late Pleistocene fossil and referable to a Recent taxon; it is not a tadornine, contra the possibility raised in Chapter 2, see Appendix 2. It therefore has no bearing on the early evolution of tadornines.

An important discovery in the Faunal Zone A (c. 25 Ma) Sticky Site, at Riversleigh, revealed the first occurrence of fossil anseranatids in Australia. As mentioned in Chapter 1, the Magpie Goose is the sole living representative of the Anseranatidae but, to date, has no Tertiary fossil record in Australia. The family has been reported from Eocene sites in England and France (Olson 1999; Mourer-Chauviré et al. 2004), so a Tertiary record in Australia is important to delimit the time Anseranas has been a member of Australian faunas. These new fossils are described in Chapter 6 as a new genus Eoanseranas, but synapomorphies with Anseranas suggest that this lineage has existed in Australia for at least the last 25 Ma, thus considerably extending its age range in Australia from the previous oldest record in the 5-4 Ma Early Pliocene Chinchilla Sands (Rich et al. 1991; Boles 2006).

Anseriformes from the Middle to Late Miocene of Australia are so far only known from the Alcoota LF, and then only from three bones. They are relatively uninformative 328

elements and indicate merely the presence of an undetermined tadornine and a small anatid about the size of Malacorhynchus.

9.4 Description and phylogenetic interpretation of the waterfowl from the Pliocene of Australia.

Pliocene avifaunas containing anseriforms are, as for earlier Tertiary ones, also of limited number in Australia. Those from Bluff Downs and Chinchilla in northwest Queensland have been described previously (Rich & Van Tets 1982; Baird et al. 1991; Rich et al. 1991; Boles & Mackness 2004). While several extinct taxa were named by De Vis from the Chinchilla LF (De Vis 1888, 1889, 1905), all have been subsequently referred to modern taxa (Olson 1977b; Rich & Van Tets 1982; Van Tets & Rich 1990; Rich et al. 1991).

In South Australia, the Pliocene is represented by faunas in the Tirari Formation derived from the sediments at Lake Palankarinna (Mampuwordu Member, Palankarinna Fauna) and Lake Kanunka (Kanunka Fauna) (Tedford et al. 1986; Tedford & Wells 1990; Tedford et al. 1992). These South Australian Pliocene faunas are described in Chapter 7. Nine modern taxa are present in either the Kanunka or the Toolapinna Faunas as follows: Anseranas semipalmata, Cereopsis novaehollandiae, Cygnus atratus, Tadorna tadornoides, Biziura lobata, Oxyura australis, Anas cf A. castanea, A. cf A. gracilis and Aythya australis. A new genus and species of oxyurine (Tirarinetta kanunka) is described from the Kanunka Fauna, based on a humerus. It was the only extinct waterfowl taxon found in Pliocene avifaunas.

In Chapter 8, an isolated fossil tadornine humerus from the Late Pliocene Parilla Sands, at Bookmark Cliffs on the Murray River, is described as Tadorna cf T. tadornoides.

The South Australian Pliocene avifaunas are more taxonomically diverse than the preceding Miocene ones, but the total numbers of individuals in this avifaunal assemblage is still small. Ten species in nine genera, recorded in Chapter 7 from just 19 specimens will, therefore, certainly under-estimate the diversity in the source biota. Modern faunas in the Lake Eyre Basin region include minimally the following additional taxa Malacorhynchus membranaceus, Stictonetta naevosa, Anas superciliosa, A. rhynchotis, Dendrocygna eytoni, and Chenonetta jubata (Marchant & Higgins 1990; Reid et al. 1990). The main result from this analysis is that Australian Pliocene anseriform faunas are dominated by Recent species. 329

Only one of 10 species is extinct. This contrasts markedly with Early Miocene anseriforms, four species/two genera in 61 specimens, which is therefore a considerably less diverse fauna. Thus, whereas Rich (1976) did not have the data to determine the timing of, or indeed if there had been, an avifaunal turnover in the Tertiary, the present data show that such a turnover occurred in the Late Miocene. The Early Miocene St Bathans Fauna of New Zealand is now known to have six species in four genera, plus a further two undescribed species/genera derived from an anseriform assemblage of near 2000 specimens. This Miocene waterfowl fauna has a very different composition to the Recent New Zealand fauna, also attesting to substantial turnover.

The trend of Pliocene faunas being dominated by modern taxa is found in other regions of the world. Howard’s (1964) list of 28 species of Pliocene anseriforms, including three genera and 18 species that were extinct, may under-estimate the presence of modern taxa in fossil avifaunas for two reasons: 1, faunal analyses of Pliocene assemblages show a predominance of extant species (see below); 2, a number of Pliocene fossils, named as extinct taxa, have been transferred more recently to modern taxa (Chapter 1). Perhaps the most diverse Pliocene fauna known is that from the Early Pliocene Yorktown Formation (4.8-3.7 Ma) in Lee Creek Mine, North Carolina. From this site, Olson & Rasmussen (2001) recorded 18 species of waterfowl in 10 genera, of which only one species was extinct. The San Diego Formation, California, 3.0-1.8 Ma, has two recorded anatid taxa, both of which are extant (Chandler 1990; Olson & Rasmussen 2001). A 2.5-2.0 Ma fauna from Florida included 10 extant species (five genera) and three extinct species (one extant, two extinct genera) (Emslie 1992). In contrast, the Big Sandy Formation, Arizona, 6.1-4.6 Ma (Late Miocene) has at least eight anatid taxa (five genera) listed from it, but only three undetermined Anas species are possibly modern species (Bickart 1990). These examples show that Pliocene avifaunas contrast markedly with Miocene ones by being dominated by modern genera and species.

These observations from the palaeontological record suggest that species longevity in anseriforms often may be in the range 2.5-4.8 Ma. Such an age range is borne out by analyses of mitochondrial genetic data among modern sister taxa of birds, assuming a conventional avian molecular clock of 2% divergence between lineage pairs per million years. Such estimates for allopatric and phylogeographically distinct pairs of conspecific avian populations yielded 17 divergence dates in the range 1.0-4.2 Ma out of 37 taxa investigated (Avise & Walker 1998). While some of these data relate to allopatric taxa now 330

considered sister species, e.g. Branta canadensis at 1.1 Ma (Sangster et al. 2005), most are for conspecific lineages, indicating that species longevity is likely to be longer in all cases where sister taxa are morphologically divergent species. Moreover, the mtDNA data suggests speciation may be protracted over several million years, although some speciation events have occurred in the last 250,000 years (Klicka & Zink 1997; Avise & Walker 1998; Johnson & Cicero 2003). Such protracted speciation events is consistent with slight morphological differences noted by Olson & Rasmussen (2001) in Early Pliocene modern taxa.

9.5 The identification of the extent and timing of faunal turnover and the origin of the modern fauna.

Australasia is established as a centre of anseriform diversity in the Late Oligocene, with an anseranatid, a radiation of oxyurines, a tadornid, and other large undetermined anatids now known to have been present. Anserines were also probably present, given the presence of a Cereopsis-like taxon in the Early Miocene of New Zealand. This contrasts with the diversity in Europe, which is known from comparatively rich fossil deposits, especially from St Gérand Le Puy in France (Cheneval 1983, 1984), to have at most three oxyurines and an anserine in the uppermost Oligocene, and supports the suggestion by Olson (1989) that anseriforms had a southern origin.

The data presented in Chapters 2-8 suggests that anseriform diversity increased in Australasia from the Early Miocene through to the Middle-Late Miocene, with modern diversity achieved by the Pliocene. Anseriform diversity is similarly low in the fauna from St Gérand Le Puy (Cheneval 1983, 1984), but by the close of the Miocene, the European diversity of anseriforms was approaching that of the present (Mlíkovský 2002; Chapter 1). A similar pattern is found in Asia and in the North American fauna (Mlíkovský & Švec 1986; Chapter 1; and above).

Clearly, the Middle-Late Miocene is an important period in the evolution of Anseriformes, so the absence of Australian faunas of this age containing significant anseriform fossils is unfortunate. Discovery and description of faunas of this age will therefore be most significant in understanding the evolution of this group.

The dramatic climate changes experienced during the Pleistocene and the concomitant aridification of Australia (e.g. White 2006) appear to have had minimal impact 331

on the diversity of anseriforms. Only the single species of oxyurine Tirarinetta kanunka is known to have become extinct in this period. among other birds are also few, and mostly affected the – four flamingos and one palaelodid (Miller 1963; Rich et al. 1987; Baird & Vickers-Rich 1998) – but also included a small pelican, (Rich & van Tets 1981). It seems likely that the aridification of Central Australia precipitated these extinctions, with complete drying of Lake Eyre for an extended period, first in the penultimate glaciation 150-130 ka (OIS 6), and with subsequent dry phases in OIS 3 (c.50 ka) and OIS 3-2 (30-12 ka) (Nanson et al. 1992, 1998; Magee et al. 1995; Magee & Miller 1998; Magee et al. 2004; Maroulis et al. 2007). All were relatively large-bodied taxa, and the phoenicopteriforms, of which living taxa are specialist filter feeders in shallow lakes and colonial, would be most at risk from drying of lakes. The impact of these environmental changes on other avian groups, such as Pelecaniformes, Gruiformes, and Charadriiformes, is not known, because their taxonomic biodiversity has not been the subject of any comprehensive analyses.

The apparent evolutionary constancy within waterfowl over the last 4 Ma contrasts markedly with mammals in Australia in which there has been significant evolution of taxa and associated faunas in the same time period. Such widely differing groups as Diprotodontidae, Macropodinae and rodents diversified greatly during the Pliocene (e.g. as reviewed in Rich et al. 1991; Aplin 2006; Archer & Hand 2006; Cooke 2006). Why there should be this contrast with birds is a matter for some conjecture. The first appearances of colubrid snakes (Scanlon 2006), and rodents (Aplin 2006), and major turnover in bat communities (Hand 2006), in the Early Pliocene, signal the first significant period of extensive faunal interchange with the Northern Hemisphere since Australia was part of Gondwana. This explains diversification of rodents but does not well explain rapid speciation in the large browsing diprotodontids or macropodines. Also, the first occurrences of (Ephippiorhynchus), ibis (Threskiornis), swamphens (Porphyrio), swans (Cygnus) and anatines (Anas and Aythya) in the Early Pliocene of Australia (Boles 2006; Chapter 7), indicates interchange with avifaunas from the north at this time also, yet there is no rapid evolution apparent in avian waterfowl lineages. While it is well known that avifaunas undergo rapid speciation of taxa on islands (e.g. Slikas et al. 2002; Worthy & Wragg 2003; Steadman 2006, and references therein), continental avifaunas reveal little speciation during the Pliocene (e.g. Bickart 1990; Chandler 1990; Olson & Rasmussen 2001). It is tempting to speculate that the continent-wide mobility necessary for waterfowl to survive while 332

exploiting stochiastic and widely separated water resources has not allowed diversification by allopatric speciation. The exception is allopatric speciation within genera shared between Australia and New Zealand (e.g. Oxyura, Chenonetta, Malacorhynchus and Biziura). Otherwise, the Australian fauna is marked by low diversity genera.

9.6 Towards a new hypothesis of generic relationships within anseriforms: Is there congruence with Livezey’s (1997a) morphology-based phylogenetic hypothesis?

Unlike many other groups of birds, such as galliforms (Crowe et al. 2006), gruiforms (Fain et al. 2007) and accipitrids (Griffiths et al. 2007), there is as yet no comprehensive genetic-based phylogenetic analysis of anatid genera. There are a number of genetic analyses based on limited subsets of taxa (e.g. Sraml et al. 1996; Johnson & Sorenson 1999; McCracken et al. 1999; Sorenson et al. 1999; Donne-Goussé et al. 2002; McCracken & Sorenson 2005), but none include either a majority of genera or all the relatively basal taxa of interest here. Moreover, all of these genetic studies had relatively small datasets based mainly on 1–3 mitochondrial genes. In the absence of a comprehensive phylogeny based on genetic data, the morphology-based phylogeny advanced by Livezey (1997a), which was derived from a series of analyses of constituent anseriforms groups (Livezey 1986, 1989ab, 1991, 1995ab, 1996a-c, 1997c), has gained general acceptance, with the exceptions of a few taxa (e.g. Callaghan & Harshman 2005). There is, however, much evidence in available mitochondrial data for contrary generic relationships of some taxa, such as the placement of Cereopsis in anserines next to Coscoroba (Donne-Goussé et al. 2002) and indications that Oxyura, Biziura and Malacorhynchus are all relatively basal taxa (e.g. Sraml et al. 1996; McCracken et al. 1999; McCracken & Sorenson 2005). It is therefore important to independently test Livezey’s (1997a) hypothesis of anseriform phylogenetic relationships based on a new analysis of morphological data. Analyses described in Chapters 3 and 5, while primarily to ascertain the phylogenetic relationships of fossil taxa, developed a new hypothesis of anatid generic relationships, summarized in Figure 9.1. 333

Fig. 9.1. The consensus tree obtained by Bayesian analysis of the data set with Presbyornis included, as reported in Chapter 5. The phylogenetic constraint used in Chapter 3 is employed and 35 characters were treated as ordered. Posterior probabilities or credibility values are shown above 334

the lines for clades if >0.50; Bootstrap values for the same clades, if >50%, are given below the lines.

The tree shown in Figure 9.1, which was derived from the most inclusive data set in terms of taxa and characters and the favoured analyses (Chapter 5), provides strong support for some quite different generic arrangements from those proposed by Livezey (1997a). This analysis was based on a substantially different set of characters and taxa to those used by Livezey, so it provides a robust test of his hypothesis. Presbyornis was included in the analysis to further test the dataset, as previous inferences about its relationships were based on other very different character sets.

9.6.1 Relationships of Presbyornis

Presbyornis, geological range Cretaceous-Eocene, was confirmed as sister to Anatidae (above Anseranatidae), as found previously (Ericson 1997c; Livezey 1997b). As the oldest member of Anatidae is Romainvillia, of Late Eocene age (Mayr in press), it is possible that Anatidae is derived directly from presbyornithid-like ancestors, as suggested by Olson & Feduccia (1980). An Eocene origin (56-34 Ma) for Anatidae at first would appear to have little support from analyses of genetic data, as studies such as Pereira & Baker (2006), Slack et al. (2006) and Brown et al. (2008), suggest Anatidae split from Anseranatidae deep in the Cretaceous. But, these studies actually suggest the lineage leading to Anatidae split from its common ancestor with Anseranatidae at that time. Because Presbyornithidae is sister to Anatidae, the origin of Anatidae relates to the age of their common ancestor, and so may be very much younger than the Anatidae-Anseranas split. In fact, as all autapomorphies of Presbyornithidae are plesiomorphic within Anseriformes (Ericson 1997), then the possibility that Anatidae has its common ancestor within Presbyornithidae cannot be excluded, in which case Anatidae may have originated as recently as in the Eocene. The divergence time of 60 Ma for Dendrocygna, the basal anatid in Brown et al.’s (2008) analysis, may constrain the origin of Anatidae to between 100 and 60 Ma. This may, however, be an over-estimate because divergence dates derived from molecular analyses can be very much younger depending on the methods used (e.g. Ericson et al. 2006). Estimates are likely to vary considerably in future as techniques develop. At present, the fossil record, which reveals the most basal anatid (Romainvillia) in the Late 335

Eocene MP 20 (c. 34 Ma) (Mayr in press), provides the best data on the origin of Anatidae, and suggests it derived from within Presbyornithidae during the Eocene.

9.6.2 Relationships of Cnemiornis and Cereopsis

Livezey (1997a) concluded that Cnemiornis was a basal anseriform and placed it in Cnemiornithidae as sister to Anatidae. Cereopsis was placed in its own tribe, sister to the rest of Anserinae, and Cereopsis and Cnemiornis were separated in the linear sequence by Dendrocygninae and Dendrocheninae. This contrasts with the long held view that Cereopsis and Cnemiornis are closely related taxa, (e.g. Owen 1866; Sharpe 1899; Shufeldt 1913; Phillips 1923; Worthy et al. 1997). Cereopsis has been listed next to geese in its own subfamily (e.g. Shufeldt 1913, 1914; Phillips 1923; Peters 1931), or tribe (e.g. Carboneras 1992), but is now widely considered part of Anserinae (Johnsgard 1961; Woolfenden 1961; Mayr & Cottrell 1979; Clements 1981; Marchant & Higgins 1990; Dickinson 2003; Christidis & Boles 2004, 2008; Callaghan & Harshman 2005), although Condon (1975) placed Cereopsis in Cereopsinae separated from Cygninae by Stictonettinae in the linear sequence. In a new arrangement based on DNA-DNA hybridization, Sibley & Ahlquist (1990) and Sibley & Monroe (1990) included Cereopsis with Anser and Branta in their eclectic Anserini, which in a departure from convention, was listed first in Anatinae, and so geese were separated from Cygninae, which was listed first in Anatidae.

In the analyses presented here, Cnemiornis and Cereopsis were found to be successive lineages of basal anatids below other anserines and more derived than Presbyornis. It is noteworthy, however, that when characters demonstrably related to diving were removed (Chapter 3), these three lineages formed a single clade. A number of shared synapomorphies for Cnemiornis and Cereopsis cannot so easily be dismissed either: 1, the tympanic cavity is closed anteriorly by the fusion of the zygomatic process with the ala parasphenoidalis (elsewhere in Anatidae, seen only in Malacorhynchus); 2, the palatines are convex laterally; 3, the premaxilla lacks a posterior process extending under the jugal; 4, there is a large pneumatic foramen laterally at the base of the ischium; 5, the procoracoidal foramen is large; 6, and the acromion on the scapula is reduced (Worthy et al. 1997). Given these synapomorphies, of which 2 and 3 were not included in the current analyses, it seems likely that modifications towards flightlessness and large size in Cnemiornis have obscured 336

phylogenetic signal in the data set used. On balance it is likely that Cnemiornis and Cereopsis are indeed sister taxa forming a clade sister to other anserines.

Further resolution of this issue will benefit from inclusion in analyses of the South American endemic ‘swan’ Coscoroba, which has recently been shown to be sister to Cereopsis (Donne-Goussé et al. 2002; St. John et al. 2005; Callaghan, in Kear 2005: 257). Moreover, this relationship and the discovery of anserines in the St Bathans Fauna (19-16 Ma) that appear to have greater affinity to Cereopsis than to Northern Hemisphere geese (Chapter 4) suggest a southern radiation of geese long separated from that in the north. True geese, Anser and Branta, and cygnines only make an appearance in the Northern Hemisphere record about the Middle Miocene and are preceded there by the enigmatic Cygnopterus (Late Oligocene–Early Miocene), which is in need of a phylogenetic analysis of its relationships. Swans do not appear in the Australian record until the Pliocene (Chapter 7), at which time there were faunal influxes into Australia of bats (Hand 2006), rodents (Aplin 2006), and colubrid snakes (Scanlon 2006), suggesting that until then Australia was well separated from northern continents. Thus any resolution of phylogenetic understanding of anserines, whether based on genetic or morphological data, will need to include Coscoroba and Cereopsis to capture this Southern Hemisphere radiation. In addition, because large size and flightlessness have likely obscured much of the phylogenetic signal in the morphology of Cnemiornis, its relationships should be further explored with recourse to more genetic data. In particular, a search for the synapomorphic presence the ‘chicken repeat 1’ nuclear gene found in Coscoroba and Cereopsis (St. John et al. 2005) could be made to test the sister group relationship with Cereopsis found by Worthy et al. (1997). Discovery of additional fossils of the Cereopsis-like anserine in the St Bathans Fauna is also likely to be important in resolving these relationships in the same way as inclusion of fossil oxyurine taxa has helped resolve that group (see below).

9.6.3 Relative position of Dendrocygna

Livezey (1997a) placed Dendrocygna in its own tribe, sister to Thalassornithini, in the subfamily Dendrocygninae, as the most basal member of Anatidae. The analyses presented in Chapters 3 and 5, in contrast, consistently found anserines to be more basal than Dendrocygna with strong Bootstrap and Bayesian support (e.g., Figure 9.1). Such a relatively basal position for anserines (variously as either tribes or subfamilies) was 337

generally accepted until the 1960s (e.g. Sharpe 1899; Shufeldt 1913, 1914; Phillips 1923; Peters 1931; Delacour & Mayr 1945; Verheyen 1955a-d; Brodkorb 1964). Johnsgard (1961, 1968, 1978) modified this arrangement, so that Anserinae included, successively, Dendrocygna, Thalassornis, Cygnus, Coscoroba, Anser, Branta, Cereopsis and Stictonetta.

The current trend to list Dendrocygna basal in Anatidae (e.g. Mayr & Cottrell 1979; Marchant & Higgins 1990; Carboneras 1992; Christidis & Boles 1994, 2008; Dickinson 2003; Callaghan & Harshman 2005) derives from two influential works in 1961. Woolfenden (1961: 100) listed Dendrocygna most basal as it shared several skeletal characters with Anseranas. The same relationship was found by Johnsgard (1961) in a study based on behavioural characters. The basal placement of Dendrocygna was further reinforced by the referral of Thalassornis to Dendrocygninae on the basis of shared reticulate tarsi, plain-coloured downy young, tracheal structure and certain skeletal features, such as the straight dorsolateral margin of the quadrate (Johnsgard 1968; Raikow 1971). Further support was found for this basal position of Dendrocygna and Thalassornis with DNA-DNA hybridization data (Sibley et al. 1988): Dendrocygnidae was placed before Anatidae (Sibley & Ahlquist 1990; Sibley & Monroe 1990).

Livezey (1986), using many characters derived from Woolfenden (1961), found only weak support for the basal position of Dendrocygna (3 synapomorphies). Subsequently, he found in an analysis including Cnemiornis, a different relationship: Cnemiornis was the most basal taxon, and other Anserinae were sister to the remaining Anatidae (Livezey 1989a). But, if Cnemiornis was excluded, the position of anserines was unresolved with their position above or below Dendrocygna equally likely. Similarly, the sequence of these taxa was unresolved in the Majority-rule bootstrapped consensus tree obtained by Livezey (1996a) while investigating anserine relationships. Thus, Livezey’s data did not provide robust support for his preferred placement of Dendrocygna basal to anserines (Livezey 1997a).

The analyses presented in Chapters 3 and 5 revealed that all the characters listed by Woolfenden linking Dendrocygna to Anseranas have a wider distribution than was stated, and most are plesiomorphic and so not necessarily evidence for a close relationship. Similarly, the features listed by Raikow (1971) to align Thalassornis with Dendrocygna are also plesiomorphic (Chapter 3, 5), and are found in other basal oxyurines as herein defined. Significantly, oxyurines were not included in several of the pertinent analyses (e.g. Livezey 1995b, 1996a) that led to the phylogeny advanced by Livezey (1997a). In summary, while 338

the previous evidence for a basal position for Dendrocygna is weak, the present analysis provides robust support for a basal position of anserines below Dendrocygna (Figure 9.1). Moreover, the emergence in the fossil record of anserines as the oldest crown-group anatids supports this position.

9.6.4 Relationships of Plectropterus

Plectropterus was placed in a tribe early in Tadorninae by Livezey (1997a). The possible relationship of Plectropterus as sister to Dendrocygna found in Chapter 5 is new. Its affinity has been problematic in the past: Peters (1931) placed it in Plectropterinae, with Anseranas, between Cygninae and Cereopsinae. It has sometimes been included with the ‘perching ducks’ Cairinini (e.g. Delacour & Mayr 1945; Johnsgard 1961; Carboneras 1992), and in Anatinae, separate from Tadorninae (Mayr & Cottrell 1979). Recently, it has usually been related to members of Tadorninae (Woolfenden 1961; Sibley & Ahlquist 1990; Sibley & Monroe 1990; Dickinson 2003), but Livezey (1986) placed it in a monotypic subfamily Plectropterinae. Most recently, it has been treated as a monotypic genus and tribe of uncertain relationships in Tadorninae (e.g. Livezey 1997a; Callaghan & Harshman 2005). Livezey’s (1996b) analysis that addressed Plectropterus affinities did not include any Dendrocygna species, so the relationship found here (Chapter 5; Figure 9.1), was not explored.

The attraction of Plectropterus to Dendrocygna in my analyses may result from homoplasy related to their similar large terrestrial morphotype (Chapter 5). The presence in Plectropterus of such highly conservative characters as the scutellated tarsi and an asymmetrical syringeal bulla (Livezey 1996b), both lacking in Dendrocygna, is strong evidence against a close relationship between these taxa. Nevertheless, there appears to be a strong phylogenetic signal in the data for a relatively basal position for Plectropterus, so any further analyses exploring its relationships should include representative taxa of those more primitive than tadornines.

9.6.5 The oxyurine radiation

The most significant insight into anseriform relationships found in this thesis is the concept of a single clade of diverse oxyurines that includes most well known fossil Neogene fossil ‘ducks’. The hitherto difficult to classify Stictonetta was identified as the most basal 339

taxon in the clade, followed successively by the European Mionetta, one of the oldest fossil taxa, and then Malacorhynchus. More deeply nested in this clade is a grade of the Australasian Oligo-Miocene fossils (Pinpanetta, Dunstanetta, and Manuherikia), then as sister to Manuherikia, is a clade of the extant highly specialised divers Nomonyx+Oxyura and Biziura+Thalassornis.

Previously, Stictonetta has been classed as an aberrant anatine (Peters 1931; Delacour & Mayr 1945), included in Anserinae (Johnsgard 1961, 1968, 1978; Mayr & Cottrell 1979; Carboneras 1992), listed in its own subfamily (Condon 1975; Sibley & Ahlquist 1990; Sibley & Monroe 1990; Christidis & Boles 1994; Callaghan & Harshman 2005), or included in Oxyurinae (Marchant & Higgins 1990).

The relationships of Malacorhynchus have been equally problematic, and it is usually listed as an aberrant anatine (Peters 1931; Mayr & Cottrell 1979; Delacour & Mayr 1945; Woolfenden 1961; Johnsgard 1961, 1968, 1978; Condon 1975; Marchant & Higgins 1990; Turbott 1990; Dickinson 2003; Christidis & Boles 1994; Callaghan & Harshman 2005).

Oxyura, Nomonyx, Biziura, and often Thalassornis, have generally been associated together and placed at the end of Anatinae (e.g. Delacour & Mayr 1945; Woolfenden 1961; Brodkorb 1964; Johnsgard 1961, 1968; Clements 1981; Carboneras 1992; Dickinson 2003), or after Anatinae, in Oxyurinae (Peters 1931; Condon 1975; Mayr & Cottrell 1979; Turbott 1990). As noted above, Thalassornis has, since the 1970s, been placed in Dendrocygninae following Johnsgard (1968) and Raikow (1971) (e.g. Mayr & Cottrell 1979; Sibley & Monroe 1990). The placement of Oxyurinae between Dendrocygna and anserines (Sibley et al. 1988; Sibley & Ahlquist 1990; Sibley & Monroe 1990) was followed by Marchant & Higgins (1990) and Christidis & Boles (1994). Callaghan & Harshman (2005), following McCracken et al. (1999), placed Biziura incertae sedis in Anatidae, and Oxyura, Nomonyx, and Heteronetta in Oxyurinae after Stictonettinae and before Tadorninae. Most recently, Christidis & Boles (2008) retained Biziura as a basal monotypic anatid, but placed Oxyura tentatively at the end of Anatinae. Clearly, the taxonomic affinities of these groups are far from settled.

Livezey’s (1997a) phylogenetic classification had the various taxa recognised herein as oxyurines widely separated in the linear sequence. Thalassornis (Thalassornithini) was included in Dendrocygninae; Stictonetta (Stictonettinae) followed Anserinae before 340

Tadorninae; Malacorhynchus, in its own tribe, was placed incertae sedis in Anatinae; and Oxyura, Nomonyx and Biziura, (Oxyurini) came at the end of Anatinae, after Aythini and Mergini.

In the analyses reported in Chapters 3, there was a tendency for all ‘oxyurines’ to associate together, but Thalassornis, Biziura and Malacorhynchus+Stictonetta, tended to disassociate from the main oxyurine clade in some analyses. In Chapter 5, where analyses were based on a larger character set and more taxa, a single oxyurine clade, with moderate support, was obtained, as shown in Figure 9.1. Thalassornis was consistently sister to Biziura, and this pair sister to Oxyura and Nomonyx, together forming the ‘traditional oxyurines’ with strong bootstrap and Bayesian support, especially in the analyses of Chapter 5. The more inclusive clade, of Stictonetta and all other oxyurines (fossil and recent), had moderate support, as did successively nested clades between Stictonetta and the traditional oxyurines. The incomplete nature of the six fossil taxa was almost certainly responsible for reducing support in these clades, and as these become better known, resolution within this whole clade can be expected to improve.

This newly identified clade demonstrates the importance of fossil taxa in understanding the relationships of modern taxa. Not only do the data presented herein overturn previous hypotheses of species relationships, but they may lead to an improved classification of Anseriformes, as the use of fossils was predicted to by Olson (1985: 217). A working hypothesis for such a classification is presented below (Table 3). The highly autapomorphic nature of the monotypic taxa Malacorhynchus and Stictonetta should be expected given they are the last survivors of ancient lineages. This hypothesis of a single clade of oxyurines simplifies anatid evolution, as it requires only two independent acquisitions of diving behaviour, one in oxyurines more derived than Malacorhynchus, and one on the mergine/aythyine lineage (Chapter 5).

Another significant outcome from this result is the discovery that most, if not all, known Oligocene–Middle Miocene ‘ducks’ are members of this oxyurine radiation. In the Late Miocene, oxyurines gave way to a new radiation of anatines out of which was spawned the now dominant Anas group. Australasia remains as the stronghold for oxyurines, possibly because it was effectively separated from other continents from the Late Oligocene until the Pliocene, so limiting the exposure of waterfowl therein to the expansion that replaced oxyurines elsewhere. This is supported by the observation that while there is in Australia essentially no Late Miocene and only a very limited Pliocene anseriform fauna, the only 341

documented extinction of a taxon in the Pliocene was the oxyurine Tirarinetta kanunka (Chapter 6). This extinction followed the entry into Australia, presumably from a Northern Hemisphere source, of Anas (at least four species), Aythya (at least one species) and Cygnus (at least one species). The discovery of fossil avifaunas in Australasia containing anseriforms that are of Late Miocene age and expanding the Pliocene record will be instrumental in revealing details of this faunal turnover.

9.6.6 A grade of tadornines

In his Tadorninae, Livezey (1997a) included tribe Merganettini for Hymenolaimus, Tachyeres and Merganetta; tribe Plectropterini for Plectropterus and ; Euryanatini for Euryanas (=Chenonetta finschi herein); and Tadornini for the sheldgeese (Cyanochen, Alopochen, , Chloephaga) and shelducks (Pachyanas, Tadorna). Livezey (1986) found little resolution of tadornines, but the scheme above derives from a study with 114 characters (39 skeletal, 4 and syrinx, 71 of and soft parts) and 19 taxa (Livezey 1996b). In this study, a strict consensus of 40 shortest trees found Hymenolaimus and Plectropterus unresolved with respect to a clade of core tadornines (Cyanochen, Alopochen, Neochen, Chloephaga and Tadorna) and most other taxa in the comparisons. Only in Majority Rule trees, or when complex weighting schemes were applied to the data, did these tadornine taxa form a clade (Livezey 1996b), and so they were united under Tadorninae (Livezey 1997a).

In the present analyses, Recent taxa usually included in Tadorninae (Chloephaga, Tadorna, Alopochen, Chenonetta and Hymenolaimus) formed a paraphyletic grade of taxa on the trees (e.g. Figure 9.1), but with little support for any particular sequence of taxa. In Chapter 3, in a weakly supported arrangement, Chloephaga and Alopochen were sister taxa and, in turn, were sister to a clade of Tadorna species. In Chapter 5, the more comprehensive datasets used resulted in the South American Chloephaga consistently, and with moderate support, taking a position basal to the oxyurine lineage (Figure 9.1), indicating that sheldgeese are only distantly related to shelducks. This differs from the finding by Livezey (1996b) that Tadorna is the sister to a larger group of tadornines including Alopochen and Chloephaga. In Chapter 5, Alopochen always was found to associate with Tadorna, but with little support. An enlarged character set chosen to 342

investigate shelduck-sheldgeese relationships is needed to resolve this issue, but as the affinity of Miotadorna was the primary interest, this aspect was not pursued.

Hymenolaimus was placed with moderate support highest in a paraphyletic grade of ‘tadornines’ (sensu Livezey 1996b, 1997a), as the sister to Anatinae in the analyses herein (Figure 9.1). In contrast, Livezey (1996b, 1997a) identified Hymenolaimus as a relatively basal tadornine sister to Tachyeres+Merganetta, which clade was, in turn, sister to the remaining tadornines.

Similarly, the relationship for Chenonetta found herein also departs markedly from that in Livezey (1997a). Firstly, Chenonetta jubata and Chenonetta (=Euryanas) finschi were found consistently as sister taxa, though with weak support. Livezey had these taxa widely separated, but this likely resulted from his methodology. Livezey’s (1986) first study placed Chenonetta jubata effectively unresolved in a group of anatines only weakly separated from the tadornines and in a later analysis, it was included within the framework of an investigation of anatine relationships (Livezey 1991). The relationships of ‘Euryanas’ were investigated in only one analysis that did not include C. jubata (Livezey 1989a), wherein it was found to have a sister relationship with unspecified ‘Tadorninae’ + ‘Anatinae’, but was more derived than Plectropterus, Stictonetta, Thalassornis, Dendrocygna and Anserinae. Thus, Livezey did not include both Chenonetta jubata and C. finschi in any analysis, despite a long history of these taxa being identified as closely related (e.g., Lydekker 1891; Oliver 1955; Howard 1964). The results found here support the analysis and results of Worthy & Olson (2002) that identified these as sister taxa. Moreover, Chenonetta was found here to form a clade with moderate support, wherein it was sister to Hymenolaimus and Anatinae, and thus more deeply nested than shelducks and sheldgeese. Thus Chenonetta, includes C. jubata and C. finschi, and should be arranged in a linear sequence after Tadorna and before Hymenolaimus. The data herein strongly suggest that the inclusion of Chenonetta and Hymenolaimus in Tadorninae would render that subfamily paraphyletic. Their phylogenetic relationships to other anatids would more correctly be expressed by their inclusion as basal taxa within Anatinae.

9.6.7 Towards a revised phylogeny of Anseriformes

The data presented in the preceding chapters and as discussed above invites a revision of the phylogenetic classification of Anseriformes. I have compiled a generic level 343

classification, derived from those advocated by Livezey (1997a) and Callaghan & Harshman (2005), which includes fossil taxa and adopts relationships shown by the results herein (Table 3). This table serves several purposes. Firstly, it identifies for the first time in a single place all valid fossil anseriform genera, identifying where they are poorly known (incertae sedis) and so inviting study of their relationships. In particular, the Oligo-Miocene European putative anserines, Cygnopterus, Cygnavus, and the Middle Miocene Sansan anseriforms, stand out as deserved of further study. Secondly, it identifies the phylogenetic sequence suggested by the results herein and so can serve as a framework for further analyses.

344

Table 3. A phylogenetic classification of anseriform genera including fossils, based on Livezey (1997a) and Kear (2005), and modified according to the results herein reported. † signifies an extinct taxon.

Order ANSERIFORMES (Wagler, 1831) Family incertae sedis Genus †Palaeopapia Harrison & Walker, 1979 Genus †Petropluvialis Harrison & Walker, 1976 Genus †Paracygnopterus Harrison & Walker, 1979 Genus †Paranyroca Miller & Compton, 1939 Genus †Eutelornis Ameghino, 1895 Genus †Eoneornis Ameghino, 1895

Suborder Anhimae Wetmore & Miller, 1926 Family Anhimidae Stejneger, 1885: screamers Genus Anhima Brisson, 1760 Genus †Chaunoides Alvarenga, 1999 Genus Illiger, 1811

Suborder Anseres Wagler, 1831 Family Anseranatidae (Sclater, 1880) Subfamily incertae sedis Genus †Anserpica Mourer-Chauviré, Berthet & Hugueney, 2004 Subfamily †Anatalavinae Olson, 1999 Genus †Anatalavis Olson & Parris, 1987 Subfamily Anseranatinae Sclater, 1880 Genus †Eoanseranas n. gen. Genus Anseranas Lesson, 1828

Family †Presbyornithidae Wetmore, 1926 Genus †Presbyornis Wetmore, 1926 Genus †Telmabates Howard, 1955 Genus †Headonornis Harrison & Walker, 1976

Family Anatidae Leach, 1820 Subfamily incertae sedis Genus †Loxornis Ameghino, 1895 Genus †Cayaoa Tonni, 1980 Genus †Geochen Wetmore, 1943 A Genus †Sinanas Yeh, 1980

Subfamily †Romainvilliinae (Lambrecht, 1933) Genus †Romainvillia Lebedinsky, 1927

Subfamily Anserinae Vigors, 1825: Swans and Geese Tribe incertae sedis Genus †Cygnopterus Lambrecht, 1931 Genus †Cygnavus Lambrecht, 1931 Genus †Guguschia Aslanova & Burczak-Abramovicz, 1968 Genus †Aminornis Ameghino, 1899 Tribe Cereopsini Vigors, 1825 Genus Cereopsis Latham, 1802 Genus †Cnemiornis Owen, 1866 Genus Coscoroba Reichenbach, 1853 345

Tribe Cygnini Vigors, 1825 Genus †Afrocygnus Louchart et al. 2005 Genus Cygnus Bechstein, 1803 Tribe Anserini Vigors, 1825 Genus †Presbychen Wetmore, 1930 Genus Anser Brisson, 1760 Genus Branta Scopoli, 1769 Genus †Eremochen Brodkorb, 1961

Subfamily Dendrocygninae Reichenbach, 1849: Whistling-Ducks Genus Dendrocygna Swainson, 1837

Subfamily Oxyurinae Phillips, 1926: Stiff-tailed Ducks Genus Stictonetta Reichenbach, 1853 Genus †Dendrochen Miller, 1944 Genus †Mionetta Livezey & Martin, 1988 Genus Malacorhynchus Swainson, 1831 Genus †Pinpanetta n. gen. Genus †Tirarinetta n. gen. Genus †Manuherikia Worthy, Tennyson, Jones, McNamara & Douglas, 2007 Genus †Dunstanetta Worthy, Tennyson, Jones, McNamara & Douglas, 2007 Genus Oxyura Bonaparte, 1828 Genus Nomonyx Ridgeway, 1880 Genus Heteronetta Salvadori, 1865 Genus Biziura Stephens, 1824 Genus Thalassornis Eyton, 1838

Subfamily Tadorninae Reichenbach, 1849: Shelducks Tribe Plectropterini (Eyton, 1838)B Genus Plectropterus Stephens, 1824 Genus Sarkidiornis Eyton, 1838 Tribe Tadornini (Reichenbach, 1849-1850) Genus †Australotadorna n.gen. Genus †Miotadorna Worthy, Tennyson, Jones, McNamara & Douglas, 2007 Genus †Anabernicula Ross, 1935 Genus †Telornis Ameghino, 1899C Genus †Anserobranta Kurokin & Ganea, 1972 D Genus Chloephaga Eyton, 1838 Genus Neochen Oberholser, 1918 Genus Cyanochen Bonaparte, 1856 Genus Alopochen Stejneger, 1885 Genus †Centriornis Andrews, 1897 Genus Tadorna Oken, 1817 Genus †Brantatadorna Howard, 1963

Subfamily Anatinae Leach, 1820: Ducks Tribe incertae sedis Genus Salvadorina Rothschild & Hartert, 1894 Genus †Matanas Worthy, Tennyson, Jones, McNamara & Douglas, 2007 Tribe Euryanatini (Livezey, 1989) Genus Chenonetta Brandt, 1836 Tribe Merganettini (Bonaparte, 1853) Genus Tachyeres Owen, 1875 Genus Merganetta Gould, 1842 Genus Hymenolaimus Gray, 1843 346

Tribe Anatini Leach Genus Cairina Fleming, 1822 Genus Asarcornis Salvdori, 1895 Genus Pteronetta Salvadori, 1895 Genus Nettapus Brandt, 1836 Genus Aix Boie, 1828 Genus Anas Linnaeus, 1758 Genus †Chelychelynechen Olson & James, 1991 Genus †Ptaiochen Olson & James, 1991 Genus †Thambetochen Olson & James, 1991 Genus †Pachyanas Oliver, 1955 Genus Amazonetta Boetticher, 1929 Genus Callonetta Delacour, 1936 Genus Lophonetta Riley, 1914 Genus Speculanas Boetticher, 1929 Genus Mareca Stephens, 1824 Tribe Aythyini Delacour and Mayr, 1945 Genus Marmaronetta Reichenbach, 1853 Genus Rhodonessa Reichenbach, 1853 Genus Netta Kaup, 1829 Genus Aythya Boie, 1822 Tribe Mergini Rafinesque, 1815: Sea Ducks Genus Polysticta Eyton, 1836 Genus Somateria Leach, 1819 Genus †Chendytes Miller, 1925 Genus †Megalodytes Howard, 1992E Genus Histrionicus Lesson, 1828 Genus Camptorhynchus Bonaparte, 1838 Genus Melanitta Boie, 1822 Genus Clangula Leach, 1819 Genus Bucephala Baird, 1858 Genus Mergellus Selby, 1840 Genus Lophodytes Reichenbach, 1853 Genus Mergus Linnaeus, 1758

Notes A. Subfamilial affinities indeterminate, either Anatinae or Anserinae (Olson & James 1991) B. Position tentative, may deserve subfamilial rank. C-E. provisional placements,

9.7 Improvements to the fossil record of anseriforms

Within the course of this project, considerable new data have been accumulated bearing on the fossil record of Anseriformes. Ten new fossil species have been added to the record, significantly expanding the diversity known from the Late Oligocene and Early Miocene, as highlighted in Table 4. These taxa establish Australasia as an important centre in early anseriform evolution and lead to the prediction that unrecognised diversity yet will be found in other southern continents, such as Africa and South America. 347

The analyses presented herein have resulted in the following insights into anseriform evolution: Dendrocheninae is no longer recognised and taxa from which it was previously comprised are now included in Oxyurinae; the diversity of Miocene oxyurines is more than doubled revealing that Early-Middle Miocene ‘ducks’ are dominated by this group; the first undoubted Oligocene-Miocene tadornines are described, with the oldest in the Late Oligocene; the first Miocene Southern Hemisphere anserines are identified; and the oldest acceptable record for Anatinae is now from the Early Miocene. Further, the data suggest there was a radiation of Southern Hemisphere anserines evolving in parallel with those in the north, and that tadornines arose in the south. Together these observations support the suggestion by Olson (1989) that Anseriformes evolved first in the Southern Hemisphere.

The phylogenetic analyses of the fossil taxa with recent genera provide some useful calibration points for future genetic analyses investigating divergence dates. The well dated Mionetta and Pinpanetta indicates that the world-wide radiation of oxyurines began no later than in the Late Oligocene. Similarly, the confirmation of Miotadorna as a tadornine, either sister to Tadorna or of similar evolutionary grade to shelducks and sheldgeese, indicates minimally an Early Miocene origin for them. The identification of Australotadorna, as a tadornine, even if conservatively identified as a sister to the shelduck+sheldgeese clade, gives an upper bound to the origin of this group in the Late Oligocene 26-24 Ma.

In Australasia, knowledge of anseriform evolution will clearly benefit from new collections of Oligo-Miocene age, if only so that existing taxa can be better known. But, particularly, what is needed are faunas of Middle-Late Miocene age, to address this critical period of faunal turnover, within which period modern anatine genera evolved and by the end of which modern faunas existed.

348

Table 4. The temporal distribution of fossil taxa in Anatidae that are older than the Pliocene. These include new taxa described herein (shaded), with revised subfamily attributions. If there is doubt concerning the subfamilial attribution, the genus is preceded by a ‘?’; if the generic attribution of a species is in doubt, a. ‘?’ is placed after the genus.

Anserinae Dendrocygninae Oxyurinae Tadorninae Anatinae Merginae and Aythyinae Oligocene Cygnopterus affinis ?Telornis impressus ?Cygnopterus lambrechti Guguschia nailiae Cygnavus formosus Aminornis excavatus Late Oligocene- Cygnopterus alphonsi Mionetta blanchardi Australotadorna Early Miocene alecwilsoni Cygnavus senckenbergi Mionetta consobrina Mionetta natator Dendrochen robusta Dendrochen integra Pinpanetta tedfordi Pinpanetta vickersrichae Pinpanetta fromensis Early Miocene NZ anserine Manuherikia lacustrina Miotadorna Matanas enrighti sanctibathansi Manuherikia minuta Manuherikia douglasi Dunstanetta johnstoneorum

Middle Miocene Anserobranta? robusta Aythya? chauvirae cf Tadorna Nördlinger Megalodytes morejohni Mergus? miscellus Ries & Calvert Fm Cygnus atavus Megalodytes sp. Japan Cygnus sp. 349

Presbychen abavus Anser sp.

Late Miocene Cygnus csakvarensis Dendrocygna sp. Oxyura? doksana Anserobranta Dendronessa sp. Mergus sp. tarabukuni Cygnus mariae Oxyura cf O. dominica Anas sp. 1& 2 Florida Bucephala ossivallis Afrocygnus chauvirae Anas sp. 1, 2, 3 Arizona Aythya shihubas Anser thraceiensis Aythya sp. Florida Anser arenosus Anser arizonae Anser? eldaricus Anser? udabnensis Branta woolfendeni Branta? thessaliensis Branta sp. Anabernicula sp.

350

351

CHAPTER 10 References

Agnolin, F. L. 2004. Revisión sistemática de algunas aves deseadenses (Oligoceno Medio) descriptas por Ameghino en 1899. Revista del Museo Argentino de Ciencias Naturales, n.s. 6(2): 239-244. Alvarenga, H. M. F. 1999. A fossil screamer (Anseriformes: Anhimidae) from the Middle Tertiary of South Eastern Brazil. In S. L. Olson (ed.), Avian paleontology at the close of the 20th century. Proceedings of the 4th International Meeting of the Society of Avian Paleontology and Evolution Washington, D. C., 4-7 June 1996. Smithsonian Contributions to 89: 223-230, 10 figures. Alvarez, R. 1977. A Pleistocene avifauna from Jalisco, Mexico. Contributions from the Museum of Paleontology, University of Michigan 24(19): 205-20. Alvarez, R. & Olson, S. L. 1978. A new merganser from the Miocene of Virginia (Aves: Anatidae). Proceedings of the Biological Society of Washington 91: 522-532. Aplin, K. P. 2006. Ten million years of rodent evolution in Australasia: Phylogenetic evidence and a speculative historical . Pp 707-744, in Merrick, J. R., Archer, M., Hickey, G. M. & Lee, M. S. Y. (eds.), Evolution and Biogeography of Australasian Vertebrates, Auscipub Pty Ltd, Oatlands, NSW, 942 pp. Apte, S.; Smith, P. J. & Wallis, G. P. 2007. Mitochondrial phylogeography of the New Zealand freshwater crayfishes, Paranephrops spp. Molecular Ecology 16: 1897-1908. Archer, M. & Hand, S. J. 2006. The Australian radiation. Pp 575-646, in Merrick, J. R., Archer, M., Hickey, G. M. & Lee, M. S. Y. (eds.), Evolution and Biogeography of Australasian Vertebrates, Auscipub Pty Ltd, Oatlands, NSW, 942 pp. Archer, M.; Arena, D. A.; Bassarova, M.; Beck, R. M. D.; Black, K.; Boles, W. E.; Brewer, P.; Cooke, B. N.; Crosby, K.; Gillespie, A.; Godthelp, H.; Hand, S. J.; Kear, B. P.; Louys, J.; Morrell, A.; Muirhead, J.; Roberts, K.; Scanlon, J. D.; Travouillon, K. J. & Wroe, S. 2006. Current status of species-level representation in faunas from selected fossil localities in the Riversleigh World Heritage Area, northwestern Queensland. Alcheringa Special Issue 1: 1- 17. Archer, M.; Arena, R.; Bassarova, M.; Black, K.; Bramwell, J.; Cooke, B.; Creaser, P.; Crosby, K.; Gillespie, A.; Godthelp, H.; Gott, M.; Hand, S. J.; Kear, B.; Krikmann, A.; Mackness, B.; Muirhead, J.; Musser, A.; Myers, T.; Pledge, N.; Wang, Y. & Wroe, S. 1999. The evolutionary history and diversity of Australian mammals. Australian Mammalogy 21: 1-45. Archer, M.; Hand, S. J.; Godthelp, H. & Creaser, P. 1997. Correlation of the Cainozoic sediments of the Riversleigh World Heritage Fossil Property, Queensland, Australia. In Aguilar, J. –P.; 352

Legendre, S. & Michaux, J. (eds), Actes du Congrès BiochroM’97, Mémoires et Travaux de l’Institut de Montpellier de l’Ecole Pratique des Hautes Etudes 21: 131-152. Aslanova, S. M. & Burczak-Abramowicz, N. I. 1968. A fossil swan from the Maykopian of Azerbaydzhan. Acta Zoologica Cracoviensia 13: 325-338, pls XI-XIV. Avise, J. C. & Walker, D. 1998. Pleistocene phylogeographic effects on avian populations and the speciation process. Proceedings of the Royal Society of London, Series B 265: 457-463. Baird, R. F. & Vickers-Rich, P. 1998. (Aves: from the Middle to Late Cainozoic of Australia. Alcheringa 22: 135-151. Baird, R. F.; Rich, P. V. & van Tets G.F. 1991. Localities yielding avian assemblages of Quaternary age in Australia. Pp 850-870, in Rich, P.V., Monaghan, J.M., Baird, R.F., Rich, T.H. (eds), Vertebrate Palaeontology of Australasia. Pioneer Design Studio and Monash University Publications Committee, Melbourne, 1437 pp. Baker, A. J.; Daugherty, C. H.; Colbourne, R. & McLennan, J. L. 1995. Flightless brown kiwis of New Zealand possess extremely subdivided population structure and cryptic species like small mammals. Proceedings of the National Academy of Science of the United States of America 92: 8254-8258. Baker, A. J.; Huynen, L. J.; Haddrath, O.; Millar, C. D. & Lambert, D. M. 2005. Reconstructing the tempo and mode of evolution in an extinct clade of birds with ancient DNA: The giant of New Zealand. Proceedings of the National Academy of Science of the United States of America 102: 8257- 8262. Barker, F. K.; Cibois, A.; Schikler, P.; Feinstein, J. & Cracraft, J. A. 2004. Phylogeny and diversification of the largest avian radiation. Proceedings of the National Academy of Sciences of the United States of America 101(30): 11040-11045. Becker, J. J. 1987. The fossil birds of the Late Miocene and Early Pliocene of Florida. 1. Geology, Correlation, and Systematic overview. Pp 159-171, in C. Mourer-Chauviré (ed.), L’Évolution des Oiseaux d’Après le Temoignage des Fossils. Documents du Laboratoire Geologique de Lyon 99, 248 pp. Beddard, F. E. 1898. The Structure and Classification of Birds. Longmans, Green, and Co., London. 548 pp. Bickart, K. J. 1990. The birds of the Late Miocene-Early Pliocene Big Sandy Formation, Mohave County, Arizona. Ornithological Monographs No. 44: 1-72. Bocheski, Z. 1997. List of European fossil bird species. Acta Zoologica Cracoviensia 40: 293-333. Boeuf, O. & Mourer-Chauviré, C. 1992. Les oiseaux des gisements d’âge Pliocène supérior de Chilhac, Haute-Loire, France. Comptes rendus de l'Académie des sciences, Paris 314(2): 1091-1096. Boev, Z. 1998. Fossil birds of Dorkovo – an Early Pliocene site in the Rhodope Mts, (Southern Bulgaria). Geologica Balcanica 28: 53-60. 353

Boev, Z. 2000. Cygnus verae sp. n. (Anseriformes: Anatidae) from the Early Pliocene of Sofia (Bulgaria). Acta Zoologica Cracoviensia 43: 185-192. Boev, Z. N. & Koufos, G. D. 2006. Aves [The late Miocene vertebrate locality of Perivolaki, Thessaly, Greece.] Palaeontographica Abteilung A, 276: 11-22. Boev, Z. N. 2002. Neogene avifauna of Bulgaria. Pp 29-40, in Zhonge Zhou & Fucheng Zhang (eds), Proceedings of the 5th symposium of the Society of Avian Paleontology and Evolution, Beijing, 1-4 June 2000, 311 pp. Science Press, Beijing. Boles, W. E. 1992. Revision of Dromaius gidju Patterson and Rich, 1987, with a reassessment of its generic position. In Campbell, K. E., Jr (ed.), Papers in avian paleontology honouring Pierce Brodkorb. Natural History Museum of Los Angeles County, Science Series 36: 195-208. Boles, W. E. 1993a. A new cockatoo (Psittaciformes: Cacatuidae) from the Tertiary of Riversleigh, northwestern Queensland, and an evaluation of rostral characters in the systematics of parrots. Ibis 135: 8-18. Boles, W. E. 1993b. A logrunner Orthonyx (Passeriformes: Orthonychidae) from the Miocene of Riversleigh, northwestern Queensland. Emu 93: 44-49. Boles, W. E. 1993c. Pengana robertbolesi, a peculiar from the Tertiary of Riversleigh, northwestern Queensland, Australia. Alcheringa 17: 19-25. Boles, W. E. 1995a. The world’s oldest songbird (Aves: Passeriformes). Nature 374: 21-22. Boles, W. E. 1995b. A preliminary analysis of the Passeriformes from Riversleigh, Northern Queensland, Australia, with the description of a new species of lyrebird. Courier Forschungsinstitut Senckenberg 181: 163-170. Boles, W. E. 1997a. Fossil songbirds (Passeriformes) from the Early Eocene of Australia. Emu 97: 43-50. Boles, W. E. 1997b. A kingfisher (Halcyonidae) from the Miocene of Riversleigh, Northwestern Queensland, with comments on the evolution of kingfishers in Australo-Papua. Memoirs of the Queensland Museum 41: 229-234. Boles, W. E. 1997c. Riversleigh birds as palaeoenvironmental indicators. Memoirs of the Queensland Museum 41: 241-246. Boles, W. E. 1997d. Hindlimb proportions and locomotion of Emuarius gigju (Patterson & Rich, 1987) (Aves: Casuariidae). Memoirs of the Queensland Museum 41: 235-240. Boles, W. E. 1998. A Budgerigar Melopsittacus undulatus from the Pliocene of Riversleigh, northwestern Queensland. Emu 98: 32-35. Boles, W. E. 1999. A new songbird (Aves: Passeriformes: Oriolidae) from the Miocene of Riversleigh, northwestern Queensland. Alcheringa 23: 51-56. Boles, W. E. 2001. A swiftlet (Apodidae: Collocaliini) from the Oligo-Miocene of Riversleigh, northwestern Queensland. Memoir of the Association of Australasian Palaeontologists 25: 45-52. 354

Boles, W. E. 2005a. A review of the Australian fossil storks of the genus Ciconia (Aves: Ciconiidae), with a description of a new species. Records of the Australian Museum 57: 165- 178. Boles, W. E. 2005b. A new flightless gallinule (Aves: Rallidae: Gallinula) from the Oligo-Miocene of Riversleigh, northwestern Queensland, Australia. Records of the Australian Museum 57: 179–190. Boles, W. E. 2005c. Fossil honeyeaters (Meliphagidae) from the Late Tertiary of Riversleigh, north- western Queensland. Emu 105: 21-26. Boles, W. E. 2006. The avian fossil record of Australia: an overview. Pp 387-411, in Merrick, J. R., Archer, M., Hickey, G. M. & Lee, M. S. Y. (eds.), Evolution and Biogeography of Australasian Vertebrates, Auscipub Pty Ltd, Oatlands, NSW, 942 pp. Boles, W. E. & Ivison, T. J. 1999. A new genus of the dwarf (Galliformes: Megapodiidae) from the late Oligocene of Central Australia. Proceedings of the 4th International meeting of the Society of Avian Paleontology and Evolution. Washington, USA, June 1996. Smithsonian Contributions to Paleobiology 89: 199-206. Boles, W. E. & Mackness, B. 1994. Birds from the Bluff Downs Local Fauna, Allingham Formation, Queensland. Records of the South Australian Museum 27: 139-149. Brodkorb, P. 1958a. Fossil birds from Idaho. Wilson Bulletin 70: 237-242. Brodkorb, P. 1958b. Birds from the Middle Pliocene of McKay, Oregon. Condor 60: 252-255. Brodkorb, P. 1962. The systematic position of two Oligocene birds from Belgium. Auk 79: 706-707. Brodkorb, P. 1964. Catalogue of fossil birds. Part 2 (Anseriformes through Galliformes). Bulletin of the Florida State Museum, Biological Sciences 8: 195-335. Brodkorb, P. 1978. Catalogue of Fossil Birds, Part 5. Bulletin of the Florida State Museum, Biological Sciences 23: 139-228. Brown, B.; Gaina, C. & Müller, R. D. 2006. Circum-Antarctic palaeobathymetry: Illustrated examples from the Cenozoic to recent times. Palaeogeography, Palaeoclimatology, Palaeoecology 231: 158-168. Brown, J. W.; Rest, J. S.; Garcia-Moreno, J.; Sorenson, M. D. & Mindell D. P. 2008. Strong mitochondrial DNA support for a Cretaceous origin of modern avian lineages. BMC Biology 2008, 6:6 doi:10.1186/1741-7007-6-6. Brush, A. H. 1976. Waterfowl feather proteins: analysis of use in taxonomic studies. Journal of Zoology, London 179: 467-498. Burczak-Abramovich, N. I. & Gadzyev, D. V. 1978. Anser eldaricus sp. nova from the Upper Sarmatian Hipparion fauna of Eldar. Acta Zoologica Cracoviensia 23: 67-78, pl. XV. Callaghan, D. & Harshman, J. 2005. Taxonomy and systematics, Pp 14-26, in Kear, J. (ed.), Ducks, Geese and Swans. Oxford University Press, Oxford, UK, 2 vols, 908 pp. 355

Campbell, H. & Hutching, G. 2007. In Search of Ancient New Zealand. and GNS Sciences, Wellington, New Zealand. 240 pp. Carboneras, C. 1992. Family Anatidae. Pp 536–628, in del Hoyo, J., Elliot, A. & Sargatal, J. (eds), Handbook of the Birds of the World. Vol. 1. Ostrich to Ducks. Lynx Edicions, Barcelona, 696 pp. Chandler, R. M. 1990. Fossil birds of the San Diego Formation, Late Pliocene, Blancan, San Diego County, California. Ornithological Monographs No. 44: 73-161. Cheneval, J. 1983. Les Anatidae (Aves, Anseriformes) du gisement Aquitanien de Saint-Gérand-le- Puy (Allier, France). Pp 85-98, in E. Buffetaut, J. M. Mazin & E. Salmon (eds): Actes du Symposium Paléontologique Georges Cuvier, Montbéliard, France, 1982. Ville de Montbéliard edit, Montbéliard, France. 548 pp. Cheneval, J. 1984. Les oiseaux aquatiques ( à Anseriformes) du gisement aquitanien de Saint-Gérand-le-Puy (Allier, France). Révision Systematique. Palaeovertebrata 14: 33-115, 6 figs, 13 tabl., 9 pl. Cheneval, J. 1987. Les Anatidae (Aves, Anseriformes) du Miocene de France. Révision Systématique et évolution. Pp 137-157, in C. Mourer-Chauviré (ed.), L’Évolution des Oiseaux d’Après le Temoignage des Fossils. Documents du Laboratoire Geologique de Lyon 99, 248 pp. Christidis, L. & Boles, W. E. 1994. The Taxonomy and Species of Birds of Australia and its Territories. Royal Australasian Ornithologist’s Union, Monograph 2. 122 pp. Christidis, L. & Boles, W. E. 2008. Systematics and Taxonomy of Australian Birds. Csiro Publishing, Collingwood, 277 pp. Clarke, J. A.; Tambussi, C. P.; Noriega, J. I.; Erickson, G. M. & Ketchman, R. A. 2005. Definitive fossil evidence for the extant avian radiation in the Cretaceous. Nature 433: 305-308. Clements, J. 1981. Birds of the World: a Checklist. Croom Helm, London, 562 pp. Condon, H. T. 1975. Checklist of the Birds of Australia, Part 1. Non-passerines. Royal Australasian Ornithologists Union, Melbourne, 311 pp. Cooke, B. 2006. Kangaroos. Pp 647-672, in Merrick, J. R., Archer, M., Hickey, G. M. & Lee, M. S. Y. (eds.), Evolution and Biogeography of Australasian Vertebrates, Auscipub Pty Ltd, Oatlands, NSW, 942 pp. Cooper, A. & Penny, D. 1997. Mass Survival of Birds Across the Cretaceous-Tertiary Boundary - Molecular Evidence. Science 275(5303): 1109-1113. Cooper, A. J. & Cooper, R. A. 1995. The Oligocene bottleneck and New Zealand biota: genetic record of a past environmental crisis. Proceedings of the Royal Society of London, Series B 261: 293-302. 356

Cooper, A.; Lalueza-Fox, C.; Anderson, S.; Rambaut, A.; Austin, J. & Ward, R. 2001. Complete mitochondrial sequences of two extinct moas clarify ratite evolution. Nature 409: 704-7. Cooper, R. A. & Millener, P. R. 1993. The New Zealand biota: historical background and new research. Trends in Ecology and Evolution 8: 429-433. Cracraft, J. 2001. Avian evolution, Gondwana biogeography and the Cretaceous-Tertiary mass extinction event. Proceedings of the Royal Society of London, Series B 268: 459-469. Cracraft, J.; Barker, F. K.; Braun, M.; Harshman, J.; Dyke, G. J.; Feinstein, J.; Stanley, S.; Cibois, A.; Schikler, P.; Beresford, P.; Garcia-Moreno, J.; Sorenson, M. D.; Yuri, T. & Mindell, D. P. 2004. Phylogenetic relationships among modern birds (Neornithes). Towards an avian tree of life. Pp 468-489, in: Cracraft, J. & Donoghue, M. (eds), Assembling the Tree of Life. Oxford University Press, London, 576 pp. Crowe, T. M.; Bowie, R. C. K.; Bloomer, P.; Mandiwana, T. G.; Hedderson, T. A. J.; Randi, E.; Pereira, S. L. & Wakeling, J. 2006. , biogeography and classification of, and character evolution in, gamebirds (Aves: Galliformes): effects of character exclusion, data partitioning and missing data. Cladistics 22: 495-532. De Vis, C. W. 1888. A glimpse of the post-Tertiary avifauna of Queensland. Proceedings of the Linnean Society of New South Wales, Series 2, 3: 1277-1292. De Vis, C. W. 1889. Additions to the list of fossil birds. Proceedings of the Royal Society of Queensland 6: 55-58. De Vis, C. W. 1905. A contribution to the knowledge of the extinct avifauna of Australia. Annals of the Queensland Museum 6: 3-25. Delacour, J. 1964. The Waterfowl of the World. Vol. 4. County Life Limited, London, 364 pp., 15 pls. Delacour, J. & Mayr, E. 1945. The family Anatidae. Wilson Bulletin 57: 3-55. Dickinson, E. C. (ed.) 2003. The Howard and Moore Complete Checklist of the Birds of the World (3rd ed). Christophe Helm, London. 1039 pp. Donne-Goussé, C.; Laudet, V. & Hänni, C. 2002. A molecular phylogeny of Anseriformes based on mitochondrial DNA analysis. Molecular Phylogenetics and Evolution 23: 339-356. Douglas, B. J. 1986. Lignite resources of Central Otago. Manuherikia Group of Central Otago, New Zealand: Stratigraphy, depositional systems, lignite resource assessment and exploration models. New Zealand Energy Research and Development Committee Publication, 104 pp, 2 vols. Douglas, B. J.; Lindqvist, J. K.; Fordyce, R. E. & Campbell, J. D. 1981. Early Miocene terrestrial vertebrates from Central Otago. Geological Society of New Zealand Newsletter 53: 17. 357

Driskell, A.; Christidis, L.; Gill, B. J.; Boles, W. E.; Barker, F. K. & Longmore, N. W. 2007. A new endemic family of New Zealand passerine birds: adding heat to a biodiversity hotspot. Australian Journal of Zoology 55: 73-78. Dyke, G. J. 2001. The fossil waterfowl (Aves: Anseriformes) from the Eocene of England. American Museum Novitates 3354: 1-15. Emslie, S. D. 1992. Two new Late Blancan avifaunas from Florida and the extinction of the Wetland Birds in the Plio-Pleistocene. In Campbell, K. E. (ed.), Papers in avian paleontology honoring Pierce Brodkorb. Natural History Museum of Los Angeles County, Science Series 36: 249-269. Ericson, P. G. P. 1997. Systematic relationships of the palaeogene family Presbyornithidae (Aves: Anseriformes). Zoological Journal of the Linnean Society 121: 429-483. Ericson, P. G. P. 2000. Systematic revision, skeletal , and paleoecology of the New World early tertiary Presbyornithidae (Aves: Anseriformes). PaleoBios 20: 1-23. Ericson, P. G. P.; Anderson, C. L.; Britton, T.; Elzanowski, A.; Johansson, U. S.; Kallersjo, M.; Ohlson, J. I.; Parsons, T. J.; Zuccon, D. & Mayr, G. 2006. Diversification of : integration of molecular sequence data and fossils. Biology Letters 2: 543-547. Ericson, P. G. P.; Christidis, L.; Cooper, A.; Irestedt, M.; Jackson, J.; Johansson, U. S. & Norman, J. A. 2002. A Gondwanan origin of passerine birds supported by DNA sequences of the endemic New Zealand wrens. Proceedings of the Royal Society of London, Series B 269: 235-241. Eyton, T. C. 1838. A Monograph on the Anatidae-or Duck Tribe. Longman, Orme, Brown, Green and Longman, London, x + 178 pp. Eyton, T. C. 1867. Osteologia Avium: or a Sketch of the Osteology of Birds. Wellington, London. i- vi, x, 1-229 pp; 113 pls. Eyton, T. C. 1869. Osteologia Avium: or a Sketch of the Osteology of Birds. Supplement, &c. Wellington, London, 18 pls. Eyton, T. C. 1873-75. Osteologia Avium: or a Sketch of the Osteology of Birds. Supplement II. Wellington, London, 3 pt: 51 pls, with descriptive letterpress. Fain, M. G. & Houde, P. 2004. Parallel radiations in the primary clades of birds. Evolution 58: 2558-2573. Fain, M. G.; Krajewski, C.; & Houde, P. 2007. Phylogeny of “core Gruiformes” and the resolution of the Limpkin-Sungrebe problem. Molecular Phylogenetics and Evolution 43: 515-529. Falla, R. A. 1953. The Australian element in the avifauna of New Zealand. Emu 53: 36-46. Fleming, C. A. 1962. History of the New Zealand landbird fauna. Notornis 9: 270-274. Fleming, C. A. 1979. The Geological History of New Zealand and its Life. Auckland University Press and Oxford University Press, Auckland, 141 p. 358

Fordyce, R. E. 1991. A new look at the fossil vertebrate record of New Zealand. Pp 1191-1316, in P. V. Rich, J. M. Monaghan, R. F. Baird, T. H. Rich (eds), Vertebrate Palaeontology of Australasia. Pioneer Design Studios in cooperation with the Monash University Publications Committee, Melbourne, 1437 pp. Fordyce, R. E. 2003. Fossils and the history of life. Pp 35-64, in J. Darby, R. E. Fordyce, A. Mark, K. Probert & C. Townsend (eds), The Natural History of Southern New Zealand. Otago University Press, Dunedin, NZ, 387 pp. Fordyce, R. E. & Jones, C. M. 1990. Penguin history and new fossil material from New Zealand. Pp 419-446, in J. Darby (ed.), Penguin biology. Academic Press, San Diego.

Frith, H. J. 1964. Taxonomic relationships of Stictonetta naevosa (Gould). Nature 202: 1352-3.

Frith, H. J. 1977. Waterfowl in Australia. A.H. & A.W. Reed, Sydney. 328 pp. Fürbringer, M. 1888. Untersuchungen zur Morphologie und systematik der Vogel II. Allgemeiner Theil Van Holkema, Amsterdam, 2 vol., 1751 pp, 30 pl. Gadow, H. 1892. On the classification of birds. Proceedings of the Zoological Society of London, 1892: 229-256. Gaina, C.; Müller, R. D.; Royer, J. –Y.; Stock, J.; Hardebeck, J. & Symonds, P. 1998b. The tectonic evolution of the Tasman Sea: A tectonic puzzle with thirteen pieces. Journal of Geophysical Research 103 (B6): 12,413-12,433. Gaina, C.; Roest, W. R.; Müller, R. D. & Symonds, P. 1998a. The opening of the Tasman Sea: A gravity anomaly Animation. Earth Interactions 2(4): 1-23. Gibbs, G. 2006. Ghosts of Gondwana. The history of life in New Zealand. Craig Potton Publishing, Nelson, New Zealand, 232 pp. Godthelp, H.; Archer, M.; Cifelli, R.; Hand, S. J. & Gilkeson, C. E. 1992. Earliest known Australian Tertiary mammal fauna. Nature 356: 514-516. Graf, D. L. & Foighil, D. Ó. 2000. Molecular phylogenetic analysis of 28S rDNA supports a Gondwanan origin for Australasian Hyriidae (: Bivalvia: Unionoida).Vie et Milieu 50(4): 245-254. Griffiths, C. S.; Barrowclough, G. F.; Groth, J. G. & Mertz, L. A. 2007. Phylogeny, diversity, and classification of the Accipitridae based on DNA sequences of the RAG-1 exon. Journal of Avian Biology 38: 587-602. Hand, S. J. 2006. Bat beginnings and biogeography: the Australasian record. Pp 673-705, in Merrick, J. R., Archer, M., Hickey, G. M. & Lee, M. S. Y. (eds.), Evolution and Biogeography of Australasian Vertebrates, Auscipub Pty Ltd, Oatlands, NSW, 942 pp. Hand, S.; Beck, R.; Worthy, T.; Archer, M. & Sigé, B. 2007. Australian and New Zealand bats: The origin, evolution, and extinction of bat lineages in Australasia. Journal of Vertebrate Paleontology 27(3) Abstracts: 86A. 359

Harrison, C. J. O. & Walker, C. A. 1976. Birds of the British Upper Eocene. Zoological Journal of the Linnean Society 59: 323-351. Harrison, C. J. O. & Walker, C. A. 1979a. New names for two Upper Eocene bird genera. Tertiary Research 2: 110. Harrison, C. J. O. & Walker, C. A. 1979b. Birds of the British Lower Oligocene. Tertiary Research Special Papers 5: 29-39. Harrison, G. L. (Abby); McLenachan, P. A.; Phillips, M. J.; Slack, K. E.; Cooper, A. & Penny, D. 2004. Four new avian mitochondrial help get to basic evolutionary questions in the Late Cretaceous. Molecular Biology and Evolution 21(6): 974-983. Heads, M. 2006. Panbiogeography of Nothofagus (Nothofagaceae): Analysis of the main species massings. Journal of Biogeography 33: 1066-1075. Heizmann, E. P. J. & Hesse, A. 1995. Die mittelmiozänen Vogel- und Säugetierfaunen des Nördlinger Ries (MN6) und des Steinheimer Beckens (MN7) – ein Vergleich. Courier Forschungsinstitut Senckenberg 181: 171-185. Helbig, A. J.; Kocum, A.; Ingrid Seibold, I. & Braun, M. J. 2005. A multi-gene phylogeny of aquiline eagles (Aves: ) reveals extensive at the genus level. Molecular Phylogenetics and Evolution 35: 147-164. Hennig, W. 1966. Phylogenetic Systematics. University of Illinois Press, Urbana, Illinois. 263 pp. Hiroshige, M.; Hajime, N.; Yuji, T. & Yoshikazu, H. 2001. Preliminary note on the Miocene flightless swan from the Haraichi Formation, Tomioka Group of Annaka, Gunma, Japan. Bulletin of Gunma Museum of Natural History 5: 1-8. Hiroshige, M.; Yoshikazu, H. & Yuji, T. 2004. Osteological note on the completely prepared fossil “Annaka Short-winged Swan” from the Miocene Tomioka Group, Japan. Bulletin of Gunma Museum of Natural History 8: 35-56. Holdaway, R. N. 1994. An exploratory phylogenetic analysis of the genera of the Accipitridae, with notes on the biogeography of the family. Pp 601-649, in Meyburg, B. –U. & Chancellor, R. D. (eds), Raptor Conservation Today. World Working Group on Birds of Prey and , Berlin. Hope, S. 2002. The Mesozoic radiation of Neornithines. Pp 339-388, in Chiappe, L. M. & Witmer, L. M. (eds), Mesozoic Birds: Above the Heads of Dinosaurs. University of California Press, Berkeley, California, 536 pp. Howard, H. 1964. Fossil Anseriformes. Pp 233-326, in Delacour, J. (ed.), The Waterfowl of the World. Country Life Ltd, London, United Kingdom, 364 pp. Howard, H. 1992. New records of Middle Miocene anseriform birds from Kern County, California. In Campbell, K. E. (ed), Papers in avian paleontology honoring Pierce Brodkorb. Natural History Museum of Los Angeles County, Science Series, 36: 231-237. 360

Huxley, T. H. 1867. On the classification of birds; and on the taxonomic value of the modifications of certain of the cranial bones observable in that . Proceedings of the Zoological Society of London 1867: 415-472. Johnsgard, P. A. 1961. The taxonomy of Anatidae – a behavioural analysis. Ibis 103: 71-85. Johnsgard, P. A. 1968. Waterfowl: Their Biology and Natural History. University of Nebraska Press, Lincoln, xviii + 138 pp. Johnsgard, P. A. 1978. Ducks, Geese and Swans of the World. University of Nebraska Press, Lincoln and London, xxii + 404 pp.

Johnson, K. P. & Sorenson, M. D. 1999. Phylogeny and biogeography of dabbling ducks (Genus: Anas): a comparison of molecular and morphological evidence. Auk 116(3): 792-805. Johnson, N. K. & Cicero, C. 2003. New mitochondrial DNA data affirm the importance of Pleistocene speciation in North American birds. Evolution 58(5): 1122-1130. Kear, J. (ed.) 2005. Ducks, Geese and Swans. Oxford University Press, Oxford, UK. 2 vols, 908 pp. Kear, J. & Murton, R. K. 1976. The origins of Australian waterfowl as indicated by their photoresponses. Pp 83-97, in Frith, H. J. & Calaby, J. H. (eds), Proceedings of the 16th International Ornithological Congress. Australian Academy of Science, Canberra, 765 pp. Klicka, J. & Zink, R. M. 1997. The importance of Recent ice ages in speciation: a failed paradigm. Science 277: 1666-1669. Knapp, M.; Mudaliar, R.; Havell, D.; Wagstaff, S. J. & Lockhart, P. J. 2007. The drowning of New Zealand and the problem of Agathis. Systematic Biology 56(5): 862-870. Kurochkin, E. N. 1985. Birds of the Central Asia in Pliocene. The Joint Soviet-Mongolian Paleontological Expedition (Transactions) [In Russian] 26: 5-119. Kurochkin, E. N.; Dyke, G. J. & Karhu, A. A. 2002. A new Presbyornithid bird (Aves: Anseriformes) from the late Cretaceous of Southern Mongolia. American Museum Novitates 3386: 1-11. Kurochkin, Ye. N. 1968. New Oligocene birds from Kazakhstan. Paleontological Journal (a translation of Paleontologicheski Zhurnal) 2: 83-90. Lambrecht, K. 1933. Handbuch der Palaeornithologie. Berlin, Gebrüder, Borntraeger. 1024 pp. Landis, C. A.; Campbell, H. J.; Begg, J. G.; Mildenhall, D. C.; Paterson, A. M. & Trewick, S. A. 2008. The Waipounamu Erosion Surface: questioning the antiquity of the New Zealand land surface and terrestrial fauna and flora. Geological Magazine 145(2): 173-197. Lee, D. E.; Bannister, J. M. & Lindqvist, J. K. 2007. Late Oligocene-Early Miocene leaf macrofossils confirm a long history of Agathis in New Zealand. New Zealand Journal of Botany 45: 565-578. Lianhai, H. 1985. Upper Miocene birds from Lufeng, Yunnan. Acta Anthropologica Sinica IV(2): 118-126. 361

Livezey, B. C. 1986. A phylogenetic analysis of Recent anseriform genera using morphological characters. Auk 105: 681-698. Livezey, B. C. 1989a. Phylogenetic relationships of several subfossil Anseriformes of New Zealand. Occasional papers of the Museum of Natural History of the University of Kansas 128: 1-25. Livezey, B. C. 1989b. Phylogenetic relationships and incipient flightlessness in the extinct Auckland Islands merganser. Wilson Bulletin 101: 410-435. Livezey, B. C. 1991. A phylogenetic analysis and classification of Recent dabbling ducks (Tribe Anatini) based on comparative morphology. Auk 108: 471-507. Livezey, B. C. 1993. Morphology of flightlessness in Chendytes, fossil seaducks (Anatidae: Mergini) of coastal California. Journal of Vertebrate Paleontology 13(2): 185-199. Livezey, B. C. 1995a. Phylogeny and comparative ecology of the stiff-tailed ducks (Anatidae: Oxyurini). Wilson Bulletin 107: 214-234. Livezey, B. C. 1995b. A phylogenetic analysis of the whistling and white-backed ducks (Anatidae: Dendrocygninae) using morphological characters. Annals of the Carnegie Museum 64: 65-97. Livezey, B. C. 1996a. A phylogenetic analysis of geese and swans (Anseriformes: Anserinae), including selected fossil species. Systematic Biology 45: 415-450. Livezey, B. C. 1996b. A phylogenetic reassessment of the tadornine-anatine divergence (Aves: Anseriformes: Anatidae). Annals of the Carnegie Museum 65: 27-88. Livezey, B. C. 1996c. A phylogenetic analysis of modern pochards (Anatidae: Aythyini). Auk 113: 74-93. Livezey, B. C. 1997a. A phylogenetic classification of waterfowl (Aves: Anseriformes), including selected fossil species. Annals of the Carnegie Museum 67: 457-496. Livezey, B. C. 1997b. A phylogenetic analysis of basal Anseriformes, the fossil Presbyornis, and the interordinal relationships of waterfowl. Zoological Journal of the Linnean Society 121: 361-428. Livezey, B. C. 1997c. A phylogenetic analysis of modern shelducks and sheldgeese (Anatidae: Tadornini). Ibis 139: 51-66. Livezey, B. C. 1998. A phylogenetic analysis of the Gruiformes (Aves) based on morphological characters, with an emphasis on the rails (Rallidae). Philosophical Transactions of the Royal Society London B 353: 2077-2151. Livezey, B. C. & Martin, L. D. 1988. The systematic position of the Miocene anatid Anas [?] blanchardi Milne- Edwards. Journal of Vertebrate Paleontology 8(2): 196-211. Livezey, B. C. & Zusi, R. L. 2007. Higher-order phylogeny of modern birds (, Aves: Neornithes) based on comparative anatomy. II. Analysis and discussion. Zoological Journal of the Linnean Society 149: 1-95. 362

Louchart, A.; Vignaud, P.; Likius, A.; Mackaye, H. T. & Brunet, M. 2005. A new swan (Aves: Anatidae) in Africa, from the latest Miocene of Chad and Libya. Journal of Vertebrate Paleontology 25: 384-392. Lydekker, R. 1891. Catalogue of Fossil Birds in the (Natural History). British Museum (Natural History), London, 368 pp. Madsen, C. S.; McHugh, K. P. & de Kloet, S. W. R. 1988. A partial classification of the waterfowl (Anatidae) based on single-copy DNA. Auk 105: 452-459. Magee, J. W. & Miller, G. H. 1998. Lake Eyre palaeohydrology from 60 ka to the present: beach ridges and glacial maximum aridity. Palaeogeography, Palaeoclimatology, Palaeoecology 144: 307-329. Magee, J. W.; Bowler, J. M.; Miller, G. H. & Williams, D. L. G. 1995. Stratigraphy, sedimentology, chronology and palaeohydrology of Quaternary lacustrine deposits at Madigan Gulf, Lake Eyre, South Australia. Palaeogeography, Palaeoclimatology, Palaeoecology 113: 3-42. Magee, J. W.; Miller, G. H.; Spooner, N. A. & Questiaux, D. 2004. Continuous 150 k.y. monsoon record from Lake Eyre, Australia: Insolation-forcing implications and unexpected Holocene failure. Geology 32: 885-888. Marchant, S. & Higgins, P. J. (co-ordinators) 1990. Handbook of Australian, New Zealand & Antarctic Birds, vol. 1, Ratites to Ducks. Oxford University Press, Melbourne, 1400 pp. Maroulis, J. C.; Nanson, G. C.; Price, D. M. & Pietsch, T. 2007. Aeolian-fluvial interaction and climate change: source-bordering dune development over the past ~100 ka on Cooper Creek, central Australia. Quaternary Science Reviews 26: 386-404. Martin, L. D. & Mengel, R. M. 1980. A new goose from the Late Pliocene of Nebraska with notes on variability and proportions in some recent geese. Papers in Avian Paleontology honouring Hildegarde Howard. Contributions in Science, Natural History Museum of Los Angeles County 330: 75-85. Mayr, G. 2005. The Paleogene fossil record of birds in Europe. Biological Review 80: 515-542. Mayr, G. in press. Phylogenetic affinities and morphology of the late Eocene anseriform bird Romainvillia stehlini Lebedinsky, 1927. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. Mayr, E. & Cottrell, G. W. 1979. [Peter’s] Check- of the World, Vol. 1, Second Edition. Museum of Comparative Biology, Cambridge Massachusetts, xvii + 547 pp. Mayr, G. & Smith, R. 2001. Ducks, rails, and limicoline waders (Aves: Anseriformes, Gruiformes, Charadriiformes) from the lowermost Oligocene of Belgium. Geobios 34: 547-561. McCracken, K. G. & Sheldon, F. H. 1998. Molecular and osteological heron phylogenies: sources of incongruence. Auk 115: 127-141. 363

McCracken, K. G. & Sorenson, M. D. 2005. Is homoplasy or lineage sorting the source of incongruent mtDNA and nuclear gene trees in the stiff-tailed ducks (Nomonyx-Oxyura). Systematic Biology 54: 35-55. McCracken, K. G.; Harshman, J.; McClellan, D. A. & Afton, A. D. 1999. Data set incongruence and correlated character evolution: an example of functional convergence in the hind-limbs of stiff-tail ducks. Systematic Biology 48: 683- 714. Megirian, D.; Murray, P.; Schwartz, L. & Von Der Borch, C. 2004. Late Oligocene Kangaroo Well Local Fauna from the Ulta Limestone (new name), and climate of the Miocene oscillation across central Australia. Australian Journal of Earth Sciences 51: 701-741. Meyer, A. B. 1879-1897. Abbildungen von Vogel-Skeletten. Herausgegeben mit Unterstutzung der Generaldirection der Königl. Sammlungen für Kunst und Wissenschaft in Dresden, Berlin and Dresden. Vol. 1: xiv + 71 pp., 120 pls; vol. 2: xxi + 120 pp., 122 pls. Mildenhall, D. C. 1989. Summary of the age and palaeoecology of the Miocene Manuherikia Group, Central Otago, New Zealand. Journal of the Royal Society of New Zealand 19: 19-29. Mildenhall, D. C. & Pocknall, D. T. 1989. Miocene-Pleistocene spores and pollen from Central Otago, South Island, New Zealand. New Zealand Geological Survey Palaeontological Bulletin 59: 1-128. Millener, P. R. 1991. The Quaternary avifauna on New Zealand. Pp 1317-1344, in Vickers-Rich, P. V.; Monaghan, J. M.; Baird, R.F.; Rich, T. H. (eds), Vertebrate Palaeontology of Australasia. Pioneer Design Studios in cooperation with the Monash University Publications Committee, Melbourne, 1437 pp. Miller, A. H. 1948. The whistling swan in the Upper Pliocene of Idaho. Condor 50: 132. Miller, A. H. 1963. The fossil flamingos of Australia. Condor 65: 289-299. Miller, A. H. & Compton, L. V. 1939. Two fossil birds from the Lower Miocene of South Dakota. Condor 41: 153-156. Miller, L. 1930. A fossil goose from the Ricardo Pliocene. Condor 32: 208-209. Miller, L. 1944. Some Pliocene birds from Oregon and Idaho. Condor 46: 25-32. Miller, L. 1952. The avifauna of the Barstow Miocene of California. Condor 54: 296-301. Miller, L. 1961. Birds from the Miocene of Sharktooth Hill, California. Condor 63: 399-402. Milne-Edwards, A. 1863. Mémoire sur la distribution géologique des oiseaux fossils et description de quelques especies nouvelles. Annales des Sciences Naturelles 4(20): 132-176. [Abbreviated versions in Bulletin de la Sociétié Philomatique 1863: 61-64; Comptes Rendus des Séances Hebdomadaires de l’Académie des Sciences (Paris) 56: 1219-1222; Revue des Sociétés Savantes (1)4: 1-5.] Milne-Edwards, A. 1867-71. Recherches Anatomiques et Paléontologiques pour Server à l’Historie des Oiseaux Fossils de la France. Masson, Paris, 1: 472 pp, 96 pl; 2: 627 pp, pl. 97-200. Mlíkovský, J. 2002. Cenozoic Birds of the World, Part 1: Europe. Ninox Press, Praha, 406 pp. 364

Mlíkovský, J. & Švec, P. 1986. Review of the Tertiary waterfowl (Aves: Anseridae) of Asia. Vestnik Cestoslovenske Spolecnosti Zoologicke 50: 259-272. Molnar, R. E. & Pole, M. 1997. A Miocene crocodilian from New Zealand. Alcheringa 21: 65-70. Molnar, R. E. & Wiffen, J. 1994. Polar faunas from New Zealand. Cretaceous Research 15: 689-706. Mourer-Chauviré, C. 2004. Review of: Cenozoic Birds of the World, Part 1: Europe. Auk 121: 623- 627. Mourer-Chauviré, C.; Berthet, D. & Hugueney, M. 2004. The Late Oligocene birds of the Créchy Quarry (Allier, France), with a description of two new genera (Aves: Pelecaniformes: Phalacrocoracidae, and Anseriformes: Anseranatidae). Senkenbergiana lethaea 84: 303-315. Murray, P. & Megirian, D. 1992. Continuity and contrast in Middle and Late Miocene vertebrate communities from the Northern Territory. The Beagle, Records of the Northern Territory Museum of Arts and Sciences 9: 195- 218. Murray, P. F. & Vickers-Rich, P. 2004. Magnificent Mihirungs, the Colossal Flightless Birds of the Australian Dreamtime. Indiana University Press, Bloomington and Indianapolis, 410 pp. Nanson, G. C.; Callen, R. A. & Price, D. M. 1998. Hydroclimatic interpretation of Quaternary shorelines on South Australian playas. Palaeogeography, Palaeoclimatology, Palaeoecology 144: 281-305. Nanson, G. C.; Price, D. M. & Short, S. A. 1992. Wetting and drying of Australia over the past 300 ka. Geology 20: 791-794. Noriega, J. I. 1995. The avifauna from the “Mesopotamian” (Ituzaingó Formation, Upper Miocene) of Entre Rios Province, Argentina. Courier Forschungsinstitut Senckenberg 181: 141-148. Noriega, J. I. & Tambussi, C. P. 1995. A Late Cretaceous Presbyornithidae (Aves Anseriformes) from Vega Island, Antarctic Peninsula; biogeographic interpretations. Ameghiniana 32: 57- 61. Oliver, W. R. B. 1955. New Zealand Birds. A.H. & A.W. Reed, Wellington, 661 pp. Olson, S. L. 1977a. Notes on the subfossil Anatidae from New Zealand, including a new species of pink-eared duck Malacorhynchus. Emu 77: 132-135. Olson, S. L. 1977b. The identity of the fossil ducks described from Australia by C. W. De Vis. Emu 77: 127-131. Olson, S. L. 1985. The fossil record of birds. Pp 79-238, in Farner, D. S., King, J. R. & Parkes, K. C. (eds), Avian biology, vol. 8. Academic Press, New York, 256 pp. Olson, S. L. 1989. Aspects of global avifaunal dynamics during the Cenozoic. Pp 2023-2029, in Ouellet, H. (ed), Acta XIX Congressus Internationalis Ornithologici, volume 2, University of Ottawa Press, Ottawa. 365

Olson, S. L. 1994. A giant Presbyornis (Aves: Anseriformes) and other birds from the Paleocene of Maryland and Virginia. Proceedings of the Biological Society of Washington 107: 429-435. Olson, S. L. 1999. The anseriform relationships of Anatalavis Olson and Parris (Anseranatidae), with a new species from the Lower Eocene London Clay. Pp 231-243, 9 figures, in Olson, S. L. (ed.), Avian paleontology at the close of the 20th century. Proceedings of the 4th International Meeting of the Society of Avian Paleontology and Evolution Washington, D. C., 4-7 June 1996. Smithsonian Contributions to Paleobiology 89. Olson, S. L. & Feduccia, A. 1980. Presbyornis and the origin of the Anseriformes (Aves: Charadriomorphae). Smithsonian Contributions to Zoology 323: 1-24. Olson, S. L. & James, H. F. 1991. Descriptions of thirty-two new species of birds from the Hawaiian Islands: Part 1. Non-Passeriformes. Ornithological Monographs 45: 1-88. Olson, S. L. & Parris, D. C. 1987. The Cretaceous birds of New Jersey. Smithsonian Contributions to Palaeobiology 63, 22 pp. Olson, S. L. & Rasmussen, P. C. 2001. Miocene and Pliocene birds from the Lee Creek Mine, North Carolina. Smithsonian Contributions to Paleobiology 90: 233-365. Owen, R. 1866. On Dinornis (Part X): containing a description of part of the skeleton of a flightless bird indicative of a new genus and species (Cnemiornis calcitrans, Ow.). Transactions of the Zoological Society of London V(5): 395-404, Pls. LXIII-LXVII. Parris, D. C. & Hope, S. 2002. New interpretations of the birds from the Navesink and Hornerstone Formations, New Jersey, USA (Aves: Neornithes). Pp 113-124, in Zhonge Zhou & Fucheng Zhang (eds), Proceedings of the 5th symposium of the Society of Avian Paleontology and Evolution, Beijing, 1-4 June 2000. Science Press, Beijing. Pereira, S. L. & Baker A. J. 2006. A mitogenomics timescale for birds detects variable phylogenetic rates of molecular evolution and refutes the standard molecular clock. Molecular Biology and Evolution 23: 1731-1740. Peters, J. L. 1931. Check-list of Birds of the World, Vol. 1. Harvard University Press, Cambridge, 345 pp. Phillips, J. C. 1923. Natural History of the Ducks, Volume 1, Plectropterinae, Dendrocygninae, Anatinae. Longmans, Green and Company, London, xi + 264 pp. Phillips, J. C. 1926. Natural History of the Ducks, Volume 4, Fuligulinae (concluded), Oxyurinae, Merganettinae and Merginae. Longmans, Green and Company, London, xii + 489 pp. Pledge, N. S. 1984. A new Miocene vertebrate faunal assemblage from the Lake Eyre Basin. A preliminary Report. Australian Zoologist 21(4): 345-355. Pledge, N. S. & Tedford, R. H. 1990. Vertebrate fossils, in Tyler M. J., Twidale C. R., Davies M., Wells C.B. (eds). Natural History of the North East Deserts. Occasional Publications of the Royal Society of South Australia 6: 199-209. 366

Pole, M. 1989. Early Miocene floras from Central Otago, New Zealand. Journal of the Royal Society of New Zealand 19: 121-125. Pole, M. 1992a. Early Miocene flora of the Manuherikia Group, New Zealand. 1. Ferns. Journal of the Royal Society of New Zealand 22: 279-286. Pole, M. 1992b. Early Miocene flora of the Manuherikia Group, New Zealand. 2. Conifers. Journal of the Royal Society of New Zealand 22: 287-302. Pole, M. 1992c. Early Miocene flora of the Manuherikia Group, New Zealand. 3. Possible cycad. Journal of the Royal Society of New Zealand 22: 303-306. Pole, M. 1993a. Early Miocene flora of the Manuherikia Group, New Zealand. 4. Palm remains. Journal of the Royal Society of New Zealand 23: 283-288. Pole, M. 1993b. Early Miocene flora of the Manuherikia Group, New Zealand. 5. Smilacaceae, Polygonaceae, Elaeocarpaceae. Journal of the Royal Society of New Zealand 23: 289-302. Pole, M. 1993c. Early Miocene flora of the Manuherikia Group, New Zealand. 6. Lauraceae. Journal of the Royal Society of New Zealand 23: 303-312. Pole, M. 1993d. Early Miocene flora of the Manuherikia Group, New Zealand. 7. Myrtaceae. Journal of the Royal Society of New Zealand 23: 313-328. Pole, M. 1993e. Early Miocene flora of the Manuherikia Group, New Zealand. 8. Nothofagus. Journal of the Royal Society of New Zealand 23: 329-344. Pole, M. 1993f. Early Miocene flora of the Manuherikia Group, New Zealand. 9. Miscellaneous leaves and reproductive structures. Journal of the Royal Society of New Zealand 23: 345-391. Pole, M. 1993g. Early Miocene floras of the Manuherikia Group, New Zealand. 10. Paleoecology and stratigraphy. Journal of the Royal Society of New Zealand 23: 393-426. Pole, M. 1994. The New Zealand flora-Entirely long-distance dispersal. Journal of Biogeography 21: 625-635. Pole, M. 1997. Miocene conifers from the Manuherikia Group, New Zealand. Journal of the Royal Society of New Zealand 27: 355-370. Pole, M. & Douglas, B. 1998. A quantitative palynostratigraphy of the Miocene Manuherikia Group, New Zealand. Journal of the Royal Society of New Zealand 28: 405-420. Pole, M.; Douglas, B. & Mason, G. 2003. The terrestrial Miocene biota of southern New Zealand. Journal of the Royal Society of New Zealand 33: 415-426. Raikow, R. J. 1971. The osteology and taxonomic position of the white-backed duck, Thalassornis leuconotus. Wilson Bulletin 83: 270-277. Reid, J. R. W.; Badman, F. J. & Parker, S. A. 1990. Birds. Pp 169-182, in Natural History of the North East Deserts, Tyler, M. J.; Twidale, C. R.; Davies, M. & Wells C. B. (eds). Occasional Publications of the Royal Society of South Australia 6. 367

Rest, J. S.; Ast, J. C.; Austin, C. C.; Waddell, P. J.; Tibbets, E. A.; Hay, J. M. & Mindell, D. P. 2003. Molecular systematics of primary reptilian lineages and the tuatara mitochondrial genome. Molecular Phylogenetics and Evolution 29: 289-297. Rich, P. V. 1976. The history of birds on the island continent Australia. Pp 53-82, in Frith, H. J. & Calaby, J. H. (eds), Proceedings of the 16th International Ornithological Congress. Australian Academy of Science, Canberra, 765 pp. Rich, P. V. 1979. The Dromornithidae, an extinct family of large ground birds endemic to Australia. Bureau of Mineral Resources, Geology and Geophysics, Australia. Bulletin 184: 1-196. Rich, P. & van Tets, J. [G.F.] 1982. Fossil birds of Australia and New Guinea: their biogeographic, phylogenetic and biostratigraphic input. Pp 235-384, in Rich, P. V. & Thompson, E. M. (eds), The fossil vertebrate record of Australasia. Monash University Offset printing Unit, Clayton, Victoria, Australia, 759 pp. Rich, P. V. & van Tets, G. F. 1981. The fossil pelicans of Australia. Records of the South Australian Museum 18: 235-264. Rich, P. V.; van Tets, G. F.; Rich, T. H. V. & McEvey, A. R. 1987. The Pliocene and Quaternary flamingoes of Australia. Memoirs of the Queensland Museum 25: 207- 225. Rich, T. H.; Archer, M.; Hand, S. J.; Godthelp, H.; Muirhead, J.; Pledge, N. S.; Flannery, T. F.; Woodburne, M. O.; Case, J. A.; Tedford, R. H.; Turnbull, W. D.; Lundelius, E. L.; Rich, L. S. V.; Whitelaw, M. J.; Kemp, A. & Rich, P. V. 1991. Australian Mesozoic and tertiary terrestrial mammal localities. Pp 1005-1070, in Rich, P.V., Monaghan, J.M., Baird, R.F., Rich, T.H. (eds), Vertebrate Palaeontology of Australasia. Pioneer Design Studio and Monash University Publications Committee, Melbourne, 1437 pp. Roelants, K. & Bossuyt, F. 2005. Archaeobatrachian paraphyly and Pangaean diversification of crown-group frogs. Systematic Biology 54: 111-126. Sangster, G.; Collinson, J. M.; Helbig, A. J.; Knox, A. G.; & Parkin D. T. 2005. Taxonomic recommendations for British Birds: third report. Ibis 147: 821-826. Scanlon, J. D. 2006. Origins and radiations of snakes in Australasia. Pp 309-330, in Merrick, J. R., Archer, M., Hickey, G. M. & Lee, M. S. Y. (eds.), Evolution and Biogeography of Australasian Vertebrates, Auscipub Pty Ltd, Oatlands, NSW, 942 pp. Sharpe, R. B. 1899. A Hand-list of the Genera and Species of Birds. Vol. 1. British Museum (Natural History), London, xxi + 303 pp. Shepherd, L. D. & Lambert, D. M. 2007. The relationships and origins of the New Zealand wattlebirds (Passeriformes, Callaeatidae) from DNA sequence analyses. Molecular Phylogenetics and Evolution 43: 480-492. Short, L. L. 1970. A new anseriform genus and species from the Nebraska Pliocene. Auk 87: 537- 543. Shufeldt, R. W. 1909. Osteology of Birds. New York State Museum Bulletin 130: 1-381. 368

Shufeldt, R. W. 1913. On comparative osteology of Cereopsis novaehollandiae. Emu 12: 209-237. Shufeldt, R. W. 1914. Contributions to the study of the “tree-ducks” of the genus Dendrocygna. Zoologische Jahrbücher (Systematik, Geographie und Biologie der Tiere) 38: 1-70. Sibley, C. G. & Ahlquist, J. E. 1990. Phylogeny and Classification of Birds: A Study in Molecular Evolution. Yale University Press, New Haven, CT, xxiii + 976 pp. Sibley, C. G.; Ahlquist, J. E. & Monroe, B. L. 1988. A classification of the living birds of the world based on DNA-DNA hybridization studies. Auk 105: 409-423. Sibley, C. G. & Monroe, B. L. Jr. 1990. Distribution and taxonomy of birds of the World. Yale University Press, New Haven, CT, xxiv + 1111 pp. Slack, K. E.; Jones, C. M.; Ando, T.; Harrison, G. L.; Fordyce, R. E.; Arnason, U. & Penny, D. 2006. Early penguin fossils, plus mitochondrial genomes, calibrate avian evolution. Molecular Biology and Evolution 23(6): 1144-1155. Slikas, B.; Olson, S. L. & Fleischer, R. C. 2002. Rapid independent evolution of flightlessness in four species of Pacific Island rails (Rallidae): an analysis based on mitochondrial sequence data. Journal of Avian Biology 33: 5-14. Sorenson, M. D.; Cooper, A.; Paxinos, E. E.; Quinn, T. W.; James, H. F.; Olson, S. L. & Fleischer, R. C. 1999. Relationships of the extinct moa-nalos, flightless Hawaiian waterfowl, based on ancient DNA. Proceedings of the Royal Society of London, Series B 266: 2187-2193. Sraml, M.; Christidis, L.; Easteal, S.; Horn, P. & Collet, C. 1996. Molecular relationships within Australasian waterfowl (Anseriformes). Australian Journal of Zoology 44: 47-58. St. John, J.; Cotter, J. -P. & Quinn, T. W. 2005. A recent chicken repeat 1 retrotransposition confirms the Coscoroba-Cape Barren goose clade. Molecular Phylogenetics and Evolution 37: 83-90. Steadman, D. W. 2006. Extinction & Biogeography of Tropical Pacific Birds. University of Chicago Press, Chicago and London, xiv + 594 pp. Stirton, R. A.; Tedford, R. H. & Miller, A. H. 1961. Cenozoic stratigraphy and vertebrate paleontology of the Tirari Desert, South Australia. Records of the South Australia Museum 14: 19-61. Stirton, R. A.; Tedford, R. H. & Woodburne, M. O. 1967. A new Tertiary formation and fauna from the Tirari Desert, South Australia. Records of the South Australia Museum 15: 427-462. Sutherland, R. 1999. Basement geology and tectonic development of the greater New Zealand region: an interpretation from regional magnetic data. Tectonophysics 308: 341-362. Švec, P. & Mlíkovský, J. 1986. First record of the genus Oxyura (Aves: Anseridae). asopis pro Mineralogii a Geologii 31: 403-407, pl. 1. Tambussi, C. & Noriega, J. 1996. Summary of the avian fossil record from Southern South America. Münchner Geowissenschaftliche Abhandlungen, Reihe A, 30: 245-264. 369

Tambussi, C. & Acosta Hospitaleche, C. 2007. Antarctic birds (Neornithies) during the Cretaceous- Eocene tines. Revista de la Asociación Geológica Argentina 62(4): 604-617. Tavares, E. S.; Baker, A. J.; Pereira, S. L. & Miyaki, C. Y. 2006. Phylogenetic relationships and historical biogeography of neotropical parrots (Psittaciformes: Psittacidae: Arini) inferred from mitochondrial and nuclear DNA sequences. Systematic Biology 55: 454-470. Tedford, R. H.; Archer, M.; Bartholomai, A.; Plane, M.; Pledge, N.S.; Rich, T.; Rich, P. & Wells, R. T. 1977. The discovery of Miocene vertebrates, Lake Frome area, South Australia. BMR Journal of Australian Geology & Geophysics 2: 53-57. Tedford, R. H. & Wells, R. T. 1990. Pleistocene deposits and fossil vertebrates from the “Dead Heart of Australia”. Memoirs of the Queensland Museum 28: 263-284. Tedford, R. H.; Wells, R. T. & Barghoorn, S. F. 1992. Tirari Formation and contained faunas, Pliocene of the Lake Eyre Basin, South Australia. The Beagle, Records of the Northern Territory Museum of Arts and Sciences 9: 173-194. Tedford, R. H.; Williams, D. & Wells, R. T. 1986. Late Cenozoic sediments and fossil vertebrates. Pp 42-72, in Wells, R.T. & Callen, R.A. (eds) The Lake Eyre Basin – Cainozoic sediments, fossil vertebrates and , landforms, silcretes and climatic implications. Australasian Sedimentologists Group Field Guide Series No. 4. Geological Society of Australasia, Sydney, 177 pp. Tonni, E. P. 1980. Un nuevo anseriforme de sedimentos marinos Terciarios de Chubut, Argentina. El Hornero 12: 11-15. Travouillon, K.; Archer, M.; Hand, S. J. & Godthelp, H. 2006. Multivariate analyses of Cenozoic mammalian faunas from Riversleigh, northwestern Queensland. Alcheringa Special Issue 1: 323-349. Turbott, E. G. (convener) 1990. Checklist of the and the Ross Dependency, Antarctica. Third Edition. Ornithological Society of New Zealand, Inc., Random Century, New Zealand, xv + 247 pp.. van Tets, G. F. & Rich, P. V. 1990. An evaluation of De Vis’ fossil birds. Memoirs of the Queensland Museum 28: 165-168. Verheyen, R. 1955a. La systématique des Ansériformes basée sur l’ostéologie comparée. Institut Royal des Sciences Natuelles de Belgique Bulletin 31(35): 1-18. Verheyen, R. 1955b. La systématique des Ansériformes basée sur l’ostéologie comparée. Institut Royal des Sciences Natuelles de Belgique Bulletin 31(36): 1-16. Verheyen, R. 1955c. La systématique des Ansériformes basée sur l’ostéologie comparée. Institut Royal des Sciences Natuelles de Belgique Bulletin 31(37): 1-22. Verheyen, R. 1955d. La systématique des Ansériformes basée sur l’ostéologie comparée. Institut Royal des Sciences Natuelles de Belgique Bulletin 31(38): 1-16. 370

Vickers-Rich, P. 1991. The Mesozoic and Tertiary history of birds of the Australian plate. Pp 721- 808, in Rich, P. V., Monaghan, J. M., Baird, R. F. & Rich T. H. (eds), Vertebrate palaeontology of Australasia. Pioneer Design Studio and Monash University Publications Committee, Melbourne, 1437 pp. Waters, J. M. & Craw, D. 2006. Goodbye Gondwana? New Zealand biogeography, geology, and the problem of circularity. Systematic Biology 55: 351-356. Wetmore, A. 1943. Remains of a swan from the Miocene of Arizona. Condor 45: 120. White, M. E. 2006. Environments of the geological past. Pp 17-50, in Merrick, J. R., Archer, M., Hickey, G. M. & Lee, M. S. Y. (eds.), Evolution and Biogeography of Australasian Vertebrates, Auscipub Pty Ltd, Oatlands, NSW, 942 pp. Wiens, J. J.; Bonett, R. M. & Chippindale, P. T. 2005. Ontogeny discombobulates phylogeny: Paedomorphosis and higher-level salamander relationships. Systematic Biology 54: 91-110. Wiley, E. O. 1981. Phylogenetics: The Theory and Practice of Phylogenetic Systematics. J. Wiley and Sons, New York, New York, xv + 439 pp. Woodburne, M. O.; Macfadden, B. J.; Case, J. A.; Springer, M. S.; Pledge, N. S.; Power, J. D.; Woodburne, J. M. & Springer, K. B. 1994. Land mammal biostratigraphy and magnetostratigraphy of the Etadunna Formation (Late Oligocene) of South Australia. Journal of Vertebrate Paleontology 13: 483-515. Woolfenden, G. E. 1961. Postcranial osteology of the waterfowl. Bulletin of the Florida State Museum. (Biol. Sci.) 6: 1-129. Worthy, T. H. 1988. Loss of flight ability in the extinct New Zealand duck Euryanas finschi. Journal of Zoology, London 215: 619-628. Worthy, T. H. 1995. Description of some post-cranial bones of Malacorhynchus scarletti, a large extinct pink-eared duck from New Zealand. Emu 95: 13-22. Worthy, T. H. 1997. A mid-Pleistocene rail from New Zealand. Alcheringa 21: 71-78. Worthy, T. H. 2002. The New Zealand musk duck (Biziura delautouri Forbes 1892). Notornis 49: 19-28. Worthy, T. H. 2004. The Holocene fossil waterfowl fauna of Lake Poukawa, North Island, New Zealand. Tuhinga 15: 77-120. Worthy, T. H. 2005. A new species of Oxyura (Aves: Anatidae) from the New Zealand Holocene. Longmann Symposium. Memoirs of the Queensland Museum 51: 255-272. Worthy, T. H. & Gill, B. J. 2002. New distributional records of the extinct New Zealand duck Malacorhynchus scarletti (Anatidae). Records of the Auckland Museum 39: 49-52. Worthy, T. H. & Holdaway R. N. 2002. The Lost World of the Moa: Prehistoric Life of New Zealand. Indiana University Press, Indiana, xxxiii + 718 pp. 371

Worthy, T. H. & Lee, M. S. Y. 2008. Affinities of Miocene (19-16 Ma) waterfowl (Anatidae: Manuherikia, Dunstanetta and Miotadorna) from the St Bathans Fauna, New Zealand. Palaeontology 51(3): 677-708. Worthy, T. H. & Olson S. L. 2002. Relationships, adaptations, and habits of the extinct duck ‘Euryanas’ finschi. Notornis 49: 1-17. Worthy, T. H.; Edwards, A. R. & Millener, P. R. 1991. The fossil record of moas (Aves: Dinornithiformes) older than the Otira (last) Glaciation. Journal of the Royal Society of New Zealand 21: 101-118. Worthy, T. H.; Holdaway, R. N.; Sorenson, M. D. & Cooper, A. C. 1997. Description of the first complete skeleton of the extinct New Zealand goose Cnemiornis calcitrans Owen, (Aves: Anatidae), and a reassessment of the relationships of Cnemiornis. Journal of Zoology, London 243: 695-723. Worthy, T. H.; Tennyson, A. J. D.; Archer, M.; Musser A. M.; Hand, S. J.; Jones, C.; Douglas, B. J. McNamara, J. A. & Beck, R. M. D. 2006. Miocene mammal reveals a Mesozoic ghost lineage on insular New Zealand, southwest Pacific. Proceeding of the National Academy of Sciences of the United States of America 103(51): 19419-19423. Worthy, T. H.; Tennyson, A. J. D.; Hand, S. J. & Scofield, R. P. in press. A new species of the diving duck Manuherikia and evidence for geese (Aves: Anatidae: Anserinae) in the St Bathans Fauna (Early Miocene), New Zealand. Journal of the Royal Society of New Zealand. Worthy, T. H.; Tennyson, A. J. D.; Jones, C. & McNamara, J. A. 2002a. A diverse early-Miocene (15-20 Ma) terrestrial fauna from New Zealand reveals snakes and mammals. IPC2002, Geological Society of Australia Abstracts 68: 174-175. Worthy, T. H.; Tennyson, A. J. D.; Jones, C. & McNamara, J. A. 2002b. The Early-Miocene (15-20 Ma) Manuherikia Group reveals New Zealand’s first diverse Tertiary terrestrial fauna. P. 58 in H. R. Grenfell (ed.) Abstracts and Programme, Geological Society of New Zealand Annual Conference, “Northland 2002”, Geological Society of New Zealand Miscellaneous Publication 112A. Worthy, T. H.; Tennyson, A. J. D.; Jones, C.; McNamara, J. A. & Douglas, B. J. 2007. Miocene waterfowl and other birds from Central Otago, New Zealand. Journal of Systematic Palaeontology 5(1): 1-39. Worthy, T. H.; Wragg, G. M. 2003. A new species of Gallicolumba: Columbidae from Henderson Island, Pitcairn Group. Journal of the Royal Society of New Zealand 33: 769-793.

372

373

’ Miocene mammal reveals a Mesozoic ghost lineage on insular

New Zealand, southwest Pacific

By Trevor H. Worthy*†, Alan J.D. Tennyson‡, Michael Archer§, Anne M. Musser¶, Suzanne

J. Hand§, Craig Jones , Barry J. Douglas**, James A. McNamara††, and Robin M.D. Beck§



*

]

"

$

**

††



Proceeding of the National Academy of Sciences of the United States of America2006, 103(51): 19419-19423.

A Worthy, T.H., Tennyson, A.J.D., Archer, M., Musser, A.M., Hand, S.J., Jones, C., Douglas, B.J., McNamara, J.A. & Beck, R.M.D. (2006) Miocene mammal reveals a Mesozoic ghost lineage on insular New Zealand, southwest Pacific. Proceeding of the National Academy of Sciences of the United States of America, v. 103(51), pp. 19419-19423

A NOTE: This publication is included on pages 374-378 in the print copy of the thesis held in the University of Adelaide Library. A It is also available online to authorised users at: A http://dx.doi.org/10.1073/pnas.0605684103 379

Miocene mammal reveals a Mesozoic ghost lineage on insular

New Zealand, southwest Pacific

Supporting Information

DESCRIPTION AND COMPARISONS

The very fragmentary nature of the material and lack of comparative characters (notably almost all dental features) means that any conclusions regarding the placement of the St

Bathans (SB) mammal within current mammaliaform phylogenies, for example (1,2), must be tentative. We also note that ontogenetic variation as described for Haldanodon femora

(2) may be significant. Nevertheless, the grade of mammal or near-mammal represented can be bracketed, based on the available evidence. Comparisons that follow are made in the context of the phylogenetic framework of Luo et al. (1, fig. 1) and subsequent phylogenetic analyses (2,3,4,5) that have employed updated and modified versions of the original matrix used by Luo et al. (1), and which have supported broadly similar topologies.

Notes on phylogenetic framework. The composition of crown-group Mammalia is currently the subject of much debate. The relative phylogenetic positions of and are unstable, placed either within crown Mammalia or outside of it

(see [1] for a review). This is in part dependant on the placement of monotremes, which, as living mammals, root the tree of crown Mammalia. Of particular relevance to this study is the position of Eutriconodonta: eutriconodontans (including jenkinsi (6) and

Gobiconodon ostromi [7]) are included within crown-group Mammalia by Luo et al. (1; the topology employed here) but are considered by others (6,8) to lie outside crown-group

Mammalia, being regarded by these authors to be more primitive than monotremes but more derived than morganucodontans. In fact, Luo et al. (1) found only weak support for inclusion of Eutriconodonta within crown-group Mammalia. Their alternative placement of

1 380

Eutriconodonta outside Mammalia does not differ significantly from their most parsimonious tree in which Eutriconodonta falls within Mammalia (1, p. 34).

Femur (Supporting Information Fig. 4A-J)

(a) Fovea for the acetabular ligament on femoral head (fc in Supporting Information Fig.

4): The St Bathans (SB) femur has a diffuse, rugose depression on the ventromedial part of the head, interpreted here as a fovea capita for the acetabular ligament connecting the femur to the hip socket (Fig. 2; Fig. 4H). A fovea capita that appears roughly similar in development and position to that of the SB mammal is present in morganucodontans (9,10) where it has been described as ‘a shallow, and in some specimens, a rugose depression on the ventromedial aspect of the head’ (9, p. 407). The fovea is marginally more medial in morganucodontans than in the SB femur, where the fovea is more ventrally positioned. A ventrally placed fovea capita (on the flexor surface of the femur) is present in multituberculates (11). A fovea capita is present in Megazostrodon (1) (considered by some to be a docodont, 12) and holotherians ( (1,2,10,13); Vincelestes (1);

Didelphis (1); Pucadelphys (1,14); and Erinaceus (1). A fovea capita is absent in

Probainognathus (1); tritylodontids (10,15,16); Sinoconodon (1), Haldanodon (2);

Jeholodens (1,10); (7,10); and monotremes (Ornithorhynchus and

Tachyglossus).

Possession of a fovea capita is variable between groups: it is present in the most primitive known mammals (morganucodontans and possibly docodonts) but is not seen in several more derived groups (e.g., Eutriconodonta and monotremes). Its presence may also be variable within groups: it is found in Megazostrodon but not in the docodont Haldanodon

(2). Degree of development of the fovea capita varies within groups; for example, in some basal marsupials the fovea capita is described as faint (Pucadelphys and Monodelphis) but in others it can be deep and well defined (e.g., Metachirus) (14). It is therefore difficult to

2 381

assess the significance of the fovea capita in the SB mammal although it is at least as derived as morganucodontans in this mammalian-like feature.

(b) Orientation and shape of head/development of neck: The head of the St Bathans (SB) femur is a rounded knob (sub-spherical) emerging from the femoral shaft without support of a well-developed neck (Fig. 2; Fig. 4G and H). The head has a slight mediodorsal inclination and is not inclined at a definite angle to the shaft.

The head and neck region of femora in certain eucynodonts and basal mammals is very similar to that of the SB mammal e.g., the advanced eucynodonts Probainognathus (1),

Oligokyphus (2,15), Bienotherium (17) (Fig. 4A and B), and Bienotheroides (18).

Dorsomedial orientation of the head is seen in and primitive mammals (9,10).

In morganucodontans, a hemispherical head is dorsomedially reflected from a very short neck (9,10) (Fig. 4E and F); the neck is regarded as absent by (1). The sub-spherical rather than globular shape of the femoral head in Haldanodon (Fig. 4C and D) may be a fossorial (19), but alternatively may be a retained plesiomorphic condition (2).

Head shape appears to be related to ontogenetic development, as older individuals of

Haldanodon have a more rounded head than smaller, presumably younger specimens (2).

In Ornithorhynchus the head is sub-spherical and the neck is incipient. In the eutriconodontan Jeholodens the femur lacks a constricted neck and the head is oriented dorsally (1,2). The unusual femur of the eutriconodontan Gobiconodon has a bulbous femoral head (compressed post-mortem) and broad, flattened, neck (2,7,10), which in many ways is very different from those of morganucodontan-like mammals (10).

The derived mammalian state of a spherical, medially-inflected head on an elongate, well-defined neck (Fig. 4I and J) is almost universal in crown-group Mammalia more derived than monotremes and eutriconodontans, where the femur is known (1,20), including cimolodontan multituberculates (1,11); (1,2,20,21);

Henkelotherium (1,2,13); Vincelestes (1); marsupials (e.g., Pucadelphys [1,14]); and

3 382

placentals (e.g., Erinaceus [1]). An exception in therian mammals is the recently described symmetrodont Akidolestes, in which the dentition, shoulder girdle and forelimb are typically therian but the hindlimb (including the head, neck and trochanteric region of the femur) is similar to that of morganucodontans, eutriconodontans and, to a lesser extent, monotremes

(3). This has been interpreted as convergence due to parallel locomotor adaptations (3).

The SB femur lacks the apomorphic state of a spherical head and well developed neck seen in all crown group Mammalia exclusive of monotremes, Jeholodens and probably

Akidolestes, exhibiting a plesiomorphic state generally similar to that of Haldanodon, morganucodontans, monotremes and certain very advanced eucynodonts such as tritylodontids.

(c) Orientation and form of greater trochanter: The greater trochanter of the SB femur arises from the lateral side of the shaft (directed dorsolaterally per [1]) and is comparatively large and triangular (wing-shaped) (Fig. 2; Fig. 4G and H). It is low in height, not quite reaching the level of the top of the femoral head. There is a distinct muscle scar running proximally in an arc from the distolateral margin of the dorsal surface of the greater trochanter to the middle of the shaft (similar to the morphology of this area in Haldanodon

(2, p. 230, fig. 12A). The greater trochanter is proximally directed with a distinct apex (agt in Fig. 4), as in Haldanodon, morganucodontans and other early non-therian mammals (2).

Presence of a third trochanter (as in the femur of Haldanodon, 2) cannot be determined because the specimen is broken proximal to this point. There is no digital fossa developed on the ventral aspect of the greater trochanter (see (e) below).

Laterally enlarged, ‘wing-shaped’ greater trochanters are characteristic of tritylodontids (9,15,17,18); Therioherpeton (3); morganucodontans (9,10); Haldanodon

(2,22); and Ornithorhynchus (23). Ornithorhynchus differs, however, in that the greater trochanter is proportionately much larger than in these other taxa and extends much further distally, and most likely represents a fossorial/aquatic adaptation. Enlargement of the

4 383

greater (as well as lesser) trochanters is a primitive configuration not seen in therian mammals (9) or in multituberculates (11); in these mammals the greater trochanter appears in dorsal/anterior view as a proximal continuation of the femoral shaft rather than as a triangular wing (9). The greater trochanter in Jeholodens is much smaller than in

Haldanodon (2) and would therefore be relatively smaller than that of the SB femur. The height of the greater trochanter at or near the level of the head is similar to tritylodontids

(15, 17); tritheledontids (Hopson, pers. comm. in [3]); Therioherpeton (24); Haldanodon (2) and juvenile Ornithorhynchus. This is not so in morganucodontans, where the head is higher than the greater trochanter (9,10).

The greater trochanter is oriented more dorsally/anteriorly (losing any lateral orientation) and is more erect in multituberculates (1,11); Zhangheotherium (1,10,20,21);

Henkelotherium (1,10); Vincelestes (1); marsupials (e.g., Pucadelphys (14); and placentals

(e.g., Erinaceus [1]).

(d) Development of notch between greater trochanter and femoral head: There is a moderately deep notch between the greater trochanter and the femoral head (Fig. 2; n in Fig.

4) similar in extent to the notch in Haldanodon (2). A distinct notch between the greater trochanter and the head is absent in most cynodonts (incipient in Cynognathus [25]), but is present in some eucynodonts (e.g., Pachygenelus [26]) and tritylodontids [15,17,26,27]). A notch is present in morganucodontans (9,26) and in Haldanodon (2), as well as crown-group

Mammalia (e.g., 1). A well developed notch was considered a synapomorphy for

Mammaliamorpha (Tritylodontidae + Mammalia), here including the eucynodont taxa listed above (27). In this mammal-like character, therefore, the SB femur is more derived than most (but not all) eucynodonts.

(e) Ventral intertrochanteric fossa (itf in Fig. 4): The SB femur has a gently concave, shallow intertrochanteric fossa on the ventral aspect of the shaft between greater and lesser trochanters (Fig. 2). The limits of this fossa appear less well defined than in Morganucodon,

5 384

where a more distinct intertrochanteric ridge is formed. Curved, incised grooves appear to be muscle attachment scars (as described by [28] for the Stonesfield femur). The ventral part of the head overhangs the fossa, as described in Oligokyphus (15).

In basal mammals with a shallow intertrochanteric fossa (e.g., Morganucodon), the m. obturator externus inserts onto the fossa near the centre of the shaft (23). In

Ornithorhynchus the m. obturator internus attaches broadly over the centre of the shallow intertrochanter fossa and extends ventrally towards the lesser trochanter. The

The gluteus medius and gluteus maximus insert on the greater trochanter, and appear to be the corollories of the caudo-ilio-femoralis in reptiles. In most mammals, only the m. obturator externus inserts into the excavated digital fossa (23,28), which is a pocket or deep fossa formed by the inturned medial margin on the ventral aspect of the greater trochanter (25). An intermediate stage is described for the Late Purbeck femora

(23), in which the fossa has deepened and is more excavated than in basal mammals but a true digital fossa is not developed.

In general form, the intertrochanteric fossa on the SB femur is similar to that of

African cynodonts (29); tritylodontids (15,17,28); morganucodontans (9,10); monotremes

(28 and personal observation); and probably Haldanodon (damage obscures its form and extent, 2). The femur of Gobiconodon has a damaged greater trochanter, but it lacks a deep digital fossa and apparently has a shallow centrally located intertrochanteric fossa (7).

Multituberculates have a small, possibly divided digital fossa (11). Zhangheotherium has a tall greater trochanter with an expanded, inturned apex and triangular intertrochanteric fossa

(21). The intertrochanteric fossa of Henkelotherium is described as deep (10); a digital fossa is not described. As stated above, all living mammals have an obturator fossa or pit (23,28).

The absence of a well developed digital fossa in the SB femur is a plesiomorphic trait shared with eucynodonts, basal mammals and monotremes and excludes it from most crown

Mammalia, including multituberculates (11), marsupials and placentals. Early therians show

6 385

intermediate stages in the development of a digital fossa. Lack of development of a digital fossa suggests the SB femur is at the pre-holotherian, pre-multituberculate level for this character.

(f) Position of the lesser trochanter: The lesser trochanter of the SB femur is broken proximally (at its origin; olt in Fig. 4G and H); however, it appears to have originated on the ventromedial side of the shaft distal to the origin of the greater trochanter (Fig. 2).

Unfortunately, the size and form of the lesser trochanter of the SB femur cannot be determined with any confidence since all of the trochanter except its point of origin is missing.

A ventromedial lesser trochanter distal to the greater trochanter is the primitive conformation for Cynodontia (26) and is seen in Eucynodontia, in most Cynognathia

(including some tritylodontids: Bienotherium [17] and Bienotheroides [18]) and

Probainognathia (Probainognathus, Probelesodon, and Chiniquodon [26]) (we note that

Probainognathus, contra [26], is reported to have a medial lesser trochanter by [1]). Some eucynodonts (e.g., the tritylodontid Oligokyphus [15]) and all basal mammals) have a more medially placed lesser trochanter, interpreted as a synapomorphy uniting the eucynodont

Pachygenelus + Morganucodon (26). The basal mammals Sinoconodon, Morganucodon,

Megazostrodon, and Haldanodon; the eutriconodontans Jeholodens and Gobiconodon; and

Vincelestes have a medial lesser trochanter (1). In monotremes, the lesser trochanter is ventromedial in Tachyglossus but medial in Ornithorhynchus. Oligokyphus and

Ornithorhynchus are considered specialized in sharing this medial position of the lesser trochanter (15,30).

A ventromedial or ventral position for the lesser trochanter is found in multituberculates (1,11) and therian mammals (1,30).

As a distal position of the lesser trochanter is considered plesiomorphic in cynodonts

(15), can be interpreted as retaining the plesiomorphic state of a more distal and

7 386

ventromedial lesser trochanter. That being so, femora of Oligokyphus, Ornithorhynchus and basal mammals (listed above), which have a distal and medial origin to the lesser trochanter are derived in this feature (15).

The ventromedial, distal position of the lesser trochanter of the SB femur is therefore a plesiomorphic feature shared with basal eucynodonts, Bienotherium, Bienotheroides,

Tachyglossus, and therian mammals. The SB femur differs from the tritylodontid

Oligokyphus, the Ornithorhynchus, and, to a lesser extent, Sinoconodon,

Morganucodon and Haldanodon in the ventral rather than medial position of the lesser trochanter.

(g) Ridge joining greater trochanter to femoral head: The SB femur lacks a bony ridge running from the greater trochanter to the femoral head, present in most cynodonts but absent (the derived state for cynodonts) in tritylodontids and most mammals. A short ridge is reported between the head and greater trochanter in morganucodontans (9,10) although this character was coded as absent in Morganucodon (26). For this character, therefore, the

SB femur is more derived than most (but not all) eucynodonts and Morganucodon.

(h) Epiphyses: The degree of development of muscle attachment areas and the high density of the superficial bone surface suggests that the SB femur is from an adult rather than a juvenile. A fovea capita is present, which would be on the epiphysis of the femoral head if epiphyses were present. The SB femur lacks any indication of secondary ossifications

(epiphyses): no undoubted suture lines can be seen on any part of the femur. The erosion of the bone surface on the head reveals an evenly large 'cellular' structure to the 'spongy bone ' across the zone of a potential epiphysis, rather than any narrow zone of dense bone expected at the fusion of two surfaces. However, a seamless fusion in adulthood of an epiphysis with the subsequent resorption of the bone at the symphysis to remove all sign of the former surfaces cannot be ruled out. As preserved, the structure of the SB mammal appears very similar to that of Haldanodon, which has been interpreted as lacking epiphyses (2). We

8 387

conclude that the SB mammal, therefore, probably lacked epiphyses, but that their presence cannot be ruled out. Epiphyses are not developed in ancestral tetrapods (27); cynodonts including Oligokyphus (15,27,39); morganucodontans (10,27,31); Haldanodon (2); or

Jeholodens (9). Martin suggested that the lack of epiphyses in Haldanodon indicates continuous growth throughout the life of the animal (2). Epiphyses are considered a synapomorphy of crown-Mammalia (15,27), being present in monotremes (27),

Gobiconodon (32), multituberculates (11,27), and Theria (27), although Jeholodens lacks these, as mentioned above. Although epiphyses are present in monotremes, these are avascular (unlike the epiphyses of therian mammals) and may have been independently derived from those of therians (33).

The presence or absence of epiphyses is not scored in the phylogenetic analysis presented in Supporting Information Fig. 5, and thus does not affect the placement there of the SB mammal in a trichotomy with multituberculates and immediately above the eutriconodontans Gobiconodon and Jeholedons, in which group typical mammalian-like epiphyses first appear. An absence of epiphyses on the SB femur would make it more primitive in this character than any crown-Mammalia except for Jeholodens (6) and monotremes, where epiphyses, as stated, may be independently derived. Epiphyses, and hence definitive (rather than lifelong or indeterminate) growth, are characteristic of crown- group mammals (e.g., 27,31,34).

(i) Posture: Position of the fovea capita near the top of the head (35) and lack of a well developed neck (3) suggest an abducted or sprawling femoral posture for the SB mammal.

Abduction of the hindlimb (a ‘sprawling’ posture) is seen in morganucodontans (10), monotremes (10,36), Jeholodens (6), multituberculates (11), Haldanodon (2,24), and

Akidolestes (3). The more ventral lesser trochanter of the SB femur suggests it was not as horizontally oriented as in the extreme abducted posture of monotremes or Oligokyphus

(29,30,36). Abduction of the hindlimb is a more plesiomorphic posture than that of therian

9 388

mammals (symmetrodonts except Akidolestes, ‘prototribosphenidans’, marsupials and placentals) where a more erect (parasagittal) posture is progressively developed (e.g., 3).

Mandible

(a) Dental formula: Because of damage to both specimens, the anterior lower dental formula is uncertain; the posterior lower dental formula is unknown because that part of the jaw is not preserved. MNZ S.41866, with reasonably well-preserved right alveoli, is the most informative. There are six alveoli present (Fig. 1), although the last, number 6, is incomplete. There clearly was one large, semi-procumbent incisor in the first (most anterior) alveolus. If there were other incisors, they had to be much smaller and mesial to this large . Five alveoli posterior to what we identify as the incisor region are evident. The first, alveolus 2, is large and oval, with a slight lateral bulge. It presumably represents a moderately procumbent canine, given the possible identity of the next alveoli. Of these alveoli, 3 is somewhat anteroposteriorly compressed and 4 is round in shape. Alveolus 3 is the smallest of the preserved alveoli. Alveolus 4 is larger, being only slightly smaller than 2 on this specimen (4 appears to be larger than 2 on the left side of MNZ S.40958). There is a thickened median saddle as part of the alveolar septum, which 3-4 share. From the midpoint of this saddle, two vertical ridges descend straight down into the alveoli, one on the anterior face and one on the posterior face of the septum. The same situation is evident for the common shared septum between alveoli 5 and 6. Alveoli 1-4 all show a degree of procumbency, which is more pronounced anteriorly. Alveolus 5 is somewhat round in shape

(although this varies depending on its preservation), non-procumbent, and shows some lateral bulging. Alveolus 6 is represented only by its anterior face and little can be said about its shape or size. All alveoli have a slight lateral inclination, most evident in anterior

10 389

view (Fig. 2C). There are small, round nutrient foramina between several alveoli along the lateral margin of the alveolar row.

The presence of a median saddle with vertical ridges between the last four alveoli, 3-6, suggests to us that these alveoli carried two two-rooted teeth. A similar medially-located, vertically-oriented ridge descends into the alveoli on anterior and posterior faces of the shared septum of most double-rooted teeth ( and molars) in a wide variety of mammals. These bony structures on the inter-radicular septa fit into the medial groves on each of the partially divided roots of double-rooted teeth. This interpretation is corroborated by the fact that alveolus 4 (which would be the posterior root of the most-anterior two- rooted tooth) is larger than 5 (which would hold the anterior root for the second two-rooted tooth). It is common in mammals with two-rooted premolars for the anterior root to be smaller than the posterior root, hence the relationship we see in the St Bathans specimens.

Lack of evidence for any change in alveolar depth or abrupt change in width between alveoli 3 and 4 compared with the alveoli 5 and 6 suggests that the second two-rooted tooth was a second rather than a first . Alveoli 3 and 4 indicate the roots of the first premolar were non-divergent, while alveoli 4 and 6 indicate divergent roots on the second and larger premolar. The second specimen, MNZ S.40958, although with the more complete areas of both the left and right dentaries, is somewhat more worn than MNZ S.41866. It still reveals the same structure and, in addition, preserves the distal tips of the roots, including the dental canal, in the bases of both alveoli for the first premolar in the right dentary.

We conclude that the dentition, so far as it can be determined, suggests a contiguous series of: one lower incisor, one lower canine, and at least two double-rooted teeth immediately behind the canine, which are probably premolars but whose homology is unknown. Given this interpretation, the SB mammal has a highly autapomorphic dentition.

The form of the femur strongly suggests affinity with basal mammals for the SB mammal. Most basal mammals have a dental formula of three to four lower incisors,

11 390

including most non-tritylodontid cynodonts (3,26); Sinoconodon (37), Morganucodon

(38,39), Megazostrodon (40); Dinnetherium (41); Docodon (42); basal eutriconodontans

(6,23); and holotherians (including Kuehneotherium (38), many dryolestoids (23,43) and basal marsupials and placentals (e.g., 44). While tritylodontid cynodonts are superficially similar with a single, greatly enlarged first incisor, and tiny i2-3 in some taxa (e.g., present in Oligokyphus; absent in Kayentatherium and Bienotherium), they have no canines and a wide diastema separating the incisors from the cheekteeth (15,16,17), quite unlike the SB mammal, in which all alveoli are contiguous. The eutriconodontan Gobiconodon has a lower dental formula of i1c1p1-3(-4)m1-5 where the incisor is large and procumbent (7) and so superficially has similarities with the SB mammal; however, the premolars are single- rooted and the dentaries are unfused.

(b) Mandibular symphysis: The dentaries of both lower jaws from St Bathans are fused together from the anterior of the jaw to p2 (posterior to alveolus 5). There is a series of nutrient foramina running along the dorsal midline of the jaw. Two conjoined, asymmetrical openings on the posterior aspect of the jaw symphysis form a double canal that runs through the symphysis to the anterior of the jaw. The left opening on both jaws is the larger of the two. There are four openings between the anteriormost alveoli (for i1) that appear to represent the anterior terminus of the canal: 1) a single large, square dorsal opening; 2) two tiny bilateral openings ventral to this; and 3) a ventral opening (probably single) that is intermediate in size between (1) and (2). The asymmetry of the openings suggests passage of blood vessels and nerves (A. Greer, pers. comm.) rather than a persistent meckelian cartilage as proposed for Cynognathus (39).

Basal cynodonts have an unfused jaw, but the jaw in most Eucynodontia is fused, a state that is considered a synapomorphy of the group (26, 45, 46). However, the mandibular symphysis of some eucynodonts is unfused, e.g., Pachygenelus (26), Prozostrodon (22) and

Tritylodontidae (26). The mandibular symphysis is unfused in all basal mammals,

12 391

multituberculates, monotremes and other Mesozoic crown Mammalia (e.g., 1,27,47,48). An unfused jaw was considered a synapomorphy of Mammaliamorpha (=last common ancestor of Tritylodontidae + Mammalia + descendants) (27); amended to include tritheledontids and basal mammals by (47). Some derived Cenozoic marsupials and placentals also have fused jaws (e.g., wombats, mystacinid bats, pigs, and primates).

Possession of a fused jaw in the SB mammal is a highly distinctive character that is rare in mammals (absent to date in Mesozoic mammals). It suggests one of two possibilities for the SB mammal: 1) that if derived from basal mammalian stock (as suggested by many aspects of the femur) the SB mammal has retained or secondarily developed a eucynodont- like fused jaw (plesiomorphic retention or atavistic reversal respectively); 2) that the SB mammal is one that has independently developed a specialised, fused jaw like that of some other Cenozoic mammals.

(c) Symphyseal length: In the St Bathans mandibles the symphysis extends posteriorly to the end of alveolus 5 (i.e., the middle of p2 in our interpretation). In taxa with fused jaws, the length of the symphysis varies. Within Eucynodontia, symphyseal lengths vary from short in the traversodont Scalenodontoides macrodontes (49) and Microconodon tenuirostris (a eucynodont of uncertain affinities) (50), with the mandibles fused to just posterior to the canine alveolus (alveolus 4), to long, with fusion to the 3rd postcanine

(alveolus 7) in Probainognathus (51) and Cynognathus (39).

In eucynodonts and early mammals, where the symphysis is unfused, the symphyseal length is also variable. In Oligokyphus, the symphyseal region extends over 1/3 the length of the jaw, reaching beyond the diastema to the first molariform (15). In most early mammal groups the symphyseal region extends past the canine alveolus. In

Morganucodon the symphysis extends nearly to the 2nd postcanine (alveolus 8) (39). In

Dinnetherium the symphysis is to the 7th alveolus (2nd postcanine) (7). In Docodon the symphysis is especially long, extending to the 14th alveolus (4th postcanine) (42). In

13 392

eutriconodontans (e.g., Triconodon, [23], Gobiconodon and Klamelia [7]), the symphysis extends well beyond the canine alveolus, to the 3rd postcanine in Trioracodon

(23) and Gobiconodon ostromi (7) and to what have been identified as m2-3 in Klamelia zhaopengi (52). In symmetrodonts such as Peralestes the symphysis extends to just past the canine (alveolus 4) (53). In archaic therians such as the dryolestoid Crusafontia and amphitheriid the symphysis extends to the 7th alveolus (2nd postcanine)

(23,54) respectively.

Length of the symphyseal region, regardless of whether the jaw is fused or unfused, is therefore a variable character within both eucynodonts and mammals. Symphyseal length in the SB mammal falls within the observed variation and may therefore be of little phylogenetic significance.

(d) Position of mental foramen: Given our interpretation of the dental formula as one incisor, one canine, and two premolars, the single preserved mental foramen in the SB jaws is ventral to the first premolar (between alveoli 3-4). In basal mammals, an anterior position for the mental foramen appears to be a characteristic feature. Some specimens of

Morganucodon have the first of two or more mental foramina ventral to i4 (alveolus 4)

(positions of the mental foramina vary between individuals) (39). In Dinnetherium there is a mental foramen ventral to i2-3 (41).

In the dentate ornithorhynchid dicksoni, based on a complete skull and partial dentaries, there appears to have been a long, non-dentigenous zone between the anterior end of the symphysis and the first cheektooth (p1), and a foramen lies ventral to the very distal p2 (lower dentition is not preserved) that was designated the f. mandibulare medium (55), not the mental foramen. The lower jaw anterior to this is not preserved in

Obdurodon (55) and the possibility of additional premolars, incisors or canines cannot be ruled out, although the skull is edentate anteriorly. That the skull of Obdurodon is very similar to Ornithorhynchus suggests that its dentary was probably splayed rather than

14 393

narrowed at the anterior end, and the mandibular symphysis unfused and horizontal (rather than inclined).

Some later mammals have the mental foramina in a similar position ventral to p1 (e.g., some dryolestoids but not others [43]) but in many other Mesozoic mammals (excepting derived types like multituberculates) the mental foramen appears to be in a more posterior position (data from 4). Position of the mental foramen or foramina varies within groups; within Eutriconodonta, Trioracodon ferox has the first of two mental foramina beneath p1

(23); and Gobiconodon ostromi has the first of three 3 mental foramina below p3 (7). In basal Marsupialia the position of this foramen or foramina also varies: in Sinodelphys the anteriormost mental foramen is beneath i3 (5) and in Alphadon it is below p1 (56). In the basal placentals Kennalestes, Asioryctes, Barunlestes, and Cimolestes the anteriormost mental foramen is ventral to p1 (57).

It is therefore difficult to determine a pattern of phylogenetic significance in the position of the mental foramen.

PHYLOGENETIC ANALYSIS

In an attempt to determine the phylogenetic relationships of the SB mammal, we added it to the morphological character matrix of Li and Luo (3), which represents one of the most comprehensive and up-to-date matrices for investigating the phylogeny of mammaliaforms currently available. The original matrix comprises 413 dental, cranial and post-cranial characters scored for 74 taxa, including several cynodonts, and representatives of all major mammaliaform lineages. Based on currently known specimens, the following characters could be scored for the SB mammal, with the relevant character state indicated:

25. Posterior-most mental foramen:

15 394

(0) In the canine and anterior premolar (premolariform) region (in the saddle behind the canine eminence of the mandible); (1) Below the penultimate premolar (under the anterior end of the functional postcanine row); (2) Below the ultimate premolar; (3) At the ultimate premolar and the first molar junction; (4) Under the first molar.

SB mammal: (0) In the canine and anterior premolar (premolariform) region. Note: given that the posterior portion of the mandible of the SB mammal is unknown, this character cannot be scored with certainty; however, given the large size of the preserved mental foramen, and the absence of accessory foramina anteriad to it, we have assumed that only one such foramen was present, and hence it is the posterior-most.

37. Dentary symphysis:

(0) Fused; (1) Unfused.

SB mammal: (0) Fused.

137. Number of lower incisors:

(0) Five or more; (1) Four; (2) Three; (3) Two; (4) One; (5) No incisors.

SB mammal: (4) One.

141. Lower canine - presence vs. absence and size:

(0) Present and enlarged; (1) Present and small; (2) Absent.

SB mammal: (0) Present and enlarged.

142. Number of lower canine roots:

(0) One; (1) Two.

SB mammal: (0) One.

16 395

150. Diastema separating the first and second lower premolars (defined as the first and second functioning premolar or premolariform postcanine):

(0) Absent (gap less than one tooth root for whichever is smaller of the adjacent teeth); (1)

Present, subequal to one tooth-root diameter or more; (2) Present, equal to or more than one- tooth length.

SB mammal: (0) Absent (gap less than one tooth root for whichever is smaller of the adjacent teeth).

157. Procumbency and enlargement of the lower anterior-most incisor:

(0) Absent; (1) Present (at least 50% longer than the adjacent incisor).

SB mammal: (1) Present (at least 50% longer than the adjacent incisor).

158. Enlarged diastema in the lower incisor-canine region (better developed in older individuals):

(0) Absent; (1) Present and behind the canine; (2) Present and behind the posterior incisor.

SB mammal: (0) Absent.

222. Inflected head of the femur set off from the shaft by a neck:

(0) Neck absent and head oriented dorsally; (1) Neck present, head spherical and inflected medially.

SB mammal: (0) Neck absent and head oriented dorsally.

223. Fovea for the acetabular ligament on the femoral head:

(0) Absent; (1) Present.

SB mammal: (1) Present.

17 396

224. Orientation of the greater trochanter:

(0) Directed dorsolaterally; (1) Directed dorsally.

SB mammal: (0) Dorsolaterally.

225. Position of the lesser trochanter:

(0) On medial side of the femoral shaft; (1) On the ventromedial or ventral side of the femoral shaft.

SB mammal: (1) On the ventromedial or ventral side of the femoral shaft.

Following our initial character analysis, we also decided to add the following character of the femur, which we consider to be a potentially significant feature not included by Li and

Luo (3) or other recent phylogenetic analyses of mammaliaforms (1,2,4,5):

414. Form of ventral intertrochanteric area:

(0) Gently concave, shallow intertrochanteric fossa centred on the ventral aspect of the shaft between greater and lesser trochanters; (1) Deep fossa centred on the ventral aspect of the shaft between greater and lesser trochanters; (2) Deep fossa (digital or obturator fossa) excavated on the ventral aspect of the greater trochanter.

Thrinaxodon, tritylodontids, Pachygenelus, Megazostrodon, Morganucodon,

Ornithorhynchus, Tachyglossus, Haldanodon, Akidolestes, Gobiconodon, SB Mammal:

(0) Gently concave, shallow intertrochanteric fossa centred on the ventral aspect of the shaft between greater and lesser trochanters.

Zhangheotherium, Henkelotherium: (1) Deep fossa centred on the ventral aspect of the shaft between greater and lesser trochanters.

Multituberculates, Pucadelphys, all living therians: (2) Deep fossa (digital or obturator fossa) excavated on the ventral aspect of the greater trochanter.

18 397

All other taxa: (?)

The SB mammal is currently only known from fragmentary remains (most notably, teeth have yet to be found), and so can only be scored for the above 13 characters; hence it is

~97% incomplete. However, the relationships of this taxon can be at least tentatively assessed using these few characters, given that: 1) simulated and empirical studies suggest that even highly incomplete taxa can be accurately placed in phylogenies (58); and 2) an empirical study of mammalian morphological character matrices indicates that homoplasy is equally distributed throughout dental, cranial and post-cranial partitions of the mammalian skeleton (59). Hence, the limited material of the SB mammal may still contain sufficient phylogenetic signal to broadly elucidate its relationships, and may be at least as informative as isolated teeth.

The modified matrix, comprising 414 characters and 75 taxa, was analysed using

PAUP* 4.0b10 (60), with all characters equally weighted and unordered and

‘mstaxa=polymorph' (i.e. polymorphic characters are treated as such, rather than as uncertain or variable), following (3). 1000 heuristic treesearch replicates were initially carried out, with a maximum of ten shortest trees saved per replicate. An additional heuristic search was then carried out within the set of globally shortest trees from the first stage. A total of 192 shortest trees was found (length = 1826; CI = 0.425), and their strict consensus, given in Fig. 5, is surprisingly well-resolved: the SB mammal forms a trichotomy with

Multituberculates and (trechnotherians + ).

However, detailed consideration of the characters supporting the position of the SB mammal suggests that the evidence for this arrangement is relatively weak. Under

ACCTRAN only five (137: 3->4, 222: 0->1, 223: 0->1, 224: 0->1, 225: 0->1), and under

DELTRAN only two (137 and 223), of the characters scored for the SB mammal support a

(SB mammal + Multituberculata + (Trechnotheria + Tinodon)) clade. Furthermore, under

19 398

ACCTRAN two of these characters (222: 1->0, 224: 1->0) subsequently reverse along the branch leading to the SB mammal, whilst along the branch leading to (Trechnotheria +

Tinodon) one character reverses (225: 1->0) and lower incisor number (137) increases from one (character state 4, as in the SB mammal and multituberculates) to four (character state 1, the plesiomorphic state for (Trechnotheria + Tinodon). The least homoplastic character change supporting the position of the SB mammal is presence of a fovea for the acetabular ligament on the femoral head (223, character state 1), which was acquired convergently by morganucodontans.

To test the stability of the phylogenetic position of the SB mammal, tree searches were carried out with it constrained as lying: 1) outside (defined by the common ancestor of Sinoconodon and crown mammals), 2) outside crown Mammalia, 3) within , 4) within Monotremata, 5) within crown Monotremata, 6) within

Theria, 7) within , 8) within Marsupialia (i.e. crown Metatheria), 9) within

Eutheria, 10) within (i.e. crown ). The maximum increase in tree length was 3, when the SB mammal was constrained to lie within either Metatheria or Marsupialia, which represents an increase of only ~0.2% over the optimal tree length. The Templeton test

(61), as implemented in PAUP* (60) and with the p-value the arithmetic mean of those of each shortest tree (62), indicates that none of these ten constrained alternative positions can be rejected as significantly worse than the optimal topology (p>>0.05, assuming a one-tailed test) (63).

To further investigate the position of the SB mammal, we repeated our phylogenetic analysis but scoring this taxon for only the five femoral characters, and then scoring only the eight mandibular characters. When represented by femoral characters, the SB mammal is more derived than Adelobasileus and Sinoconodon but unresolved with respect to other mammaliform groups, whereas the eight mandibular characters alone place it as the sister to

Multituberculata, with which it shares a single lower incisor (137: 1->4).

20 399

References

1. Luo, Z. -X., Kielan-Jaworowska, Z. & Cifelli, R. (2002) Acta Palaeontol. Pol. 47, 1-78.

2. Martin, T. (2005) Zool. J. Linn. Soc. 145, 219-248.

3. Li, G. & Luo, Z.-X. (2006) Nature 439, 195-200.

4. Kielan-Jaworowska, Z., Cifelli, R. L. & Luo, Z. -X. (2004) Mammals from the age of

dinosaurs, origin, evolution and structure (Columbia University Press, New York).

5. Luo, Z.-X., Ji, Q., Wible, J. R. & Yuan, C. (2003) Science 302, 1934-1940.

6. Ji, Q., Luo, Z. -X. & Ji, S. (1999) Nature 398, 326-300.

7. Jenkins, F. A. Jr. & Schaff, C. R. (1988) J. Vert. Paleontol. 8, 1-24.

8. Rowe, T. B. (1999) Nature 398, 283-284.

9. Jenkins, F. A. Jr. & Parrington, F. R. (1976) Philos. Trans. R. Soc. Lond. Ser. B 273, 387-

431.

10. Gambaryan, P. P. & Averianov, A.O. (2001) Acta Palaeontol. Pol. 46, 99-112.

11. Krause, D. W. & Jenkins, F. A. Jr. (1983) Bull. Mus. Comp. Zool., Harv. Univ. 150,

199-246.

12. McKenna, M. C. & Bell, S. K. (1997) Classification of Mammals Above the Species

Level (Columbia Univ. Press, New York).

13. Krebs, B. (1991) Berlin. Geowiss. Abh. A 133, 1-110.

14. Marshall, L. G. & Sigogneau-Russell, D. (1995) Mem. Mus. Nat. D’Hist. Natur. 165,

91-164.

15. Kühne, W. G. (1956) The Liassic Oligokyphus. (Brit. Mus. Nat. Hist.,

London; Adlard and Son, London).

21 400

16.Sues, H. -D. (1983) Unpublished PhD dissertation (Harvard University, Cambridge).

17. Young, C. C. (1947) Proc. Zool. Soc. Lond. 117, 537-597.

18. Sun, A.-L. & Li, Y.-H. (1985) Vertebr. Palasiat. 23, 135-151.

19. Reed, C. A. (1951) Am. Midl. Nat. 45, 513-617.

20. Hu, Y., Wang, Y. Luo, Z. -X. & Li, C. (1997) Nature 390, 137-142 (and suppl. info.).

21. Luo, Z.-X. & Ji, Q. (2005) J. Mamm. Evol. 12, 337-357.

22. Bonaparte, J. F. & Barberena, M. C. (2001) Bull. Mus. Comp. Zool. 156, 59-80.

23. Simpson, G. G. (1928) A Catalogue of the Mesozoic Mammalia in the Geological

Department of the British Museum (Trustees of the British Museum, London).

24. Krusat, G. (1991) in Fifth Symposium on Mesozoic Terrestrial and Biota.

Extended Abstracts, eds. Kielan-Jaworowska, Z., Heinz, N. & Nakarem, H. A. (Contrib.

Palaeontol. Mus. Univ. Oslo 364), pp. 37-38.

25. Gregory, W. K. & Camp, C. L. (1918) Bull. Am. Mus. Nat. Hist. 38, 447-

563.

26. Hopson, J. A. & Kitching, J. W. (2001) Bull. Mus. Comp. Zool. 156, 5-35.

27. Rowe, T. (1988) J. Vertebr. Paleontol. 8, 241-264.

28. Seeley, H. G. (1879) Quart. J. Geol. Soc. Lond. 35, 591-635.

29. Jenkins, F. A. Jr. (1971) Peabody Mus. Nat. Hist. Bull. 36.

30. Parrington, F. R. (1961) Proc. Zool. Soc. Lond. 137, 285-298.

31. Carroll, R. L. (1988) Vertebrate Paleontology and Evolution (W. H. Freeman and Co.,

New York).

22 401

32. Jenkins, F. A. Jr. (1984) in Mammals: Notes for a Short Course, ed. Broadhead, T. W.

(Univ. Tennessee Dept Geol. Sci., Stud. Geol. 8), pp. 32-47.

33. Thorp, B. H. & Dixon, J. M. (1991) The Anatomical Record 229, 447-452.

34. Gow, C. E. (1985) South African J. Sci. 81, 558-560.

35. Jenkins, F. A. Jr. & Camazine, S. M. (1977) J. Zool. Lond. 181, 351-370.

36. Pridmore, P. A. (1985) J. Zool. Lond. 205, 53-73.

37. Crompton, A. W. & Sun, A. -L. (1985) Zool. J. Linn. Soc. 85, 99-119.

38. Parrington, F. R. (1973) Zool. J. Linn. Soc. 52, 85-95.

39. Kermack, K. A., Mussett, F. & Rigney, H. W. (1973) Zool. J. Linn. Soc. 53, 87-175.

40. Gow, C. E. (1986) Palaeontol. Africana 26, 13-26.

41. Jenkins, F. A. Jr., Crompton, A. W. & Downs, W. R. (1983) Science 222, 1233-1235.

42. Kron, D. G. (1979) in Mesozoic Mammals: the First Two-thirds of Mammalian History,

eds. Lillegraven, J. A., Kielan-Jaworowska, Z. & Clemens, W. A. (University of

California Press, Berkeley), pp. 91-98.

43. Krebs, B. (1971) in Early Mammals, eds. Kermack, D. M. & Kermack, K. A. (Academic

Press, London), pp. 89-102.

44. Kielan-Jaworowska, Z. (1997) Lethaia 29, 249-266.

45. Hopson, J. A. (1991) in Origins of the Higher Groups of Tetrapods, eds. Schultze, H. -P.

& Trueb, L. (Comstock Publishing Associates, Ithaca & London), pp. 635-693.

46. Martínez, R. N. May, C. L. & Forster, C. A. (1996) J. Vertebr. Paleontol. 16, 271-291.

47. Wible, J. R. (1991) J. Vert. Paleontol. 11, 1-28.

23 402

48. Luo, Z. X. (1994) in In the Shadow of Dinosaurs – Early Mesozoic Tetrapods, eds.

Fraser, N. C. & Sues, H. -D. (Cambridge University Press, Cambridge and New York),

pp. 98-128.

49. Hopson, J. A. (1984) Palaeontol. Africana 25, 181-201.

50. Sues, H. -D. (2001) Bull. Mus. Comp. Zool. 156, 37-48.

51. Romer, A. S. (1970) Breviora 344, 1-18.

52. Chow, M. & Rich, T. H. V. (1984) J. Vert. Paleontol. 3, 226-231.

53. Cassiliano, M. L. & Clemens, W. A. (1979) in Mesozoic Mammals: the First Two-thirds

of Mammalian History, eds. Lillegraven, J. A., Kielan-Jaworowska, Z. & Clemens, W. A.

(University of California Press, Berkeley), pp. 150-161.

54. Kraus, M. J. (1979) in Mesozoic Mammals: the First Two-thirds of Mammalian History,

eds. Lillegraven, J. A., Kielan-Jaworowska, Z. & Clemens, W. A. (University of

California Press, Berkeley), pp. 162-171.

55. Musser, A. M. & Archer, M. (1998) Philos. Trans. R. Soc. Lond. Ser. B 353, 1063-1079.

56. Clemens, W. A., Wilson, G. P. & Molnar, R. (2003) J. Vert. Paleontol. 23, 232-237.

57. Kielan-Jaworowska, Z., Eaton, J. G. & Bown, T. M. (1979) in Mesozoic Mammals: the

First Two-thirds of Mammalian History, eds. Lillegraven, J. A., Kielan-Jaworowska, Z.

& Clemens, W. A. (University of California Press, Berkeley), pp. 182-191.

58. Wiens, J. J. (2006) J. Biomed. Inform. 39, 34-42.

59. Sanchez-Villagra, M. R. & Williams, B. A. (1998) J. Mamm. Evol. 5, 113-126.

60. Swofford, D. L. (2000) PAUP* - Phylogenetic analysis Using Parsimony (*and other

methods). Version 4.0b (Sinaur Associates, Sunderland, MA).

61. Templeton, A. R. (1983) Evol. 37, 221-244.

24 403

62. Lee, M. S. Y. (2000) Syst. Biol. 49, 829-836.

63. Goldman, N., Anderson, J. P. & Rodrigo, A. G. (2000) Syst. Biol. 49, 652-670.

25 404

Figure captions

Fig. 4. Comparison of femora of the Saint Bathans taxon, the tritylodontid Bienotherium yunnanense, the docodont Haldanodon exspectatus, the morganucodont Morganucodon

(=Eozostrodon), and the marsupial Didelphis. (A and B) Bienotherium yunnanense, dorsal view (A), ventral view with lesser trochanter broken off (B). (C and D) Haldanodon exspectatus, dorsal view (C), ventral view (D). (E and F) Morganucodon (=Eozostrodon) dorsal view (E), ventral view (F). (G and H) St Bathans taxon, dorsal view (G, position of lesser trochanter indicated by dashed lines), ventral view (H). (I and J) Didelphis, dorsal view (I), ventral view (J). The Saint Bathans femur shares a distoventral origin of the lesser trochanter with Bienotherium; possession of a fovea capita with Morganucodon (but in a slightly more ventral position); and a distinct apex at the proximolateral tip of the greater trochanter with Haldanodon and Morganucodon. Bienotherium and Haldanodon were probably fossorial, and their femora are comparatively robust (2, 17). Abbreviations: agt, apex of greater trochanter; df, digital fossa; fc, fovea capita; fn, femoral neck; itf, intertrochanteric fossa; n, notch between greater trochanter and head of femur; olt, origin of the lesser trochanter. Not to scale. A/B and E/F reversed for comparison. A and B redrawn from (17); C and D redrawn from (2) (specimen damaged); E and F, redrawn from (9); I and

J redrawn from (29).

Fig. 5. Full version of phylogeny presented in Fig. 3, which results from parsimony analysis (using PAUP*4.0b10) of a modified version of the matrix presented in (3), with the SB mammal added. The topology is a strict consensus of 192 most parsimonious trees of length 1826 (CI = 0.425). Selected higher taxa are indicated.

26 405

406 407

’ A reassessment of the fossil goose Anser scaldii Lambrecht, 1933 (Aves: Anatidae) By Trevor H. Worthy, Storrs L. Olson & Thierry Smith ‹ ‹ ‹ Bulletin of the British Ornithologists’ Club; 2008, in press

A Worthy, T.H., Olson, S.L. & Smith, T. (2007) A reassessment of the fossil goose Anser scaldii Lambrecht, 1933 (Aves: Anatidae). Bulletin of the British Ornithologists’ Club, in press.

A NOTE: This publication is included on pages 408-414 in the print copy of the thesis held in the University of Adelaide Library. 415

Top View