A new species of the diving duck Manuherikia and evidence for geese (Aves: Anatidae: Anserinae) in...

Worthy et al.—St Bathans diving duck and goose 97

A new species of the diving duck Manuherikia and evidence for geese (Aves: Anatidae: Anserinae) in the St Bathans Fauna (Early Miocene), New Zealand

Trevor H. Worthy1, Alan J. D. Tennyson2, Suzanne J. Hand3 and R. Paul Scofield4

Abstract A new species of the diving duck Manuherikia, larger than its two congeners, is described based on a humerus and several referred ulnae from the St Bathans Fauna, lower Bannockburn Formation, Manuherikia Group, of Early Miocene (19–16 Ma) age, in New Zealand. Evidence for anserines in the St Bathans Fauna is evaluated. Two coracoid fragments, a scapula and a proximal radius reveal a goose most similar to Cereopsis and Cnemiornis, so indicating the presence of the Cnemiornis lineage in New Zealand in the Early Miocene. Two more coracoids are provisionally interpreted as a small unusual anserine perhaps close to Cereopsis. These records extend the diversity of waterfowl described from the St Bathans Fauna to six named species in four genera, plus an additional two unnamed anserines, globally the richest Miocene waterfowl fauna known.

Keywords Anatidae; duck; Early Miocene; fossils; goose

INTroDucTIoNNew Zealand is well known as the land of birds and infamous for the numbers of Recent extinctions in the avifauna. Iconic taxa such as kiwi and moa have excited scientific interest since their discovery in the early 19th century. Yet they were only a few of the many unique endemic taxa revealed as both the extant and Recent fossil faunas became better known. For example, the New Zealand acanthisittid wrens and the kakapo (Strigops habroptilus) are now understood as the sister groups to all other passerines and parrots respectively (Ericson et al. 2002; Barker et al. 2004; Tavares et al. 2006). The New Zealand wrens, moa (Dinornithiformes), and kakapo, along with many other faunal components, are assumed to have had a very long history in New Zealand, and are often interpreted as relict Gondwanan vicariant taxa (e.g., Worthy & Holdaway 2002; Gibbs 2006). Interpretation of the evolutionary history of the New Zealand fauna formerly relied mainly on relative divergence of morphology from sister taxa elsewhere (e.g., Fleming 1962, 1979). More recently, such inferences have been derived from genetic data (e.g., Driskell & Christidis 2004; Miller & Lambert 2006; Tavares

1Darling Building, DP 418, Department of Earth and Environmental Sciences, The University of Adelaide, Adelaide, South Australia 5005, Australia. [email protected]

2Museum of New Zealand Te Papa Tongarewa, PO Box 467, Wellington 6140, New Zealand. [email protected]

3School of Biological, Earth and Environmental Sciences, University of New South Wales, New South Wales 2052, Australia.

4Canterbury Museum, Rolleston Ave, Christchurch 8001, New Zealand. [email protected]

R07017; Received 14 December 2007; accepted 27 March 2008; Online publication date 23 May 2008

Journal of the Royal Society of New ZealandVolume 38, Number 2, June, 2008, pp 97–114

Journal of the Royal Society of New Zealand, Volume 38, 200898

et al. 2006; Shepherd & Lambert 2007). Resolution of the identity of the nearest relatives of some previously problematic taxa, e.g., New Zealand wattlebirds (Callaeidae) and stitchbirds (Notiomystidae), has been advanced by the use of genetic comparisons (Ewen et al. 2006; Driskell et al. 2007; Shepherd & Lambert 2007). Knowledge of the evolutionary history of most New Zealand terrestrial fauna is restricted to inferences from Recent populations, as there is a poor fossil record of taxa. Despite this, New Zealand has long been believed to have maintained a terrestrial biota from the time it separated from Gondwana and became Zealandia to the present (e.g., Fleming 1979). However, recently some geologists and palaeontologists have controversially suggested it was completely submerged during the latest Oligocene–earliest Miocene 25–22 Ma requiring the modern fauna and flora to be entirely derived by long-distance dispersal, presumably mainly from Australia (Pole 1994; Heads 2006; Waters & Craw 2006; Campbell & Hutching 2007; Landis et al. 2008). The fossil record has, until recently, been silent regarding the evolutionary history of the New Zealand terrestrial biota, especially the issues of origins of the land and freshwater birds of New Zealand, despite an outstanding Quaternary record (Worthy & Holdaway 2002). However, since 2001, the 19–16 million year old (Ma) Bannockburn Formation, Manuherikia Group, in Central Otago, has revealed the rich and diverse “St Bathans Fauna” (Worthy et al. 2007). This fauna was deposited in a large shallow freshwater lake in a warm climate, with a surrounding vegetation including casuarinas (Casuarinaceae), eucalypts (Eucalyptus: Myrtaceae) and palms (Arecaceae), in addition to the typical “New Zealand” podocarps (Podocarpaceae), Nothofagus and araucarias (Araucariaceae) (Pole & Douglas 1998; Pole et al. 2003). The avifauna, as expected in a lacustrine depositional environment, was dominated by water birds, particularly Anatidae. It included five named species of ducks in four genera (Manuherikia two spp., Dunstanetta, Matanas and Miotadorna), as well as possibly a goose (?Anserinae) (Worthy et al. 2007). These were associated with several undescribed taxa of parrots, rails, charadriiforms, and passerines. Rarer taxa included a diving petrel (Pelecanoides miokuaka), a large gruiform, a pigeon, an eagle, an owlet nightjar, and a swiftlet. Phylogenetic analyses of the three best represented anatids revealed Manuherikia and Dunstanetta to be basal oxyurines and Miotadorna, as predicted, to be a tadornine close to Tadorna (Worthy & Lee 2008). One of the most distinctive members of the New Zealand Recent avifauna is the extinct goose Cnemiornis. This was one of the world’s largest anseriforms and highly modified from its volant ancestor following loss of flight ability (Worthy & Holdaway 2002). Long regarded as closely related to the Australian Cape Barren goose Cereopsis novaehollandiae (e.g., see in Worthy & Holdaway 2002), recent analyses have treated it as a near basal anseriform in its own family, e.g., Livezey (1989, 1997), or associated in the tribe Cereopsini with Cereopsis (Worthy et al. 1997). Modifications associated with reduction of pectoral elements and increased size of pelvic elements make homology of some skeletal elements difficult to interpret, and homoplasy effects obscure their interpretation, so there is doubt as to whether cereopsines form a phyletic grade basal to anserines or are monophyletic within Anserinae (Worthy & Lee 2008). A fossil ancestor of Cnemiornis would help resolve this issue. Ongoing excavations in the Bannockburn Formation continue to reveal new taxa and additional specimens of already represented taxa. Examination of specimens during the aforementioned phylogenetic analysis (Worthy & Lee 2008) and additional specimens recovered in the 2007 and 2008 excavation seasons have revealed a further species of Manuherikia. Secondly, new specimens unquestionably referrable to anserines (geese and swans) were found in 2007 and 2008. It is the purpose of this paper to describe these specimens and to evaluate their relationships.


MATErIAlS AND METHoDSSkeletal material of a wide range of Recent and fossil anatid taxa was examined during this study, as listed in Worthy & Lee (2008). The fossils described below were especially compared with the homologous elements from other St Bathans fossil taxa listed in Worthy et al. (2007) and Worthy & Lee (2008). We recognise the following subfamilies of relevance to this study within the Anatidae: Dendrocygninae (whistling ducks); Anserinae (swans and geese), in which we include the tribe Cereopsini for Cereopsis (Cape Barren goose), and Cnemiornis (New Zealand fossil geese); Tadorninae (shelducks); Anatinae (ducks); and Oxyurinae following Turbott (1990). The latter traditionally included stiff-tailed ducks, e.g., Oxyura and Biziura, but following the extended definition of Oxyurinae in Worthy & Lee (2008), here it includes also Malacorhynchus (Australasian pink-eared ducks), Stictonetta (freckled duck), and the Oligo-Miocene fossil taxa Mionetta, Manuherikia and Dunstanetta). Measurements were made with Tesa® dial callipers and rounded to 0.1 mm. Anatomical nomenclature of specific bone landmarks follows Baumel & Witmer (1993) at first mention, with English translations or names following Howard (1929) used thereafter. Details of sites are in Worthy et al. (2007).

abbreviations: FRN, Fossil record number in the archival Fossil Record File of the Geological Society of New Zealand; Ma, million years ago; indet, indeterminate. Measurements: TL, total length; PW, proximal width; SW, shaft width; DW, distal width.

institutions: CM, Canterbury Museum, Christchurch; NMNZ, Museum of New Zealand Te Papa Tongarewa, Wellington (formerly National Museum of New Zealand, Dominion Museum, and Colonial Museum), New Zealand; SAM, South Australian Museum, Adelaide, South Australia, Australia; UA, University of Adelaide, Adelaide, South Australia; UNSW, University of New South Wales, Sydney, Australia.anatomical nomenclature: a, acrocoracoid; ac, acromion; artic, articularis; bc, bicipital crest; bf, brachial fossa; bi, brachial impression; cdf, crus dorsale fossa (median crest); cf, clavicle facet; cg, capital groove; chs, coraco-humeral surface; cond, condylus; cort, coracoid tubercle; cr, capital shaft ridge; ct, capital tuberosity; dc, dorsal condyle; dcf, dorsal clavicle facet; dcp, dorsal cotylar process; dcr, deltoid crest; dpf, dorsal pneumotricipital fossa; dt, dorsal tubercle; ecp, ectepicondylar prominence; f, foramina; fal, facet for anterior articular ligament; fau, facies articularis ulnaris; ff, furcular facet; fmbb, fovea for insertion of M. biceps brachii; hf, humeral facet; hh, humeral head; L, left; lf, lateral facies; lig, ligamentum; M, musculus (re muscles); m, musculi (as part osteological name); mf, medial facies; pf, procoracoidal foramen; pnf, pneumatic foramen; proc, processus; R, right; rg, radial groove; sa, supraspinatus attachment scar; sc, scapular cotyla; tc, carpal tuberculum; tg, scapulotricipital groove; tuber, tuberculum; vc, ventral condyle; vcu, ventral cotyla of the ulna; vcl, ventral collateral ligament tuberosity; vpf, ventral pneumotricipital fossa.

rESulTS

Systematic palaeontology

OXYURINAE Phillips, 1926: Stiff-tailed ducksgenus: Manuherikia Worthy, Tennyson, Jones, McNamara & Douglas, 2007type species: Manuherikia lacustrina Worthy, Tennyson, Jones, McNamara & Douglas, 2007.The new taxon described below is referred to Manuherikia as it shares the following unique combination of humeral characters: 1, capital shaft ridge prominent, extends towards tuber.


dorsalis; 2, fossa pneumotricipitalis ventralis not pneumatic (closed); 3, crista deltopectoralis dorsally concave; 4, dorsal tubercle elevated; 5, dorsal pneumotricipital fossa broad, flattened, overhung by caput humeri; 6, crus dorsale fossa (median crest) large, prominent; 7, scar for attachment M. scapulohumeralis cranialis (supraspinatus) elongate; 8, fossa m. brachialis deep, bound medially by sharp ridge; 9, tuber. supracondylare ventralis (facet for anterior articular ligament) elevated, tilted distally; 10, attachment of M. pronator superficialis (= attachment of pronator brevis) on ventral facies level with top of facet for anterior articular ligament; 11, space between facet for anterior articular ligament and cond. dorsalis wider than facet; 12, epicondylus dorsalis forms distinct flange dorsad of dorsal condyle, no distinct proc. supracondylaris dorsalis.Character 1 is an apomorphy for the clade of anatids more derived than anserines and dendrocygnines. Character 2 is considered a synapomorphy of oxyurines more derived than the living Australian freckled duck Stictonetta.Manuherikia was referred to Dendrocheninae Livezey and Martin, 1988 by Worthy et al. (2007), but transferred to Oxyurinae following phylogenetic analyses and the synonymy of Dendrocheninae with that subfamily (Worthy & Lee 2008), and as supported by more extensive analyses (Worthy unpubl. data). Oxyurines are phylogenetically placed above Dendrocygna and below tadornines in those analyses.Manuherikia douglasi sp. nov. Fig. 1A–Eholotype: NMNZ S.42878, a complete R humerus, missing tuber. ventrale and part ventral margin crista bicipitalis, with repaired fracture in shaft distal to end of deltoid crest. Collected 20 February 2004 by THW, AJDT, Jamie Wood and Dave Mathews.type locality: Croc Site, Layer 1, in 3 m cliff, north side of small hill, left side Mata Creek, Otago, New Zealand (44°53′22″S; 169°50′16″E). Layer 1 is a c. 10 cm thick silty-sand layer c. 3.5 m above the Dunstan/Bannockburn contact, FRN is H41/f84.stratigraphy/age/fauna: Bannockburn Formation, Manuherikia Group, Early Miocene (Altonian); 19–16 Ma; St Bathans Fauna.measurements of holotype: Total length 81.7 mm, length bicipital crest from head 16.0 mm, width from ventral side of head at capital incision to dorsal side of dorsal tubercle 13.1 mm, length deltoid crest from dorsal tubercle 22.8 mm, minimum shaft width 4.8 mm, maximum distal width 10.7 mm, depth dorsal condyle 6.2 mm.etymology: For Barry Douglas who has worked for many years on the Manuherikia Group sediments and was first to find fossil bird (duck) bones in them (Douglas et al. 1981).diagnosis: Humerus larger than Manuherikia lacustrina; proximal end tilted cranially such that plane of dorsal pneumotricipital fossa about 30–40 degrees inclined from plane of shaft and more cranio-caudally compressed; incisura capitis forms shallow (not marked) notch in proximal profile; dorsal tubercle as wide as long; deltoid crest more elongate with 44% of length distad of bicipital crest; ventral pneumatic fossa closed level with the median crest—it does not extend as a deep pocket dorsad of the median crest towards the head; dorsal pneumotricipital fossa relatively wider, not excavated as a pocket under the dorsal part of the head and the margin of the dorsal tubercle; brachial fossa without secondary distoventral deepening; epicondylus dorsalis more prominent dorsally.referred material: Locality information for sites HH1a (FRN H41/f88) and HH1b (FRN H41/f90), Manuherikia River are given in Worthy et al. (2007).Ulnae: NMNZ S.50019, site HH1b, Manuherikia River, R ulna, preserves complete length with slight wear to olecranon, proc. cotylaris dorsalis, and to the ventrocranial prominence


Fig. 1 A–E, Manuherikia douglasi, holotype R humerus NMNZ S.42878 in cranial (A,E), dorsal (B), caudal (C) and proximo-caudal (D), views; F, G, referred L ulna NMNZ S.50018 in medial view (F) and ventral view (G); H, referred R ulna NMNZ S.50019 in ventral view. Abbreviations as in methods. Scale bar is 50 mm for A–C, F–H; and 25 mm for images D, E, and I.


of the cotylaris ventralis, collected 8 January 2007. TL = 78.1 mm, PW = 6.4 mm, SW = 4.1 mm, maximum DW = 8.0 mm (proximal and distal widths are under-estimates due to wear). NMNZ S.50018, site HH1a, Manuherikia River, L ulna, preserves complete length with slight wear proximally and distally, collected 8 January 2007. TL = 73.1 mm, PW = 6.7 mm, SW = 4.0 mm, maximum DW = 8.3 mm. NMNZ S.42953, Layer 1, Croc Site, Mata Creek, proximal L ulna preserving about half of total length, collected 19 February 2004, PW = 7.1 mm. NMNZ S.43945, site HH1a, Manuherikia River, proximal R ulna preserving most of the brachial impression, collected 26 February 2005, PW = 6.8 mm.remarks: Humeri. Of other anatids in the St Bathans Fauna, humeri of Manuherikia lacustrina are most similar to that of the new species. In the holotype of M. douglasi (NMNZ S.42878), the proximal end is more compressed cranio-caudally than in M. lacustrina, due to a less inflated intumescentia humeri. While both taxa have a distally narrowing shaft, that of M. douglasi is more slender. The dorsal pneumatic fossa in M. douglasi, at c. 7 mm wide and nearly twice the length of the capital groove (3.8 mm), is relatively wider than in M. lacustrina. It extends from the dorsal tubercle ventrally well past mid-width of the head, to terminate in a low ridge at the beginning of the capital groove, just above the median crest. It is not excavated under the dorsal part of the head or the margin, of the dorsal tubercle. The deltoid crest is directed partly dorsally and has an even curve to its margin with its greatest elevation above the cranial surface distad of the bicipital crest. The scar for M. latissimus dorsi caudalis (as distinct from M. latissimus dorsi cranialis) commences well proximal to the end of the deltoid crest and links to the distal end of that crest before extending further distally. The caudal facies adjacent to the scar for the supraspinatus is markedly concave, so the bone at this point is cranio-caudally thin. Distally the sulcus scapulotricipitalis is shallow caudally and barely extends onto the distal end, but wear to the lateral and medial prominences defining the groove has obscured its former extent.Manuherikia lacustrina is smaller—humerus length ranges 70.3–78.4 mm, mean 75.0 mm, n = 5 (Worthy & Lee 2008) and differs from M. douglasi as follows: ventral pneumotricipital fossa is deeply excavated as a pocket past the median crest towards the head; dorsal pneumotricipital fossa has a distinctive rounded margin in caudal view, usually excavated under dorsal part of head and margin of dorsal tubercle (only in rare examples is it not, e.g., NMNZ S.50017), plane of fossa base more in line with shaft (not caudally inclined), fossa relatively narrow (length capital groove 80–100% width of dorsal fossa), ventral margin of fossa at about mid-head width and excavated markedly below the capital groove; capital incision forms distinct notch in proximal profile; dorsal tubercle elongate; deltoid crest shorter, with less (c. 40%) of length distad of bicipital crest, profile angular, most elevated above the cranial facies opposite maximum ventral extent of bicipital crest; scar for M. latissimus dorsi caudalis commences level with or slightly above distal end of deltoid crest; caudal facies adjacent to the scar for the supraspinatus is not markedly concave; flange of dorsal epicondyle less prominent such that profile proximad of it is convex rather than concave; brachial fossa with distinct secondary deepening distoventrally; scapulotricipital sulcus is present caudally and extends around distal end. Other oxyurines in the St Bathans fauna include Manuherikia minuta, which is considerably smaller than the new species and Dunstanetta johnstoneorum, which is substantially larger (Worthy et al. 2007; Worthy & Lee 2008). Humeri of Recent taxa included within Oxyurinae, sensu Worthy & Lee (2008), differ as follows: Stictonetta has a pneumatic ventral pneumotricipital fossa. Malacorhynchus, although differing markedly in the pelvic limb (lacks diving apomorphies), is most similar to Manuherikia in humeral morphology, but is distinguished by a shorter deltoid crest, a less


well developed flange to the dorsal epicondyle, and a more elongate flexor process. Oxyura differs markedly with a much broader dorsal pneumotricipital fossa and the attachment of the pronator brevis is merged with the cranial edge of the facet for the attachment of the anterior articular ligament. Biziura and Thalassornis humeri are much more elongate, with the scapulotricipital groove present only caudally and then weakly defined. In both these taxa and Oxyura, the facet for the attachment of the anterior articular ligament is depressed and the pit for the attachment of the pronator brevis is fused into its cranial margin. In Thalassornis, unlike all other oxyurines, the gap between the facet for the attachment of the anterior articular ligament and the dorsal condyle is narrower than the facet width. Among fossil oxyurines, the Oligo-Miocene Mionetta of Europe has a similarly short dorsal tubercle, but the shaft is narrowest at mid length, the brachial depression is deeper and separated from the medial margin by a broader ridge, and the facet for the anterior articular ligament relatively is smaller. In the Australian Oligo-Miocene undescribed oxyurines, the dorsal tubercle is also about as wide as long, but the capital groove forms a shallower notch in the proximal profile, and distally it has a distinct ectepicondylar prominence (THW unpubl. data). Ulnae. The two whole referred ulnae, at 73.1 mm and 78.1 mm long and ranging 89–95% the length of the holotype humerus of M. douglasi, are slightly longer than expected, given that the mean ulna length of 62.3 mm, range 56.1–67.4, n = 25 in M. lacustrina (Worthy & Lee 2008) is 83% of mean humeral length. As predicted from the comparatively slender humerus of M. douglasi, these ulnae are very slender, with their width measurements within the ranges for ulnae of M. lacustrina (Worthy & Lee 2008). They differ further as follows (state for M. lacustrina in brackets): fossa brachialis shallow, not recessed caudally (deep, recessed caudally from ventral facies); lig. collateralis ventralis elongate, extending beside half length of brachial fossa (on a prominent oval rugosity at proximal end of brachial fossa); proc. cotylaris dorsalis distally wraps down onto shaft with no foramen under it (usually with foramen below it); incisura radialis shallow, in unworn specimens contains two distinct articular facets, a large distal one extending from below the dorsal cotyla towards a shorter second one positioned under the lip of the ventral cotyla (radial groove deep, distal facet relatively smaller than proximal one); tuber. carpale robust, distal margin further separated from cond. ventralis ulnaris than in M. lacustrina; maximum distal width is about twice maximum depth a result of a wider angle between the plane of the dorsal condyle and the dorsal facies (width and depth not so disparate, condyle close to right angle with dorsal facies). Ulnae of other waterfowl in the in the St Bathans fauna differ as follows: those of Manuherikia minuta are much smaller, and those of Dunstanetta and Miotadorna are much larger. That of Matanas is so far unknown but is expected to be comparatively more robust given the humerus morphology.

Anserinae, genus and species indeterminate ARadius (Fig. 2A–E)specimen: NMNZ S.50020 a proximal L radius. Maximum proximal width 7.8 mm, ventrodorsal depth 7.4 mm, lateromedial depth 7.3 mm, preserved length 87 mm.location: Vinegar Hill, Site 1, 44°52.627′S; 169°44.958′E, on most northwestern exposure in valley, near St Bathans, Central Otago. Collected 11 January 2007 by the UA, UNSW, NMNZ, CM team.horizon: A layer 1–5 cm thick of unconsolidated clean white sand with small white quartz pebbles, taken to equate to that shown at 55.5 m on the section in Douglas (1986), Vinegar Hill, St Bathans, Otago. In the first 0.5 m of overlying green-grey and yellow-green sandy clays, occasional bird bones were found in 2007—including at least three bones of Manuherikia


lacustrina (humerus, coracoid, sternum). A sphenodontid fragment (NMNZ S.42282) was found in the sand layer in 2001, and fish bones (eleotrid) are common.description and comparisons: The radius has the typical form of anatids: the cotyla humeralis on the caput radii is as wide as deep and the shaft leading to the proximal end is straight, without a kink. The insertion point for the M. biceps brachii does not form a distinct tuber. bicipitale radialis (bicipital tubercle) found in many birds, rather it is in a relatively proximal location and has the form of an elongate shallow fovea on the ventral facies bound laterally by a narrow ridge on its interosseus margin, as is typical of anatids. The capital tuberosity is bulbous over the dorsal facies, wider (5.3 mm) than long (4.4 mm), with its prominence aligned diagonally from the medial cotylar margin distally towards the lateral or interosseus margin. The facies artic. ulnaris extends from the dorsal facies around onto the lateral facies (interosseus margin), is about 3 mm long, and occupies about 4 mm of the 7 mm width of the lateral facies: the ventral half of the lateral facies extends uninterrupted from the cotyla down the shaft. The capital tuberosity forms a distinct bulge from the dorsomedial margin of the cotyla in proximal aspect. A prominent linea intermuscularis extends along the dorsal margin of lateral (interosseus) facies, with the interosseus facies flattened. The shaft is dorsoventrally deeper than the lateromedial width (5.5 by 3.8 mm at 35 mm from proximal end). Intermuscular lines on the preserved portion suggest that the preserved length is slightly more than half of the complete bone, assuming its proportions are similar to Cereopsis, so the expected total length is perhaps 150 mm. It is much larger than tadornines (e.g., Tadorna tadornoides and Miotadorna) and Anatines. Tadornines differ further with the capital tuberosity being about as long as wide and the ulna facet being a clearly defined rugose area about as long as wide and occupying the whole width of the interosseus facies. The fossil is the size of radii of Anseranas (the magpie goose, Anseranatidae) and anserines. Anseranas (e.g., SAM B.48035) is distinguished from the fossil by the insertion point for the M. biceps brachii forming a distinct slightly elevated facet rather than a fovea, the capital tuberosity being more massive, and the ulnar facet extending over a greater width of the lateral/interosseus facies and being more flattened. Cygnus (swans) are similar to other anatids, with the insertion point for the M. biceps brachii forming a shallow fossa, but differ in that the ulnar facet extends over most of the lateral/interosseus width along the cotylar margin but has little length down the shaft. The geese (Anser brachyrhynchus, A. caerulescens, Branta canadensis) differ markedly with the ulnar facet being crescent-shaped, most massive dorsally but extending over the whole lateral/interosseus width, and about twice as wide as long. While the ulnar facet is short in these geese, the interosseus facies distad of it is slightly elevated and rugose down the shaft to the distal end of the insertion point for the M. biceps brachii. This rugose area is absent in the fossil. Also, in Anser and Branta, the capital tuberosity is more bulbous, about as long as wide, and its lateral margin is not bound by a diagonal groove as in the fossil. In Cereopsis (e.g., SAM B.39638), the structure of the proximal radius is essentially the same as the fossil: the capital tuberosity is wider than long and diagonally oriented, the ulnar facet wraps around the dorsal-lateral angle of the head and is restricted to the dorsal half of the lateral or interosseus facies, the interosseus facies is smooth over its ventral half distad of the cotyla, i.e., it lacks any area of rugosity as seen in other anserines. The fossil differs from Cereopsis however, by its slightly smaller size, and that the shaft is relatively deeper compared to width. In conclusion, the fossil is clearly an anserine, and is more similar to Cereopsis than other anserines. Comparison of the fossil with the radius of Cnemiornis is not very meaningful, as in the latter, the wing skeleton is very reduced and associated structure of joints are much


modified by loss of discrimination of form for the various features. The radius of Cnemiornis is also very much shortened like the rest of its wing skeleton compared to the fossil.

Coracoids (Fig. 2H,I)specimens: Two cranial fragments of L coracoids: NMNZ S.50823, length of humeral facet 16.7 mm, width humeral facet 10.9 mm. NMNZ S.50824, length of humeral facet 16.2 mm, width humeral facet 10.8 mm.location: Site HH1a, Manuherikia River (FRN H41/f88) in Trench excavations. Both collected 14 January 2008 by the UA, UNSW, NMNZ, CM team.description and comparisons: Both fragments preserve the facies artic. humeralis and a few millimetres of the lateral facies distad of it, the cotyla scapularis, the dorsal part of the sulcus m. supracoracoidei, and the neck and coraco-humeral surface (sensu Howard 1929) leading cranially from the humeral facet. They lack the proc. acrocoracoideus, the incisura nervi supracoracoidei (procoracoidal foramen) and the entire ventral side opposite the above preserved features. Despite their fragmentary nature, these fragments only approach anseriforms in the combination of a deep cup-like scapular cotyla, the cranial margin of the supracoracoidal sulcus being directed at about 45 degrees to the plane of the adjacent humeral facet, and the shape of the humeral facet which overlaps the ventral facies as a distinct flange. Among Ciconiiformes, Platalea and Threskiornis (Threskiornithidae) are the most similar non-anatids, but differ with a smaller, shallower and less round scapular cotyla. In addition, in Platalea the supracoracoidal sulcus is excavated adjacent to the humeral facet and the neck is more compressed. The fossils differ markedly from coracoids of tadornines including the sympatric Miotadorna, in which the largest individuals are only slightly smaller, by the humeral facet being relatively broader, the neck and its coraco-humeral surface being relatively and absolutely thicker, and the lateral facies below the humeral facet being planar not convex. These features are all shared with anserines such as Cereopsis, Anser and Cygnus. NMNZ S.50824 preserves the dorsal part of the facies artic. clavicularis overhanging the supracoracoidal sulcus, and this part of the sulcus has no pneumatic foramina in it, unlike the condition in Cereopsis, Anser, Cygnus and Branta. However, pneumaticity of the rest of the acrocoracoid in the fossils cannot be determined in these fragments. In Cnemiornis, pneumatic foramina are variably present entering the acrocoracoid from below the clavicle facet (Worthy et al. 1997). In summary, these two fossils can be referred to an anserine, but are insufficient to determine relationships further. They are however, of expected size for the taxon that the radius described above and for which the scapula described next relates to.

Scapula (Fig. 2L)

specimen: NMNZ S.50822, a R scapula missing most of the blade. Maximum proximal width from humeral facet to acromion 15.8 mm, length humeral facet 10.8 mm, width humeral facet 6.5 mm, shaft depth 6.8 mm, and shaft thickness 4.3 mm.location: Site HH4, Manuherikia River section, 44°54.472′S; 169°51.434′E, FRN H41/f0095, collected on 8 January 2008 by the UA, UNSW, NMNZ, CM team.horizon: A 5–15 cm thick clay-silt layer notable for an abundance of shell fragments of the gastropod Glyptophysa, first exposed between February 2005 and March 2006, 25.63–25.83 m above the base of the Bannockburn Formation.description and comparisons: NMNZ S.50822 is similar to anseriforms in general shape, including a near planar medial surface, a dorsally prominent acromion, a rounded tuber. coracoideum on the cranial margin, and a large elongate humeral facet. It shares with true


geese within anserines (i.e., not swans) a large pneumatic foramen above the humeral facet on the lateral facies. Such a foramen is found in anhimids and Anseranas, but is rarely found outside of anserines in anatids, e.g., in Somateria and Cairina, which are large taxa with highly pneumatic skeletons. NMNZ S.50822 differs from similar-sized scapulae of Anseranas as follows: the shaft is more nearly planar with the humeral facet (in Anseranas, shaft is rotated markedly medial of the plane of the humeral facet); the coracoid tubercle is more prominent; the dorsal margin of the collum (shaft) is more rounded (sharp crest in Anseranas); and the acromion is thinner. It differs from all anatids except Cereopsis and Cnemiornis in that the acromion is very reduced, to the extent that the coracoid tubercle projects craniad of it. The orientation of the shaft relative to the humeral facet is similar in these taxa to the fossil. The fossil however, differs from Cereopsis in that the acromion is relatively less cranially prominent and that the furcular facet is comparatively reduced in size. The scapulae of Cnemiornis are, as for the coracoids, very reduced, concordant with the reduced wings of this flightless taxon, but like the fossil the coracoid tubercle is cranially protuberant of the acromion, the furcular facet is reduced and there is a pneumatic foramen. NMNZ S.50822 is therefore of an anserine more similar to Cereopsis and Cnemiornis than other compared taxa. It is of appropriate size for the coracoid fragments (NMNZ S.50823, S.50824), and too large for NMNZ S.42797 (see below).

Anserinae, genus and species indeterminate B

Coracoids (Fig. 2F,J)Worthy et al. (2007) described a large anatid coracoid (NMNZ S.42797; here Fig. 2F) from Site HH1a, Manuherikia River, from the St Bathans Fauna and noted some similarities with anserines. A second specimen of this taxon was found in HH1a Trench excavations in January 2008.location: Site HH1a, Manuherikia River (FRN H41/f88), main quarry (NMNZ S.42797) and in the equivalent bed in the Trench excavations (NMNZ S.50825), about 40 m east of the main quarry. Collected 15 January 2008 by the UA, UNSW, NMNZ, CM team.description and comparisons: The new specimen NMNZ S.50825 is smaller (estimated medial length 49 mm) than NMNZ S.42797 (57.5 mm), but within the range of size variation seen in modern sexually dimorphic taxa, and is otherwise similar in form. It is less perfect, lacking the acrocoracoid, and the anguli medialis et lateralis (Fig. 2J). Like all anatids, they have muscular striae diagonally traversing the dorsal surface of the sternal half of the element; they lack a distinct impressio m. sternocoracoidei; the facies artic. sternalis is broad with only a narrow ridge bordering it cranially, unlike the robust overhanging lip in Anseranas; the acrocoracoid has about equal width in its dorsal and ventral parts. Their proportions, particularly a relatively large cranial end, are similar to anserines e.g., Anser caerulescens. They differ from all anserines in having a small procoracoidal foramen that does not penetrate the corpus. In the presence of a procoracoidal foramen and their general shape these bones are similar to coracoids of Threskiornithidae, particularly those of the Australian White Ibis Threskiornis molucca and

Fig. 2 Anserine fossils from the St Bathans Fauna compared to Recent taxa. Anserine gen. et. sp. indeterminate A: proximal L radius NMNZ S.50020, A–E; (A), lateral/interosseus facies; (B), ventral facies; (C), dorsal facies; (D), dorso-lateral facies; (E), lateral/interosseus facies. Anserine gen. et. sp. indet. A, cranial part L coracoids in latero-dorsal view: H, NMNZ S.50824; I, NMNZ S.50823; R scapula in lateral view, l, NMNZ S.50822. Anserine gen. et. sp. indet. B, L coracoids in dorsal view: F, NMNZ S.42797; J, NMNZ S.50825. Cereopsis novaehollandiae: G, L coracoid, SAM B.39638; K, R scapula, SAM B.32830. Cnemiornis gracilis: M, L scapula, CM Av25001. Abbreviations as in Methods. All scale bars = 20 mm; (D) and (E) magnified 200% of (A–C).


Straw-necked Ibis Threskiornis spinicollis and Platalea spp. (spoonbills). However, coracoids of spoonbills and ibises differ markedly from them with the sternal blade having a distinct sterno-coracoidal impression, muscular striae restricted to this impression, an acrocoracoid that is markedly wider ventrally than dorsally, a wider procoracoid such that the procoracoidal foramen exits at mid-width on the dorsal facies, and a longer procoracoid. All anserines, including Cereopsis and Cnemiornis considered to be basal in this subfamily (Worthy & Lee 2008), typically have an enlarged cranial end compared to anatines, and a highly pneumatic acrocoracoid. In NMNZ S.42797 (not determinable in S.50825), the foramina opening from the supracoracoidal sulcus into the acrocoracoid are much smaller than in Cereopsis and other Recent anserines—this feature was interpreted as not supporting a close relationship with anserines (Worthy et al. 2007). However, these coracoids are more slender than Cereopsis, for example, while the medial length of NMNZ S.42797 (57.5 mm) is similar to the coracoid of Cereopsis SAM B.39638 (59.9 mm), the shaft is much more gracile (5.7 × 5.4 mm versus 9.6 mm [12.9 mm including foramen] × 9.2 mm). In accordance with their more gracile form, these coracoids have a relatively narrower humeral facet, e.g., in NMNZ S.42797 it is 14.4 mm long by 8 mm wide, ratio of 0.555. This is similar to those of the sympatric Miotadorna (mean L = 12.6, W = 6.9, ratio = 0.554; n = 9, range 0.521–0581). A previously unmentioned feature of NMNZ S.42797 is a prominent flange extending up the medial margin from the medial angle ending on the dorsal side of the rounded facies of the shaft. A very similar flange is present in Cereopsis and in Cnemiornis but is absent in Branta and Anser species. Also, the clavicle facet in this fossil cleanly overhangs the supracoracoidal sulcus for its whole depth, as it does in Cereopsis. Homology of this feature in Cnemiornis is difficult to discern due to severe reduction of the coracoid associated with flightlessness. In Branta and Anser, both with well developed acrocoracoids, the sulcus is either not overhung by the clavicle facet (Branta), or only overhung by the dorsal part of the facet (Anser).

DIScuSSIoN

Increased diversity of ducks in the St Bathans FaunaThe description of a third species of Manuherikia increases the diversity of waterfowl for the Early Miocene St Bathans Fauna (19–16 Ma) to six named species. Already the most diverse pre-Pliocene waterfowl fauna known (Worthy et al. 2007), this new species reinforces the global significance of this fauna, and extends the diversity of the Oligo-Miocene global radiation of the “stiff-tailed ducks” (Oxyurinae). Oxyurines associate as either a phylogenetic grade of taxa, or a single clade with the New Zealand fossils Manuherikia and Dunstanetta, and in which clade the European Mionetta is relatively basal (Worthy & Lee 2008; Worthy “Descriptions and phylogenetic relationships of two new genera and four new species of Oligo-Miocene waterfowl (Aves: Anatidae) from Australia” in prep. The extant monotypic Australasian genera Stictonetta, Biziura, and Malacorhynchus, and the cosmopolitan Oxyura, are descendents of that radiation that have survived to the present day. The three fossil species now described for Manuherikia differed in size, from M. minuta, one of the smallest ducks known, to M. douglasi, about the size of the extant Australasian shoveler Anas rhynchotis. M. lacustrina was of intermediate size, but closer to M. douglasi. The wing bones of M. douglasi were more modified for diving, with the humerus compressed and the dorsal pneumotricipital fossa enlarged more towards the state seen in the highly specialised diving ducks Oxyura and Biziura, than in its smaller congeners. As yet leg bones of M. minuta are still very incompletely known so the extent of its diving capability is not known. Humeri of M. minuta are, however, relatively slightly more robust than its larger congeners, but so are those of the small Australian pygmy goose (Nettapus), which is also a diver. This suggests


that there may be some tradeoffs between constraints for flight versus diving in anatids when absolute size is small. The available bill fragments, e.g., NMNZ S.42783, 43181, suggest that Manuherikia had a typical broad flat bill, unlike for example Mergus, and the os dentale of the mandible, e.g., NMNZ S.43091, 43183, 44087, has a series of parallel grooves and ridges such as is seen below the bill lamellae in modern anatines. These features and its diving adaptations suggest that Manuherikia likely foraged on invertebrates in lake bed sediments and/or among aquatic plants. In the St Bathans Fauna, Matanas enrighti is roughly of similar size to Manuherikia doug-lasi, but is more robust, with proportions of the anatine dabbling ducks, with which it shares a pneumatic ventral pneumotricipital fossa in the humerus. Matanas enrighti is relatively rare and its leg bones have not been identified, but perhaps it occupied a dabbling niche. In com-parison, the larger fossil taxon, Dunstanetta johnstoneorum, was a highly specialised diver, as much or more so than the Eurasian scoter (Melanitta nigra), with which it shared many derived diving characteristics. Other than these “ducks”, the St Bathans Fauna included a shelduck (Miotadorna sanctibathansi), which was slightly bigger than the extant New Zealand paradise shelduck (Tadorna variegata), but relatively longer legged, and was presumably a lakeshore browser.

Anserines in the St Bathans FaunaFossils described above reveal the presence of two species of anserines in the St Bathans Fauna, making a total of eight waterfowl taxa. A species of an anserine, more similar to Cereopsis and Cnemiornis than other anserines, is represented by two coracoid fragments, a scapula and half a radius. This goose may be ancestral to Cnemiornis, if not to both Cnemiornis and Cereopsis. These are inferred to be sister taxa (Worthy et al. 1997) or at least together these are successive taxa in a basal grade of anserines (Worthy & Lee 2008). Cnemiornis, being very large and quite flightless, had a highly modified skeleton with degenerate pectoral elements and greatly enlarged pelvic girdle elements, which suggest a long history in New Zealand. The present fossil record of Cnemiornis extends back only into the Mid-Pleistocene, to Oxygen Isotope Stage 17, c. 680, 000 years BP (Gill et al. 2005). That Mid-Pleistocene fossil is, however, indistinguishable from the Late Holocene form and therefore provides some evidence for a much longer evolutionary history for the taxon in New Zealand. The fossil radius from the St Bathans Fauna, in contrast to Cnemiornis from the Quaternary, is from a goose that had the ability to fly well and that was only slightly smaller than Cereopsis. Two coracoids represent a smaller species of anatid that has proportions approaching those of anserines but which differs from Cereopsis mainly by smaller size, a smaller more typical procoracoidal foramen robustly enclosed medially, rather than a large elongate foramen bound medially by a thin bone strip, and lesser pneumaticity of the acrocoracoid. The latter may be explained as a functional correlate perhaps related to less need for weight loss in this smaller-sized bird. The presence of a procoracoidal foramen is a plesiomorphic feature, present in the anseriform taxa Anhimidae, Anseranatidae, and Presbyornithidae, and within Anatidae it occurs in Cnemiornis and occasionally in Cereopsis. This feature in the fossil is restricted to the procoracoid, i.e., it does not penetrate the shaft as it does in Anseranas or Anhima. Its presence in an anserine in the Middle Miocene is therefore not unexpected and might be expected in the ancestor of either Cnemiornis and/or Cereopsis. The differences may not be phylogenetically significant and these fossils are here interpreted as a probable anserine more closely related to Cereopsis than Branta, Anser or Cygnus. Determination of these fossils as anserines sheds light on the evolution of the group in Australasia. The only anserines within the modern or Recent Australasian indigenous water-


fowl are Cereopsis and the black swan (Cygnus atratus). Cygnus is not known earlier than the Pliocene in Australia (Rich et al. 1991; Vickers-Rich 1991; T. H. Worthy unpubl. data), and the oldest record of Cereopsis is from the Pliocene in the Bluff Downs Fauna (Boles & Mackness 1994). There are no earlier anserine fossils known from Australia (T. H. Worthy unpubl. data) by which the evolutionary history of anserines in Australia may be determined. However, a Cereopsis-Cnemiornis ancestor in the Early Miocene of New Zealand makes it probable that a member of this clade will be found in the Oligo-Miocene of Australia—at this time both land masses also shared tadornines and oxyurines (T. H. Worthy unpubl. data). Cereopsines now have a demonstrated history in Australasia since the Early Miocene, but there is no evidence of other anserines in the region, such as Northern Hemisphere Anser, Branta and relatives. The earliest definitive fossil record of anserines worldwide is from the mid-Oligocene of Europe, with Cygnopterus affinis (Van Beneden, 1883) from Belgium (Lambrecht 1933; Brodkorb 1964; Howard 1964; Olson 1985; Louchart et al. 2005). Less well known are two possible swans, Guguschia nailiae Aslanova & Burczak-Abramovich, 1968 and Cygnavus formosus Kurochkin, 1968 from the European Oligocene (Aslanova & Burczak-Abramovich 1968; Kurochkin 1968), but see Louchart et al. (2005) for observations of non-cygnine features of Guguschia. By the Early Miocene, anserines are well known in European fossil sites, with Cygnopterus alphonsi Cheneval, 1984, Cygnavus senckenbergi Lambrecht, 1931, and Anserobranta? (Anas) robusta (Milne-Edward, 1867–71) (Lambrecht 1933; Kurochkin 1968; Cheneval 1987; Mlíkovský 2002; Louchart et al. 2005; Worthy et al. 2007). True swans (Cygnus), first appeared in the fossil record in the Mid-Miocene of Europe, with Cygnus atavus (Fraas, 1870) (Mlíkovský 2002). Swans were first reported from the Miocene of North America by Wetmore (1943), but the first named species appears in the Late Miocene (Bickart 1990). Recently Louchart et al. (2005) described a new genus (Afrocygnus) in Cygnini from the Late Miocene (c. 7 Ma) of Chad in Africa. Likewise, true geese (Anser and Branta) first appear in Europe in the Middle-Late Miocene (Howard 1964; Burczak-Abramovich & Gadzyev, 1978; Heizmann & Hesse 1995; Boev 2002; Mlíkovský 2002; Boev & Koufos 2006). In North America, anserines are first evidenced by Presbychen abavus Wetmore, 1930 from the Middle Miocene of California (Brodkorb 1964; Howard 1964), and both Anser and Branta first appear in the Middle-Late Miocene (Miller 1961; Bickart 1990). Cereopsine fossils have not been recovered outside Australasia. These observations show that the anserine faunas of Australasia and the Northern Hemisphere have evolved separately, at least since the Early Miocene to the Late Pliocene, when Cygnus entered the Australian fauna. Australia split from Antarctica about 35 Ma and thereafter was relatively isolated as it was carried on the Sahul Plate northwards towards Asia (e.g., Brown et al. 2006). As the continent moved north there was increasing likelihood of colonisation from the north, but major biotic interchange first appears in the latest Miocene—earliest Pliocene, with first appearances of colubrid snakes (Scanlon 2006) and rodents (Aplin 2006), and a major turnover in bat communities (Hand 2006). This then is the time that we might expect new avian groups from the north in Australasia, and among waterfowl Cygnus seems to be one such taxon. The goose fossil from the St Bathans Fauna illustrates that the cereopsine group has a much more ancient history in Australasia and is not a relatively recent immigrant. The distinctiveness of the Australasian cereopsine geese is thus explained as the product of a separate evolutionary lineage evolved in isolation probably since at least the Oligo-Miocene. These observations are supported by the novel hypothesis of a sister relationship of Cereopsis to the southern South American swan Coscoroba, which was found to be basal to swans and outside of other anserines by Donne-Goussé et al. (2002), based on mitochondrial DNA analysis. Such a relationship is also confirmed by the discovery of a nuclear gene apomorphy,


namely the shared presence of “chicken repeat 1”, a non-long terminal repeat retrotransposition in the third intron of the lactate dehydrogenase B gene, in Coscoroba and Cereopsis and not in other anserines nor anatines (St. John et al. 2005). Together these fossil and genetic data indicate a southern radiation of anserines. However, St. John et al. (2005) estimated the time that the Coscoroba/Cereopsis clade diverged from other anserines as 10.5 million years ago, based on genetic divergence. This young age is not supported by the above fossil data, nor is it supported by the presence of a cereopsine, discrete from other anserines, in the St Bathans fauna 19–16 million years ago. For these reasons, when better material of the New Zealand cereopsine/s or an Australian equivalent becomes available, it has the potential through phylogenetic analyses to resolve the relationships of Cereopsis, Coscoroba and the European Oligocene to earliest Miocene anserines. Present phylogenetic analyses, e.g., Livezey (1986, 1989, 1996, 1997) and Worthy & Lee (2008) support the basal position of Cereopsis and Cnemiornis relative to other anserines. This together with their geographic distribution might suggest a southern origin for anserines, but caution is required in accepting this interpretation as discordance between modern and fossil distributions are becoming increasing apparent, e.g., Anseranatidae, Recent in Australia, is known from the Palaeogene (Eocene and Oligocene) of Europe (Olson 1999; Mourer-Chauviré et al. 2004). For now we consider it is best to interpret the data thus: that anserines evolved by at least the Oligocene and, at least since the earliest Miocene until the Pliocene, had a separate evolutionary history in Australasia from that of anserines in the Northern Hemisphere. Lastly, these anserines place a probable ancestor of the Cnemiornis clade in New Zealand 19–16 Ma, indicating a probable dispersal earlier than this time from Australia. Thus one of the highly characteristic New Zealand taxa was emplaced in New Zealand by this time along with other taxa typical of the Recent fauna such as flightless ratites presumed to be moa ancestors, owlet nightjars closer to New Zealand taxa than any others, several parrots, honeyeaters, an endemic terrestrial land mammal, mystaciniid bats, tuatara, and skinks and geckoes (Tennyson et al. 2003; Worthy et al. 2002, 2003, 2006, 2007). Thus, if the New Zealand fauna was going to be developed by spasmodic dispersal, as advocated by, for example, Landis et al. (2008), then most of the characteristic taxa would need to have arrived in the first couple of million years, with no unique taxa arriving thereafter. However, an increasing array of genetic studies show New Zealand taxa to have diverged from sister taxa elsewhere long before the Oligocene highstand, e.g., the wattlebirds and stitchbirds (Driskell et al. 2007; Shepherd & Lambert 2007), kakapo and kaka (Tavares et al. 2006), freshwater crayfish (Apte et al. 2007), hyriid molluscs (Graf & Foighil 2000), and kauri (Knapp et al. 2007). Given these data and the now growing body of fossil evidence for the presence of uniquely New Zealand taxa in the Early Miocene, and unique radiations in ducks, e.g., Manuherikia and associated oxyurine genera not closely related to any contemporary Australian taxa (Worthy & Lee 2008; unpubl. data), it appears that the biota, at least in part, has a history in New Zealand longer than the last 22 Ma. For some, such as the tuatara, leiopelmatid frogs and the terrestrial land mammal, an origin as Gondwanan vicariant taxa is most probable.

AcKNoWlEDGMENTSWe thank many field assistants in the St Bathans Fauna project, especially notably Mike Archer, Jenny Jones, Dave Mathews, Kristen Ramsdale, Zoe Hill, Jamie Wood, Jacqui Nyugen, Julian Louys, Henk Godthelp, and Phil Creaser and the rest of the Australian field contingents of 2007 and 2008 who helped find key specimens discussed herein. We are particularly thankful for the generous support to this project by the land owners Ann and Euan Johnstone and Jack Enright. This research was funded by the Australian Research Council, and supported by the University of New South Wales and the authors’ institutions.


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A new species of the diving duck Manuherikia and evidence for geese (Aves: Anatidae: Anserinae) in...

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