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An early Miocene diversity of parrots (Aves, Strigopidae, Nestorinae) from New Zealand
Trevor H. Worthy , Alan J. D. Tennyson & R. Paul Scofield
a a b c

School of Biological, Earth and Environmental Sciences, University of New South Wales, New South Wales, 2052, Australia
b c

Museum of New Zealand Te Papa Tongarewa, P.O. Box 467, Wellington, New Zealand Canterbury Museum, Rolleston Ave., Christchurch, 8001, New Zealand

Available online: 09 Sep 2011

To cite this article: Trevor H. Worthy, Alan J. D. Tennyson & R. Paul Scofield (2011): An early Miocene diversity of parrots (Aves, Strigopidae, Nestorinae) from New Zealand, Journal of Vertebrate Paleontology, 31:5, 1102-1116 To link to this article: http://dx.doi.org/10.1080/02724634.2011.595857

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Journal of Vertebrate Paleontology 31(5):11021116, September 2011 2011 by the Society of Vertebrate Paleontology

ARTICLE

AN EARLY MIOCENE DIVERSITY OF PARROTS (AVES, STRIGOPIDAE, NESTORINAE) FROM NEW ZEALAND
TREVOR H. WORTHY,*,1 ALAN J. D. TENNYSON,2 and R. PAUL SCOFIELD3 School of Biological, Earth and Environmental Sciences, University of New South Wales, New South Wales 2052, Australia, t.worthy@unsw.edu.au; 2 Museum of New Zealand Te Papa Tongarewa, P.O. Box 467, Wellington, New Zealand, alant@tepapa.govt.nz; 3 Canterbury Museum, Rolleston Ave., Christchurch 8001, New Zealand, pscoeld@canterburymuseum.com

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ABSTRACTA new genus and three species of parrot (Psittaciformes, Strigopidae, Nestorinae) are described from the early Miocene (1916 Ma) St Bathans Fauna of Otago, New Zealand, based on 85 fossils as follows: Nelepsittacus minimus (17), N. donmertoni (60), and N. daphneleeae (6), with two additional fossils representing a fourth unnamed taxon. These taxa range from small parrots approximately the size of Cyanoramphus species to one as large as the living Nestor notabilis. Apomorphies in the coracoid, humerus, tibiotarsus, and tarsometatarsus ally Nelepsittacus with Nestor and exclude a close relationship with Strigops, the other endemic genus assumed to have had a long history in New Zealand. With only nestorine parrots represented, the St Bathans Fauna has nothing in common with the Australian psittaciform fauna, in which cacatuids and a diversity of psittacid genera exist. These data add to the growing body of evidence that the New Zealand terrestrial vertebrate fauna, at a time minimally 3 Ma after the maximal marine inundation of Zealandia in the late Oligocene, was highly endemic, with no close relationship to the closest faunas in Australia. This high degree of endemism strongly suggests that the Zealandian terrestrial biota persisted, at least in part, through the Oligocene highstand in sea level.

INTRODUCTION The diversity of parrots (Aves, Psittaciformes) in Australia is considerable, with approximately 55 indigenous species (including three recently extinct) in three families and 29 genera (Higgins, 1999; Christidis and Boles, 2008). In contrast, the New Zealand parrot fauna is depauperate, with just nine species in three genera in the whole archipelago (Gill et al., 2010). However, these few taxa include the unique kakapo Strigops habroptilus, which with the genus Nestor, also endemic to the New Zealand biogeographic region (i.e., New Zealand, Norfolk Island, and Chatham Island), forms the sister group to all other psittaciforms globally (de Kloet and de Kloet, 2005; Tavares et al., 2006; Tokita et al., 2007; Wright et al., 2008). The kakapo is the worlds heaviest and only ightless parrot; it is also nocturnal, a polygynous lek breeder, and now critically endangered (Powlesland et al., 2006). Nestor has just three described species, one on each of North, South, and Norfolk islands (Gill et al., 2010); however, an undescribed species formerly also existed on the Chatham Islands (Millener, 1999). The remaining New Zealand taxa all form part of a recent radiation of Cyanoramphus species, which diversied within the last million years (Boon et al., 2001; Kearvell et al., 2003) and are probably an example of a recent arrival from the Pacic. The fossil history of psittaciforms in New Zealand has until this study been restricted to late Pleistocene and Holocene deposits (Worthy and Holdaway, 2002). The upper early Miocene (1916 Ma) Bannockburn Formation, Manuherikia Group, in Central Otago, New Zealand, has recently been shown to contain the diverse St Bathans Fauna (Worthy et al., 2007). The fossil beds are lacustrine and were deposited near a river delta on one side of a paleolake that extended over some 5600 km2 (Douglas, 1986; Pole et al., 2003).

*Corresponding

author.

The avifauna of the St Bathans Fauna is abundant and dominated by waterfowl (Anseriformes), with a minimum of eight taxa in ve genera (Worthy et al., 2007, 2008). The avifauna also comprises such diverse taxa as moas (Dinornithiformes), a tubenose (Procellariiformes), birds of prey (Accipitriformes), rails (Rallidae), an endemic gruiform (Aptornithidae), a gull (Laridae) and other charadriiforms with features of typical waders, a heron (Ardeidae), a palaelodid (Phoenicopteriformes), pigeons (Columbidae), parrots (Psittaciformes), a swiftlet (Apodidae), an owlet-nightjar (Aegothelidae), and passerine birds (Passeriformes) (Scoeld et al., 2010; Tennyson et al., 2010; Worthy et al., 2007, 2009, 2010a, 2010b). Fish, frogs, reptiles, and mammals are also represented (Worthy et al., 2006, 2011; Jones et al., 2009; Lee et al., 2009; Schwarzhans et al., in press). The fossil record of parrots is depauperate globally (Waterhouse, 2006; Mayr, 2009, 2010a). Despite a good record of Eocene stem-group psittaciforms (Mayr and Daniels, 1998; Mayr, 2002, 2007; Waterhouse et al., 2008; Mayr et al., 2010), crown-group psittaciforms are surprisingly rare, and are Miocene or younger in age (Waterhouse, 2006). Exclusive of Pleistocene and younger taxa, these crown-group fossil psittaciforms include ve Miocene taxa from Europe, one Miocene species from North America, and one from the Pliocene of Argentina (Mayr and Gohlich, 2004; Waterhouse, 2006; Mayr, 2010a). Despite a rich Neogene fossil record and a large diversity of Recent parrots, Australia has an extremely sparse fossil record of parrots (Boles, 2006). It is restricted to a premaxilla referred to Cacatua sp. from the early-middle Miocene (Boles, 1993), and several fossils of the extant Melopsittacus undulatus from a Pliocene site (Boles, 1998), both sites in the Riversleigh World Heritage Property in northwest Queensland. The identity and relationships of the psittaciforms from the early Miocene St Bathans Fauna, already noted as including at least three taxa (Worthy et al., 2007), are therefore important in understanding the evolution of parrots in Australasia and

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WORTHY ET AL.MIOCENE PARROTS OF NEW ZEALAND the Pacic. If these parrots were recently derived from PapuaAustralian faunas by dispersal, as required by the hypothesis of complete inundation of Zealandia (the India-sized continental fragment encompassing New Caledonia and New Zealand and its sub-Antarctic outliers: Campbell and Hutching, 2007; Landis et al., 2008), various Australian genera might be expected in the New Zealand early Miocene fauna, such as the widely dispersed cacatuids (Forshaw, 1989). The presence of endemic New Zealand taxa or stem-group taxa for the Australian radiation in the early Miocene would support either a vicariant origin for nestorine and strigopine parrots in New Zealand, as suggested by, for example, Oliver (1930, 1955) and Wright et al. (2008) and dating perhaps as recently as ca. 55 million years ago (Gaina et al., 1998; Schellart et al., 2006), or a subsequent, but pre-Oligocene dispersal event. The purpose of this contribution is to describe the parrots from the St Bathans Fauna, make an assessment of their relationships, and thus inform the history of New Zealand biota. Institutional AbbreviationsAM, Australian Museum, Sydney, New South Wales, Australia; CM, Canterbury Museum, Christchurch, New Zealand; NMNZ, Museum of New Zealand Te Papa Tongarewa, Wellington (formerly National Museum of New Zealand, Dominion Museum, and Colonial Museum), New Zealand; UNSW, University of New South Wales, Sydney, New South Wales, Australia. Anatomical AbbreviationsHypotarsal tendons run in canals or sulci and are identied as follows: fhl, tendon of m. exor hallucis longus; fdl, m. exor digitorum longus; pII, m. exor perforatus digiti II; ppII, m. exor perforans et perforatus digiti II; pIII/IV and ppIII, musculi exores perforati digitorum III et IV and m. exor perforans digiti III. Common anatomical terms are abbreviated as follows: artic, articularis; cond, condylus; L, left; lig, ligamentum; m, musculus; proc, processus; R, right; tuber, tuberculum. Comparative Material

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The fossils were compared to the following Recent psittaciform taxa, with an emphasis on Australasian and southwest Pacic species, grouped according to Schweizer et al. (2010). All are in the Australian Museum collection (and which have the extra prex AM), except a few from the Canterbury Museum, as follows: Strigopidae, Strigopinae: Strigops habroptilus A.1987; CM Av36986 (2 individuals), CM Av13782. Nestorinae: Nestor meridionalis A.1985; Nestor notabilis S.343. Cacatuidae: Probosciger aterrimus O.59371; Calyptorhynchus banksii S.1006; C. lathami S.696, O.68478; C. funereus O.54157, O.71387; Callocephalon mbriatum O.71406; Lophochroa leadbeateri O.70456; Eolophus roseicapillus O.71164, O.71171; Cacatua tenuirostris O.56960, O65126; C. sanguinea O.66376, O.72020; C. galerita O.65207, O.68491; C. ducorpsii S.882; Nymphicus hollandicus O.70721. Psittacidae, Arini: Ara ararauna O.68157, O.71333; Ara macao O.65776; Amazona ochrocephala S.652, O.71410; Brotogeris chiriri O.65100; Myiopsitta monachus O.59846; Pyrrhura molinae O.71400; Cyanoliseus patagonus O.72428. Psittacini: Psittacus erithacus S.579. Micropsittini: Micropsitta pusio P.2960. Psittaculini group A: Eclectus roratus S.630, A.13044; Geoffroyus geoffroyi O.59374; Alisterus scapularis O.71194, O.72078; Aprosmictus erythropterus O.60403, O.66369; Polytelis swainsonii O.60062, O.66383; P. anthopeplus O.7009, O.70092, 71327; Psittacula krameri O.65710. Platycercini group A: Platycercus elegans O.60429, O.66111, O72691; P. eximius O.59305, O.71684; P. adscitus O.60906; P. venustus O.60399; Barnardius zonarius O.71409; Northiella haematogaster O.66341; Lathamus discolor O.68484; Psephotus haematonotus O.58124, O.60422; P. varius O.58524; P. dissimilis O.65325; Prosopeia tabuensis S.1581; Cyanoramphus novaezelandiae O.64715, O.65091; Neopsephotus (= Neophema) bourkii O.66620; Neophema elegans O.64696; N splendida O.58019; Neophema pulchella O.54385, O.60918; Pezoporus wallicus O.60945, O.65346. Loricoloriinae: Agapornis roseicollis O.58461; A. lileanae O. 57155; Lorius lory or L. hypoinochrous S.708; Charmosyna placentis S.811; Trichoglossus haematodus O.54381; T. chlorolepidotus O.67628; Glossopsitta concinna O.71416, 72391; G. pusilla O.65469; G. porphyrocephala O.70783; Cyclopsitta diophthalma O.65125, O.71166; Melopsittacus undulatus O.64782, O.71326; Chalcopsitta cardinalis S.822. Fossil Sites The locations of the fossil sites from which the St Bathans Fauna has been collected are described by Worthy et al. (2007). Most of the fossils described here derive from a stratigraphic section exposed on the banks of the Manuherikia River, Otago, New Zealand (gured in Schwarzhans et al., in press). In this section, about 30 m of the base of the Bannockburn Formation is exposed and several fossil-bearing beds have been identied, with those containing parrot bones as follows: Bed HH1a, 6.887.0 m above the base of the Bannockburn Formation, main quarry at 44.907944 S 169.858222 E, Manuherikia River section; ca. 510 cm thick sand and cobble layer with abundant bone fragments, mud rip-up clasts and stromatolite fragments, and rare fragments of hyriid bivalves. Fossil Record Number in the archival Fossil Record File of the Geological Society of New Zealand H41/f88. Bed HH1b, 9.59.58 m above the base of the Bannockburn Formation, Trench Excavation, foot of hill 50 m across terrace from river bank at 44.90780 S 169.85844 E,

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METHODS Names for specic bone landmarks generally follow Baumel and Witmer (1993). In addition, we follow the denitions for features of the distal tarsometatarsus employed by Mayr (2002): for example, trochlea accessoria for the plantarly rotated lateral rim of trochlea metatarsi IV. Interpretation of hypotarsus morphol ogy is after Mayr and Gohlich (2004) and Mayr (2010b). Measurements were made with Tesa dial callipers and rounded to 0.1 mm. We follow Gill et al. (2010) in accepting three families among Recent parrots, Strigopidae Bonaparte, 1849, with Nestorinae Bonaparte, 1849, as a subfamily, Cacatuidae Gray, 1840, and Psittacidae Illiger, 1811. However, we note that Gill et al. (2010) attributed Strigopidae to Gray (1848) in error, when it is in fact correctly attributed to Bonaparte (1849), wherein Nestorinae was listed as a subfamily of Strigopidae. On the sheet that is Bonapartes (1849) Conspectus Systematis Ornithologiae, dated Decembr. 1849 under Ordo 1. Psittaci, is listed 2. STRIGOPIDAE. (Psittacidae, p. Gr) with the subfamilies 8. NESTORINAE (Cacatuinae, p. Gr. Oceania 3 [species] and 9. STRIGOPINAE (Cacatuinae, p. Gr.) Oceania, 1. Within Psittacidae, we follow the taxonomic nomenclature advocated by Christidis and Boles (2008) and Gill et al. (2010). Analyses of molecular data have recently produced a strongly supported topology of relationships within Psittacidae, with that of Schweizer et al. (2010) corroborating and expanding on previous studies of molecular data (de Kloet and de Kloet, 2005; Tavares et al., 2006; Tokita et al., 2007; Wright et al., 2008). Here we assess observed morphological structure against the phylogenetic arrangement of taxa inferred from the topology found by Schweizer et al. (2010) and employ clade names used therein.

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JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 31, NO. 5, 2011 of trochlea metatarsi II. Collected 1 March 2010 by the UNSW/CM/NMNZ expedition. DiagnosisAs for genus. EtymologyLatin, adjective for smallest, to reect this is the smallest species in the genus. Type LocalityBed HH2b, Manuherikia River section, Otago, New Zealand: details as above. Stratigraphy/Age/FaunaBannockburn Formation, Manuherikia Group, early Miocene (Altonian); 1916 Ma; St Bathans Fauna. Measurements of Holotype (mm)Length 17.8, proximal width 4.7, maximum proximal depth 3.6; least shaft width 2.0, distal width 5.5. ParatypesNMNZ S.50842 (Fig. 1FH), a proximal R tarsometatarsus with complete hypotarsus; NMNZ S.50843 (Fig. 1E, L), a R tarsometatarsus missing the end of the trochlea metatarsi II, the plantar surface of trochlea metatarsi IV, and part of the plantar surface of crista lateralis hypotarsi. Paratype LocalityBoth from Bed HH4, Manuherikia River section (details above). Measurements of Paratypes (mm)NMNZ S.50842, proximal width 4.8, maximum proximal depth 3.4; NMNZ S.50843, length 17.5, proximal width 4.7, least shaft width 1.9, preserved distal width 5.4. Referred Material (14 Specimens)The following specimens, mostly from the Manuherikia River section, were referred to this species rather than the one described below based on their smaller size and rarity compared to that taxon. Humerus, Bed HH1b, Trench Excavation: NMNZ S.50171 (Fig. 2I, J), distal R. Ulnae, Bed HH1a: NMNZ S.42671, distal L; NMNZ S.43997, distal R; NMNZ S.52223, distal R; Bed HH4: NMNZ S.50870 (Fig. 2Q), proximal L. Radius, Mata Creek, Site 2b: NMNZ S.43916, distal L, distal width 3.0 mm. Os carpi ulnare, Bed HH1b, Trench Excavation: NMNZ S.51690, R. Coracoids, Bed HH2b: NMNZ S.52683 (Fig. 2A, B), cranial and shaft R; Bed HH1a: NMNZ S.43994 (Fig. 2E), R; Bed HH1b, Trench Excavation: NMNZ S.51801, cranial part R. Scapula, Bed HH1a: NMNZ S.43993, R, proximal width 4.9 mm. Tibiotarsus, Bed HH1a: NMNZ S.52484 (Fig. 2N), distal R. Mandibles, based on size, we also tentatively refer to this species two symphysis fragments from Bed HH1a: NMNZ S.52115 and S.52163. Description and Comparison TarsometatarsusThe holotype (Fig. 1AD, I) is stained light brown and was reassembled from three pieces. Its projecting edges (rims of trochleae, cotylar margins) are slightly worn and breakage means it is missing the medial plantar projection of trochlea metatarsi II and the medial edge of the crista medialis hypotarsi. All salient features are described in the diagnosis. The two paratypic tarsometatarsi reveal some variation in the plantar surface opposite the sulcus for the tendon of m. extensor digitorum longus. In the holotype and NMNZ S.50842 (Fig. 1FH), this area of the plantar surface is only slightly convex, but in NMNZ S.50843 (Fig. 1E, L), it is marked by a prominent convex surface. In all three available fossil tarsometatarsi, the fhl is not closed over its entire proximodistal length: a similar incomplete plantar closure was noted for Bavaripsitta ballmanni and Neophema by Mayr and Gohlich (2004) and for Psittacula krameri in this study, although G. Mayr (pers. comm., 2011) noted that specimens he examined in Forschungsinstitut Senckenberg are all completely closed, so this character may vary intraspecically. In Nelepsittacus minimus, plantar closure of fhl is incomplete proximally, creating a notch in the proximal side of the bridge closing the canal. All three available tarsometatarsi have damage prohibiting interpretation of the structure of the external or medial member of the impressiones retinaculorum extensorium; however, that the lateral impression is low suggests that its counterpart will not be

Manuherikia River section; ca. 1015 cm thick sand and cobble layer with bones abundant, but more worn than those from Bed HH1a. Fossil Record Number is H41/f0103. Bed HH2b, 21.0221.31 m above the base of the Bannockburn Formation, at 44.907861 S 169.857389 E, Manuherikia River section; a ca. 35 cm thick clay layer with abundant calcied root casts, common sh bones, rare molluscs, and infrequent bird bones, overlain by about 17 cm of ne sand. Fossil Record Number is H41/f87. Bed HH4, 25.6325.83 m above the base of the Bannockburn Formation, and 2.152.35 m above the top of a prominent bed of oncolites. Manuherikia River section, 44.907861 S 169.857233 E. Fossil Record Number is H41/f0095. A few fossils derive from a nearby outcrop in Mata Creek (Worthy et al., 2007) as follows: Mata Creek, Croc Site, Layer 1, ca. 10 cm thick sand and cobble layer 3.5 m above the base of the Bannockburn Formation, in a 3-m cliff on north side of small hill on true left of Mata Creek, Otago, 44.889500 S 169.837833 E. Fossil Record Number is H41/f84. Mata Creek, Site 2b, sh bone bed 2 m above an oncolite bed in a bluff about 200 m downstream of Croc Site, 44.890450 S 169.838400 E. Fossil Record Number is H41/f80. SYSTEMATIC PALEONTOLOGY PSITTACIFORMES Wagler, 1830 STRIGOPIDAE Bonaparte, 1849 (Kakapo and allied parrots) NESTORINAE Bonaparte, 1849 (Kaka and Kea) NELEPSITTACUS, gen. nov. Type SpeciesNelepsittacus minimus, sp. nov. DiagnosisA parrot with a tarsometatarsus with the following unique combination of characteristics. (1) A plesiomorphic pattern of hypotarsal canals, where fhl and fdl are well separated and fully enclosed plantarly, with fdl largest, pII positioned mesad and plantar of fdl and enclosed on its sides but open plantarly, and all other tendons run over the plantar surface of the hypotarsus. (2) The depth of the proximal tarsometatarsus is greater through the cotyla lateralis to the plantar extremity of the crista lateralis hypotarsi than through the cotyla medialis to the tip of the crista medialis hypotarsi where it bounds pII. (3) It has a small foramen vasculare proximale mediale opening plantarly at the distal end of fdl on the ridge bounding that canal. (4) The tuberositas m. tibialis cranialis makes a convex protuberance on the medial prole. (5) It is relatively elongate, with distal width about 30% of total length. (6) The trochlea accessoria in distal aspect expands dorsally towards its medial end and is rounded medially. (7) The foramen vasculare distale is bound on the dorsal facies by a ridge extending proximal of it, creating a shallow groove extending proximad of the foramen. (8) The fossa metatarsi I is large and deep, and the facies within it is exposed in dorsal view, not just as a notch in the prole. (9) Trochlea metatarsi II is grooved distally and plantarly, and its plantar articular surface indicates a plane of rotation for the articulating phalanx towards the trochlea accessoria. Character 7 is a synapomorphy with Nestor alone of living genera, and characters 8 and 9 are considered autapomorphies of the new genus. EtymologyTo denote the fossil as a parrot on the ancestral lineage of Nestor. From the Greek mythological gure, Neleus (Greek) father of Nestor and the word psittakos (Greek), a parrot; masculine nominative. NELEPSITTACUS MINIMUS, sp. nov. (Figs. 1AI, L; 2A, B, E, I, J, N, Q) HolotypeNMNZ S.52404 (Fig. 1AD, I), a complete L tarsometatarsus missing only the medial plantar projection

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FIGURE 1. Tarsometatarsi of Nelepsittacus species in A, E, F, J, plantar view; D, H, L, M, N dorsal view; B, G, K, proximal view; C, distal view; and I, medial view. AD, I, Nelepsittacus minimus holotype, L, NMNZ S.52404; E, L, paratype, R, NMNZ S.50843; FH, paratype, proximal R, NMNZ S.50842; K, interpretation of proximal end; J, M, N. donmertoni, holotype, R, NMNZ S.52016; and N, N. daphneleeae, paratype, part R, NMNZ S.51398. Abbreviations: clh, crista lateralis hypotarsi; fdl, m. exor digitorum longus; fhl, tendon of m. exor hallucis longus; fmI, fossa metatarsi I; fvd, foramen vasculare distale; fvm, foramen vasculare mediale; lire, lateral impressiones retinaculum extensorium; mire, medial impressiones retinaculum extensorium; pII, m. exor perforatus digiti II; ppII, m. exor perforans et perforatus digiti II; pIII/IV and ppIII, musculi exores perforati digitorum III et IV and m. exor perforans digiti III; sedl, sulcus for tendon of m. extensor digitorum longus; ta, trochlea accessoria; tmtc, tuberositas m. tibialis cranialis; TII, trochlea metatarsi II. Scale bars equal 10 mm.

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FIGURE 2. Appendicular elements of Nelepsittacus species: Coracoids in A, CE, dorsal view and B, ventral view; humeri in F, H, I, L, caudal view and G, J, K, cranial view; ulnae in M, ventral view and PR, cranial view; and N, O, tibiotarsi in anterior view. A, B, Nelepsittacus minimus, NMNZ S.52683, cranial part R coracoid; E, NMNZ S.43994, R coracoid; I, J, NMNZ S.50171, distal R humerus; N, NMNZ S.52484, distal R tibiotarsus; and Q, NMNZ S.50870, proximal L ulna. C, N. donmertoni NMNZ S.51803, cranial part L coracoid; D, NMNZ S.51291, cranial part R coracoid; F, G, NMNZ S.50396, distal L humerus; H, NMNZ S.43995, proximal R humerus; and O, NMNZ S.42784, distal R tibiotarsus. K, L, N. daphneleeae NMNZ S. 50826, holotype, distal R humerus; M, NMNZ S.50609, distal L ulna. P, species 4 NMNZ S.52411, proximal L ulna. R, Nestor notabilis AM S.343, proximal L ulna. Abbreviations: cdu, cond. dorsalis ulnaris; cl, cotyla lateralis; cms, crista m. supracoracoidei; cs, cotyla scapularis; csr, capital shaft

WORTHY ET AL.MIOCENE PARROTS OF NEW ZEALAND robust and dorsally elevated as it is in many cacatuids or other taxa with short tarsi. The moderately elongate tarsometatarsus of N. minimus distinguishes it immediately from all cacatuids, the tribe Arini, and most members of Loricoloriinae within psittacids. A hypotarsus with just two enclosed canals, for fhl and fdl, and a well-dened sulcus for pII is shared by N. minimus, strigopids, and cacatuids, but excludes many parrot genera with various apomorphic modications to this pattern. All members of Schweizer et al.s (2010) platycercine group A are characterized by what Mayr (2008) termed a platycercini hypotarsus morphology or his derived pattern A (Mayr, 2010b), in which the crista lateralis hypotarsi is developed as a prominent ange to largely enclose a canal for pIII/IV and ppIII plantarly, and the crista medialis hypotarsi is plantarly deepened, with the canal for pII usually wholly enclosed, and, plantar to it, the canal for ppII is also well dened or enclosed. Within this group there is variation with linkages of fdl to pII (e.g., Prosopeia, Cyanoramphus, Northiella, and Lathamus), and occasionally fhl may be partly open to the canal for pIII/IV and ppIII (e.g., Cyanoramphus). Forming the sister clade to these core platycercines in Schweizer et al.s (2010) topology, Neophema and Neopsephotus have very small tarsi and further differ by lacking the large plantar-medial development of the crista lateralis hypotarsi. However, both taxa have a deeper crista medialis hypotarsi than the St Bathans fossils. In Neophema, this results in enclosure of the linked canals fdl, pII, and ppII, but in Neopsephotus, fdl remains discrete and pII is open plantarly. The sister group to platycercine group A, the Loricoloriinae of Mayr (2008), which includes Cyclopsittacini (e.g., Cyclopsitta), Loriini (e.g., Lorius, Charmosyna, Glossopsitta), some Psittaculini (e.g., lovebirds, Agapornis, and hanging parrots, Loriculus), and budgerigar (Melopsittacus), are dened by the complete enclosure of all tendons within the hypotarsus, Mayrs (2010b) derived pattern B. There is variation within this group from an extreme of all canals linked as one complex (Loriini), two canals offset plantarly from each other, where fdl + fhl lie dorsally and the other tendons in a single canal more plantarly (Agapornis and Loriculus), or various other combinations of canal linkage (Mayr, 2008). We found that Cyclopsitta had a single complex canal as in Loriini, but differed from genera such as Trichoglossus by this canal being partly open plantarly, although Mayr (2008, 2010b) reported that fhl, fdl, and pII were discrete canals. Different ossication of the intervening septa may relate to ontogenetic age. We prefer to include Cyclopsitta in Mayrs group characterized by his derived pattern B hypotarsus morphology, because it is closer to this than the platycercini type where there is no plantarlateral outgrowth of the crista medialis hypotarsi towards that on the crista lateralis hypotarsi: in Cyclopsitta it appears a small step to completely ossify the plantar enclosure of the tendinal canals. The sister group of Loricoloriinae plus platycercine group A, sensu Schweizer et al. (2010), is Micropsittini plus Psittaculini Group A (includes Alisterus, Aprosmictus, Polytelis, Prioniturus, Eclectus, Geoffroyus, and Psittacula). Micropsitta differs from Nelepsittacus minimus by having a hypotarsal structure like that of Melopsittacus, as noted by Mayr (2008, 2010b). Prioniturus has fusion of fhl and fdl in a single canal, with all deeper tendons partly enclosed in a sulcus plantarly (Mayr and Gohlich, 2004).

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The remaining taxa are similar to N. minimus, with a plantarly low crista lateralis hypotarsi that lacks a medial projection. Aprosmictus and Polytelis differ from N. minimus and are like some members of platycercine group A, with fdl, pII, and ppII all linked and wholly enclosed. Eclectus has the canal for pII wholly enclosed and that for ppII distinct, but open. Alisterus has a canal structure quite similar to N. minimus, with fdl and fhl discrete and pII open plantarly. However, this genus, as in Eclectus, Psittacula, Aprosmictus, Polytelis, and Prioniturus, differs from N. minimus in the mediolateral position of pII relative to fdl. In N. minimus, pII is distinctly offset mediad of the lateral margin of fdl, whereas in all these taxa, pII is positioned entirely plantar of fdl. In this feature, N. minimus is like Nestor (pII entirely medial in N. meridionalis; pII partly overlaps fdl in N. notabilis), most cacatuids (pII entirely medial in Callocephalon, Probosciger, Cacatua, and Nymphicus; pII partly overlaps fdl in Calyptorhynchus), and members of Arini, suggesting that a relatively medial location for pII is plesiomorphic. In Strigops, however, although the external margin of pII is only slightly medial to fdl, pII broadly overlaps fdl on the lateromedial plane. All members of Arini are distinguished by a distinct pattern of the hypotarsus, in which fdl and fhl are usually the only enclosed canals, with only pII otherwise variably present (e.g., open plantarly in Ara and Myiopsitta, enclosed in Amazona, and not apparent in Cyanoliseus), and all other tendons running over the plantar surface of the hypotarsus in very poorly dened sulci. Normally fdl and fhl are close together and their combined width is relatively narrow compared to the width of the tarsometatarsus. Characteristically, fdl slightly overlaps fhl on its cranial side and sometimes the septum is not ossied (e.g., Myiopsitta, Pyrrhura). The single representative of Psittacini examined, Psittacus erithacus, has enclosed fhl, fdl, pII, and ppII, with fhl and fdl not overlapping as in Arini, and pII plantar to fdl. The close proximity of fdl and fhl in species within Arini is also seen in most cacatuids and is another character distinguishing members of this tribe from Nelepsittacus minimus. In Cacatua (galerita, tenuirostris, sanguinea, ducorpsii), Callocephalon mbriatum, and Calyptorhynchus banksii, canal fdl was close to fhl and separated by a narrow osseous septum such that fdl overlapped fhl dorsally. In other taxa (Nymphicus, Calyptorhynchus funereus, C. lathami, Eolophus roseicapillus, and Lophochroa leadbeateri), the canals were close but did not overlap each other. Only Probosciger among cacatuids was observed to have wellseparated canals. The greater depth of the proximal tarsometatarsus through the cotyla lateralis to the plantar extremity of the crista lateralis hypotarsi compared to that through the cotyla medialis to the tip of the crista medialis hypotarsi distinguishes Nelepsittacus minimus from several of the taxa with a plesiomorphic canal pattern. Strigops has the least well-developed crista lateralis hypotarsi among extant psittaciforms, and its greatest proximal depth is through the cotyla medialis to a point beside a partly enclosed pII. A relatively shallow lateral side of the proximal tarsometatarsus also characterizes all cacatuids (e.g., Nymphicus, Probosciger, Cacatua, Calyptorhynchus, Eolophus, and Callocephalon), Psittacus, most examined taxa in Psittaculini group A (e.g., Eclectus, Alisterus, Psittacula, Aprosmictus, and Polytelis), and many of the platycercine group A taxa.

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ridge; cv, cotyla ventralis; dbmr, impression of the dorsal branch of m. extensor carpi radialis; dtre, distal tuberositas retinaculi extensoris; iba, impression of brachialis anticus; ic, incisura capitis; laa, attachment lig. acrocoraco-acromion; lap, attachment lig. acrocoraco-procoracoideum; ltmf, lateral tuber. retinaculi m. bularis; pbmr, impression of the palmar branch of m. extensor carpi radialis; pcd, proc. cotyla dorsalis; pf, proc. exorius; pp, proc. procoracoideus; sh, sulcus humerotricipitalis; tbu, tuber. bicipitale ulnae; tc, tuber. carpale; td, tuber. dorsale; tsv, tuber. supracondylare ventrale. Note that in A and B the proc. procoracoideus is preserved with some sediment adhering to its ventral side to ensure its retention, but which makes it look thicker than it is. Scale bar equals 10 mm.

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JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 31, NO. 5, 2011 does not extend onto the medial facies. In Strigops, there is no evidence of such a sulcus. In summary, Nelepsittacus minimus shows greatest similarity to Nestor (both species), sharing one exclusive (among extant taxa) synapomorphy, a distinct groove extending up the shaft from the foramen vasculare distale, but is distinguished from it by several features that support its generic distinction. This similarity with Nestor and the lack of shared apomorphies with members respectively of Loricoloriinae, platycercini group A, Psittaculini group A (including Micropsitta), Arini, Cacatuidae, and its distinction from Strigops support Nelepsittacus minimus being provisionally classied in Nestorinae. HumerusThe humerus NMNZ S.50171 (Fig. 2I, J) is referred to Nelepsittacus minimus because it exhibits most of the following psittaciform features seen in distal humeri (breakage precludes assessing characters 1 and 2, included here for completeness because of their relevance to assessing other taxa hereafter): (1) the lack of a proc. supracondylaris dorsalis, with the dorsal origin of m. extensor carpi radialis (Vanden Berge and Zweers, 1993: annotation 87) instead of a variable-shaped rugosity on the dorsocranial margin proximal of the cond. dorsalis; (2) a distinct rounded scar, sometimes a tuberosity, immediately ventral to the dorsal origin of m. extensor carpi radialis and proximal to the cond. dorsalis, termed tuber. origii m. pronator supercialis by Livezey and Zusi (2006:character 1463), or a tubercle above dorsal condyle by Mayr (2004:g. 4), or the impressio of the palmar branch of m. extensor metacarpi radialis as described for passerines (Hamon, 1964; Noriega and Chiappe, 1993), but which is the ventral head of origin for m. extensor carpi radialis (Vanden Berge and Zweers, 1993); (3) a shallow fossa brachialis; (4) a large, elevated tuber. supracondylare ventrale; (5) proc. exorius bears a pair of deep pits for the origin of m. extensor ulnaris (cranially) and the origin of m. pronator profundus (caudally) on the ventral facies, which in cranial aspect presents a notch in the ventral prole due to prominence of the epicondylus ventralis on the proximal side of these pits; (6) the condyli dorsalis et ventralis and proc. exorius have equal distal extent; (7) a narrow, well-dened sulcus scapulotricipitalis caudally; and (8) a shallow fossa olecrani. The humerus NMNZ S.50171 of Nelepsittacus minimus is small (distal width 7.2 mm) and outside the range of the slightly larger and more numerous taxon described below (Table 1). It preserves the condyli dorsalis et ventralis and proc. exorius and is broken across the proximal side of tuber. supracondylare ventrale, but has the above features 38. The fossa m. brachialis sharply undercuts the tuber. supracondylare ventrale. In distocaudal view, the proc. exorius is relatively dorsoventrally wide, little raised above the adjacent sulcus humerotricipitalis, and features a distinct groove traversing its width, which forms the distal side of the ligamental attachment point. A wide proc. exorius is shared with Nestor and cacatuids, to the exclusion of other taxa, especially those of Psittaculini group A, where it is very narrow. That the proc. exorius is little elevated caudally above the adjacent sulcus humerotricipitalis distinguishes N. minimus from all psittacids and cacatuids, and is an apomorphy shared only with Nestor and Strigops among compared taxa. UlnaThese ulnae are distinguished from the following species by their smaller size (Table 2). The proximal ulna (NMNZ S.50870; Fig. 2Q) of Nelepsittacus minimus is missing the olecranon and is broken through the shaft distal to the impressio brachialis. The impressio brachialis is deep and undercuts its caudal margin. The tuber. bicipitale ulnae (Livezey and Zusi, 2006), or tuber. musculi bicipitis (Ballmann, 1969), is complex with two main parts: (1) an elongate ridge-like tuberosity starting proximally just below (distal of) the cotyla dorsalis and extends c. 2.5 mm distally and ventrally; and (2) from below the lip of the cotyla ventralis, another elongate ridge, aligned roughly at right angles to the rst, runs distad and dorsad to end close to the

The plantar development of the crista lateralis hypotarsi further distinguishes Nelepsittacus minimus from taxa with a more plesiomorphic hypotarsal structure. In N. minimus, this crista extends plantarly as a robust ridge, such that in proximal view, it is seen to laterally enclose and so dene a sulcus for ppIII/IV and ppIII that is deeper than in all cacatuids. In lateral view, this crista is seen to extend proximad with increased plantar depth so that its most plantar point is proximal to the cotyla lateralis. In plantar aspect, this crista projects proximolaterad, and is not undercut either distally or laterally. In this, it differs markedly from all cacatuids, including Nymphicus, in which the plantar surface of the crista lateralis hypotarsi markedly overhangs, or is undercut by, the shaft both distally and laterally. Nelepsittacus minimus is, however, like Nestor and Strigops in this respect. A small medial vascular foramen is shared with Nestor. A larger, more obvious foramen is seen in Strigops, Psittacus, most Loricoloriinae, including Melopsittacus, and many members of Psittaculini group A (including Eclectus, Alisterus, Aprosmictus, and Polytelis). A few platycercines also have a foramen: it may be small or absent in Platycercus elegans and Psephotus haematonotus, but is large in Northiella and distinct in Lathamus. That the tuberositas m. tibialis cranialis causes a marked convexity on the medial margin of the shaft distinguishes Nelepsittacus minimus from all cacatuids and Loricoloriinae, and many other taxa (including Neophema, Neopsephotus, Eclectus, Prosopeia, Cyanoramphus, Psephotus, Barnardius, and Platycercus). The medial prole in Strigops is only slightly convex beside the tuberositas m. tibialis cranialis. The elongate groove extending proximad from the foramen vasculare distale on the dorsal facies, formed by the presence of a ridge extending up the shaft from the trochlea metatarsi IV, is a feature of Nelepsittacus minimus observed only in Nestor among examined modern taxa. The form of the trochlea accessoria is relatively simple in Nelepsittacus minimus. In distal view, it expands signicantly dorsally towards its medial end, so partly occluding the furrow separating it from trochlea metatarsi IV, and at its medial end is rounded, not bearing a groove aligned parallel to its dorsal and ventral margins. This distinguishes it from all members of Arini, which show the least dorsal expansion among all taxa examined, with dorsal and ventral margins (as seen in medial view) of the whole trochlea accessoria being nearly parallel. Nestor has an autapomorphic condition, not shared with the fossil, of the medial end of the trochlea accessoria being greatly expanded dorsally and deeply grooved on its medial tip giving it a bid appearance. Nelepsittacus minimus differs from all psittaciforms examined by the form of the trochlea metatarsi II, which in plantar view is seen to be well grooved over its distal end, and which groove indicates that the plane of rotation of the articulating phalanx is towards the trochlea accessoria. In all other psittaciforms examined, this plane of rotation is towards a point proximal of the trochlea accessoria. Nelepsittacus minimus shares with Nestor a relatively elongate tarsometatarsus with a distal width about a third of total length, a similar pattern of hypotarsal canals, a minute foramen vasculare medialis plantarly, a marked convexity to the medial shaft surface adjacent to the tuber. m. tibialis cranialis, and the apparent synapomorphy of a groove preceding the foramen distale on the dorsal facies. It is distinguished from Nestor by three features, in addition to the form of the trochlea accessoria: (1) the fossa metatarsi I is larger and deeper, and impinges on the medial prole, extending (autapomorphically) onto the dorsal facies; (2) the crista lateralis hypotarsi is lateromedially thinner in the area bounding the fhl canal, so that the distal exit of fhl is relatively closer to the lateral facies; and (3) the distal exit of canal pII lies ventral to the bounding crista laterad of fdl and is passed on its medial side by a shallow sulcus extending from the plantar shaft facies up the medial facies to near the lip of the cotyla medialis. In Nestor, a similar sulcus undercuts the exit of pII, but

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WORTHY ET AL.MIOCENE PARROTS OF NEW ZEALAND

TABLE 1. Measurements (mm) for humeri of Nelepsittacus species from the St Bathans Fauna. Distal width maximum 7.3 7.7 8.3 8.1 7.9 10.9 Depth dorsal condyle 4.3 4.5 4.8 4.6 6.3

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Species N. minimus N. donmertoni N. donmertoni N. donmertoni N. donmertoni N. donmertoni N. donmertoni N. donmertoni N. daphneleeae

Catalog number S.50171 S.42252 S.43995 S.43996 S.50190 S.50396 S.50864 S.51422 S.50826

Proximal width 11.9

Shaft width 3.3 3.3 3.6 3.4

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proximal end of the tuberosity. This second ridge has proximal and distal parts that are contiguous, whereas in many psittaciforms, the parts are separate, as in Platycercus elegans and Psittacula. Distal ulnae of parrots vary little and the referred specimens exhibit no special features. CoracoidMeasurements (mm): NMNZ S.52683 (Fig. 2A, B), length from the sternal side of the humeral facet to the end of the proc. acrocoracoideus 6.5; width of humeral facet 2.6; NMNZ S.43994 (Fig. 2E), medial length 23.7, width sternal facet 6.5. The fossil coracoids referred to Nelepsittacus minimus have the general form of psittaciforms and share with all parrots a prominent bulge dorsally between the cranial end of the facies artic. humeralis and the sulcus m. supracoracoidei. The better-preserved NMNZ S.52683, from the same bed as the holotype, differs from nearly all coracoids of extant psittaciforms (slight hollow seen in Psittacula), by the cotyla scapularis having a shallow, but distinct central depression, rather than forming a transverse groove across the axis. The facies artic. clavicularis has a prominent ventrally located projection, extending over the sulcus m. supracoracoidei, to which the lig. acrocoraco-procoracoideum was attached. The attachment of lig. acrocoraco-acromion forms a rounded dorsally prominent bulge on the dorsal side of the facies clavicularis, but does not overhang the sulcus m. supracoracoidei. The facies artic. clavicularis markedly overhangs a pneumatic fossa that deeply penetrates the proc. acrocoracoideus and is dorsoventrally wide, such that it undercuts the base of the ventral tuberosity. The

TABLE 2. Measurements (mm) for ulnae of Nelepsittacus species from the St Bathans Fauna. Distal width dorsal condyle 3.8 3.6 3.6 4.1+ 4.4 4.2 3.9 4.5 4.3 4.3 4.5 4.3 4.1 5.9+

Species N. minimus N. minimus N. minimus N. minimus N. donmertoni N. donmertoni N. donmertoni N. donmertoni N. donmertoni N. donmertoni N. donmertoni N. donmertoni N. donmertoni N. donmertoni N. daphneleeae

Catalog number S.52223 S.50870 S.42671 S.43997 S.42615 S.42832 S.44211 S.51407 S.51136 S.51915 S.51695 S.51501 S.51317 S.52455 S.50609

Proximal width 5.1

Shaft width 2.4 2.7 2.8 2.9 3.8

Distal width maximum 4.7 4.2 4.6 5.1+ 5.1 5.3 5.2 5.3 5.2+ 5.2 5.2 5.2 6.6

+ indicates that the measurement is a minimum and would increase slightly if the bone was not worn.

proc. procoracoideus is complete on NMNZ S.52683 and shows that it extends craniomediad from the shaft a distance equivalent to the length of its origin on the shaft, the tip is broadly rounded, not pointed, and ends slightly cranial to the cotyla scapulae. These referred coracoids thus are similar to those of strigopids with a dorsally prominent bulge for the lig. acrocoraco-acromion, and so differ from other psittaciforms, which lack such a dorsal prominence. However, they differ from coracoids of strigopids, because the same tuberosity for the lig. acrocoraco-acromion is not prominent sternally over the sulcus m. supracoracoidei. This lack of a sternally directed prominence may relate to the small size of N. minimus, because in the larger taxa described below, this tuberosity is markedly protuberant sternally. Coracoids of N. minimus further differ from those of Strigops and Nestor in having greater pneumaticity of the proc. acrocoracoideus: in Nestor, pneumatic foramina are usually restricted to small area under the dorsal part of the facies artic. clavicularis; in Strigops, a small area is variably present under the ventral process extending from the facies artic. clavicularis. Coracoids of N. minimus also differ from strigopids in the form of the proc. procoracoideus. Strigops often has the proc. procoracoideus extending well cranial to the cotyla scapulae, and occasionally it is linked to the proc. acrocoracoideus via the ventral process on the facies artic. clavicularis. Nestor is characterized by a reduced proc. procoracoideus, such that in N. meridionalis it barely protrudes from the shaft and in N. notabilis, although extending mediad, it lacks any cranial projection. That the facies clavicularis markedly overhangs the sulcus supracoracoideus distinguishes Nelepsittacus from members of Arini, in which the overhang is slight. TibiotarsusThe tibiotarsus NMNZ S.52484 (Fig. 2N) of Nelepsittacus minimus is worn, and with a distal width of 4.5 mm, is of appropriate size for the holotype tarsometatarsus. Although worn, it displays typical psittaciform features as follows: distal end broad, caudally attened, and craniocaudally shallow; distal condyli of similar size, their proximodistal length about half of the distal width, and well separated from each other; a short pons supratendineus; and the distal or medial tuber. retinaculi m. bularis is conuent with, or occupies the same crista as, the distal tuberositas retinaculi extensoris. The specimen presents the following features: the tuberculi retinaculi m. bularis medialis et retinaculi extensoris is an elongate, very robust, and cranially prominent tuberosity; the tuber. retinaculi m. bularis lateralis forms a low elongate crest on the lateral shaft margin adjacent to its counterpart and separated from it by a at groove aligned with the shaft; these tuberculi are separated from the cond. lateralis by a distance slightly greater than the proximodistal length of the cond. lateralis; the sulcus extensorius is deep adjacent to these tuberculi and lies entirely within the medial half of the shaft; the pons supratendineus was probably completely ossied, with erosion breaking it in NMNZ S.52484; the incisura intercondylaris is broad and secondarily excavated into the cond. lateralis; and the cond. lateralis extends slightly distal to the cond. medialis. Nelepsittacus minimus is similar to Nestor and unlike all other psittaciforms in having the tuberculi retinaculi m. bularis medialis et retinaculi extensoris separated from the cond. lateralis by a distance greater than or equal to the proximodistal length of that condyle: in all other psittaciforms, the separation is much less than the length of the condyle. In N. minimus, the tuber. retinaculi m. bularis lateralis forms a low crest on the lateral margin adjacent to the tuber. retinaculi m. bularis medialis et tuberositas retinaculi extensoris, as seen in Strigops, Nestor, and many psittacids. However, cacatuids (except Nymphicus) are distinguished by the tuber. retinaculi m. bularis lateralis being offset distally from its medial counterpart and enlarged, with greatest development seen in Cacatua galerita, C. ducorpsii, and Eolophus roseicapillus. In the last two taxa, the crest is very prominent laterally and extends distad onto the lateral facies of the cond. lateralis. Similarly, Micropsitta and all members of

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JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 31, NO. 5, 2011 Paratype LocalityNMNZ S.50261 and S.50311 are from Bed HH1a; NMNZ S.52049 is from Bed HH1b Trench Excavation in the Manuherikia River section, Otago: for details see above. Referred Material(56 specimens; measurements in mm.) Bed HH1a (36 specimens). Humeri (Table 1): NMNZ S.42854, part proximal R, (Worthy et al., 2007:g. 14A, D); NMNZ S.43995 (Fig. 2H), proximal R; NMNZ S.43996, distal L; NMNZ S.50267, proximal R; NMNZ S.50281, distal R; NMNZ S.50396 (Fig. 2F, G), distal L; NMNZ S.51422, distal R. Ulnae (Table 2): NMNZ S.42615, distal L; NMNZ S.42832, distal L; NMNZ S.44211, distal L; NMNZ S.50265, proximal L; NMNZ S.51407, distal L; NMNZ S.51501, distal L; NMNZ S.52455, distal R. Radii: NMNZ S.44308, distal L, distal width 3.7; NMNZ S.52259, distal R, distal width 3.8; NMNZ S.52541, distal L, distal width 3.7; NMNZ S.52622, distal R, distal width 3.8; NMNZ S.52623, distal L, distal width ca. 3.6. Femora: NMNZ S.42723, proximal R, proximal width 5.4; NMNZ S.44310, distal L femur, distal width 5.9. Tibiotarsi: NMNZ S.42784 (Fig. 2O), distal R, preserved distal width 5.7 (estimated 6.4); NMNZ S.42853, worn distal L; NMNZ S.50129, distal L, preserved distal width 5.6; NMNZ S.50301, shaft L; NMNZ S.52226, distal R. Tarsometatarsi: NMNZ S.42614, trochlea metatarsi III of R; NMNZ S.42856, trochlea III and part of distal end L; NMNZ S.44309, proximal R, minimum shaft width 2.3; NMNZ S.51720, fragment of a R, with trochlea metatarsi IV and trochlea accessoria. Coracoids: NMNZ S.40458 sternal end L; NMNZ S.42663, sternal end R, width sternal facet 6.7; NMNZ S.42855, sternal end R, width sternal facet 7.0 (Worthy et al., 2007:g. 14J); NMNZ S.50277, imperfect L; NMNZ S.51409, cranial part L. Scapulae: NMNZ S.42833, L, proximal width 6.2 (Worthy et al., 2007:g. 14I). Bed HH1b, Trench Excavation (17 specimens). Humeri (Table 1): NMNZ S.50190, distal R; NMNZ S.50864, distal R; NMNZ S.51905, proximal R. Ulnae (Table 2): NMNZ S.51136, distal L; NMNZ S.51317, distal R; NMNZ S.51695, distal L; NMNZ S.51915, distal R. Carpometacarpus: NMNZ S.51936, proximal R, proximal width 6.7. Os carpi ulnare: NMNZ S.51878, L. Coracoids: NMNZ S.51291 (Fig. 2D), cranial part R; NMNZ S.51803 (Fig. 2C), cranial part L, length from the sternal side of the humeral facet to the end of the proc. acrocoracoideus 6.8, width of humeral facet 2.7; NMNZ S.51900, cranial part L, length from the sternal side of the humeral facet to the end of the proc. acrocoracoideus 7.9; width of humeral facet 2.7. Scapula: NMNZ S.51625, L, proximal width 5.8. Tibiotarsus: NMNZ S.51972, distal part shaft R. Mandible: Based on size, we tentatively refer three mandible symphysal fragments: NMNZ S.51160, S.51624, and CM Av40879. Mata Creek, Croc Site, Layer 1 (three specimens). Humerus: NMNZ S.42252, shaft L (Worthy et al., 2007:g. 14B, E). Carpometacarpi: NMNZ S.42409, proximal L, proximal width 5.1+; NMNZ S.42480, R, length 24.7. Description TarsometatarsusThe holotype NMNZ S.52016 (Fig. 1J, M) is stained black and has been reassembled from two fragments such that the trochlea metatarsi III is not correctly oriented, being directed too much in line with the shaft rather than with a medial inclination. Apart from size and the other features listed in the diagnosis, Nelepsittacus donmertoni differs from N. minimus in having a shallower fossa metatarsi I, which is entirely on the facies medialis and at right angles to the facies dorsalis, so the oor of the fossa is not visible in dorsal view. As in N. minimus, the ridge lateral of the foramen vasculare distale extends proximad to proximodistally overlap the fossa metatarsi I. HumerusThe fossil humeri attributed to Nelepsittacus donmertoni, as best exemplied by NMNZ S.50396 (Fig. 2F, G) and S.50864, exhibit all the above listed psittaciform features and are approximately the same size as Platycercus elegans (Table 1). The

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Loricoloriinae, including Melopsitticus, have a distally displaced and enlarged tuber. retinaculi m. bularis lateralis developed into a large crest. Micropsitta and all lories and kin are oral feeders, and often climb about upside down while feeding. Eolophus, like many cacatuids, is primarily a ground feeder, foraging on seeds; however, it likes to play and while doing so frequently hangs about upside down, and this species also performs a rain dance involving it hanging upside down (Higgins, 1999). Cacatua galerita also spends considerable time hanging upside down while foraging. Cacatua ducorpsii is one of the tropical species of Cacatua and is primarily a canopy feeder (Forshaw, 1989), thus is likely to spend considerable time clambering about upside down on terminal branches. Thus we interpret the development of the tuber. retinaculi m. bularis lateralis into a large crest as a homoplasy related to, and facilitating, the arboreal agility of these taxa. The likely presence of a completely ossied pons supratendineus (shown for N. donmertoni below), shared with Nestor, distinguishes it from taxa in which the osseous bridge is incomplete, including Strigops, several genera within Arini (we note Mayrs [2010b] survey indicates several taxa that have a bridge), the cacatuids Probosciger, Cacatua, Callocephalon, Eolophus, and usually in Calyptorhynchus (but ossied in C. funereus AM O.54157, O.71387), but not Nymphicus. The medial location of the sulcus extensorius in Nelepsittacus distinguishes it from Nestor and Strigops, wherein the sulcus is centred on the shaft. MandibleThe two mandible fragments tentatively referred to N. minimus, based on smaller size than those described hereafter, reveal that the symphysal zone is dorsally concave along the axial plane in the part bearing the rhamphotheca. Dorsally, the caudal part of the symphysal zone has a rounded ridge running parallel to the caudal margin of the mandible that separates the rhamphotheca from the caudal edge of the mandible. Immediately anterior to this ridge and in a median position is a shallow fossa with three foramina entering the corpus. Ventrally, the surface is crenulated and on each side a marked groove passes from the caudal end anteriorly along the sides and onto the attened ventral symphysis to end in a foramen. NELEPSITTACUS DONMERTONI, sp. nov. (Figs. 1J, M; 2C, D, FH, O) HolotypeNMNZ S.52016 (Fig. 1J, M), a R tarsometatarsus missing the cotyla lateralis, most of the hypotarsus, the trochlea metatarsi II, and the end of the trochlea accessoria. Collected between 8 and 16 January 2008, by the UNSW/CM/NMNZ expedition. DiagnosisTarsometatarsus larger (22% longer) than Nelepsittacus minimus, plantar opening of medial vascular foramen larger, sulcus for the tendon of m. extensor digitorum longus shallower as it passes the impressiones retinaculi extensorii medialis. EtymologyFor the late Don Merton (19392011), in recognition of his major contributions over many years in the conservation of New Zealand birds, and in particular of kakapo Strigops habroptilus. Type LocalityBed HH1b, Manuherikia River section, Otago, New Zealand (details above). Stratigraphy/Age/FaunaBannockburn Formation, Manuherikia Group, early Miocene (Altonian); 1916 Ma; St Bathans Fauna. Measurements of Holotype (mm)Length (actual) 21.8, estimated total length 22.0, least shaft width 2.4. ParatypesNMNZ S.50261, a proximal end and shaft of R tarsometatarsus preserving fdl and fhl, but more plantar parts of hypotarsus lost; NMNZ S.50311, a R tarsometatarsus part proximal end and shaft, with part cotyla medialis and dorsal parts fdl and fhl; NMNZ S.52049, part L tarsometatarsus, shaft, and trochlea accessoria.

WORTHY ET AL.MIOCENE PARROTS OF NEW ZEALAND origin of the dorsal branch of m. extensor carpi radialis straddles the dorsal margin and is not prominent cranially and narrowly separated from the more ventrally located impressio of the palmar branch of m. extensor carpi radialis, which is also not prominent. This latter is separated from the cond. dorsalis by an arced sulcus whose curve is proximally convex. The fossa m. brachialis is at, with the impression of brachialis anticus within it extending to the ventral margin and abutting the tuber. supracondylare ventrale in a gentle curve (rather than abruptly), and having an elliptical shape aligned along the shaft. As in most psittaciforms, but unlike Nestor, these fossils exhibit a shaft that is not markedly convex cranially at the proximal end of the fossa m. brachialis and the pits on the proc. exorius are distinct and clearly divided by a medium ridge. The at fossa m. brachialis distinguishes Nelepsittacus donmertoni from all cacatuids, in which taxa it is deeper than in all psittacids. Strigops and Nestor species share with N. donmertoni a very shallow fossa m. brachialis, but Nestor differs with greater ination/convexity of the cranial shaft facies at the level of the fossa than observed in the fossil and other psittaciforms. Nestor humeri differ more trenchantly from that of N. donmertoni in that on the proc. exorius, the pits for the origins of m. extensor ulnaris and m. pronator profundus are less distinctly separated by a ridge; the pits are clearly separated in Strigops, cacatuids, and all psittacids. The proximal humerus is best exemplied by NMNZ S.43995 (Fig. 2H), which is complete except for loss of the proximal side of the tuber. ventrale; however, the entire margin of the fossa pneumotricipitalis is preserved. It is similar to humeri of Nestor meridionalis, for example NMNZ OR.28161, in that the capital shaft ridge connects to the caput humeri just dorsal to a distinct tubercle; this ridge bears two prominent ligamental attachment scars, one overlapping with, and extending distad of, the crista m. supracoracoidei, and a second farther distally and directed towards the cranial facies; crista m. supracoracoidei elongate and proximally overhung caudally by the tuber. dorsale; incisura capitis with sulcus in distal half closed distally by a low ridge; fossa pneumotricipitalis with few pneumatic foramina penetrating its internal wall; impressio coracobrachialis deep and well dened ventrally, extending distad to a point level with the distal margin of the crista bicipitalis; and the cranial part of the crista deltopectoralis is worn so its height in indeterminate; however, it extends well distad of the crista bicipitalis and in its distal half rises from the cranial facies at a wide angle, not nearly at a right angle as in Platycercus, for example. Humeri of Strigops differ from those of Nestor by having a shorter and more elevated tuber. dorsale, a lower capital shaft ridge, a deeper sulcus in the incisura capitis, and greater pneumaticity of the fossa pneumotricipitalis. Ulnae, Radii, Os Carpi Ulnare, ScapulaeThe ulnae (Table 2) referred to Nelepsittacus donmertoni exhibit no special features and differ little from those of Platycercus. No signicant differences in ulnae were found to distinguish nestorines from psittacids. Similarly, the fossil radii, os carpi ulnare, and scapula were referred to N. donmertoni based on their size being similar to that of Platycercus elegans; no attempt was made to detect differences among psittaciforms for these elements. However, the acromion on the referred scapula is less cranially prominent than it is in Platycercus. CarpometacarpusThe referred carpometacarpi are relatively worn and are referred by their general similarity to those of parrots. The proximal end is relatively smaller than in Platycercus, but has similar proportions to that of Nestor carpometacarpi. Also, as in Nestor, the foramen in the fossa m. infratrochlearis is relatively large. CoracoidThe fossil coracoids referred to Nelepsittacus donmertoni, as best exemplied by NMNZ S.51803 (Fig. 2C), S.51900, and S.51291 (Fig. 2D), are slightly larger than those referred to N. minimus. As in N. minimus, these coracoids have a shallow and distinct central depression in the cotyla scapularis

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and a prominent ventrally located projection on the facies artic. clavicularis extending over the sulcus m. supracoracoidei to which the lig. acrocoraco-procoracoideum was attached. However, they differ from N. minimus in that the attachment of lig. acrocoraco-acromion on the dorsal side of the facies artic. clavicularis is larger, forming a distinct projection that is separated by a deep notch from its ventral counterpart. The presence of prominent tuberosities for each of these ligamental attachment points is an apomorphy shared with strigopids, in contrast to all cacatuids and psittacids in which the dorsal ligament attachment point is linear and not developed as a projection both dorsally and sternally. The structure is most similar to Nestor, which has near-equal-sized projections separated by a deep notch for these ligament attachment points. Strigops has a rounded dorsal tuberosity with a weak and variable depth notch separating it from a markedly more elongate ventral tuberosity, which is occasionally linked to the proc. procoracoideus by ossication of the lig. acrocoraco- procoracoideum. The fossils have greater pneumaticity of the acrocoracoid than does Nestor, in which pneumaticity is mainly restricted to a small area in the dorsal half of the sulcus m. supracoracoidei, under the projection for the attachment of lig. acrocoraco-acromion. This pneumatic fossa in coracoids of N. donmertoni is relatively smaller than in the coracoids referred to N. minimus, because it does not undercut the ventral ligament attachment point to the same extent. The fragment NMNZ S.42663 reveals that the sternal end of the coracoid has a prominent crista medialis extending craniad of the medial angle a distance equivalent to about half the width of the sternal end and that the impressio m. sternocoracoidei is deep, and so is similar to N. minimus (Fig. 2E). TibiotarsusThe distal tibiotarsus NMNZ S.50129 (Fig. 2O) is worn but is estimated to have been 5.8 mm wide and, similarly, NMNZ S.42784 is estimated to have been 6.4 mm wide. These tibiotarsi are larger than, but otherwise have identical morphology to, that of Nelepsittacus minimus. The fragment NMNZ S.42784 reveals that the medial tuber. retinaculi m. bularis is distinct from, and located proximal to, the distal tuberositas retinaculi extensoris and is at the same level as the tuber. retinaculi m. bularis lateralis and the proximal tuberositas retinaculi extensoris. The fragment NMNZ S.42853 shows that an ossied pons supratendineus characterizes the species, with wear opening the pons in other available specimens. MandibleThe mandible fragments, apart from being larger than those referred tentatively to N. minimus, have a different form. As in Nestor, the median plane through the symphysis reveals an anterior zone that supports the rhamphotheca, which is anterocaudally planar (not concave) and at a markedly shallower angle to the caudal section of the symphysis, which rises steeply from the caudal margin. This caudal section also features a shallow median sulcus and is less than half as long as the anterior section. Ventrally, the fragments referred to N. donmertoni lack the well-marked grooves terminating anteriorly in foramina on the median attened anterior symphysis; instead, as best shown by NMNZ S.51624, these grooves remain on the side of the symphysal area. These specimens are most similar to mandibles of Strigops in that the anterior rhamphothecal covered zone is at a relatively shallow angle to the plane of the ventral margin of the mandible and more than twice the length as the more caudal zone: in Nestor, the length of the caudal zone exceeds the anterior part. In no fossil specimen is the dorsal margin of the anterior mandible preserved, precluding ascertaining whether the characteristic notch seen in Strigops is present. Examined psittacids such as Platycercus, Prosopeia, Eclectus, and Alisterus have the full length of the symphysis forming a single axial plane at a relatively steep angle to the ventral surface of the mandible, with the rhamphothecal zone extending close to the caudal margin of the symphysis. Cacatuids have highly derived mandibles differing from the fossils in several ways, including having a foreshortened

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JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 31, NO. 5, 2011 Despite the bone having been worn before fossilization and the tubercle for the lig. acrocoraco-procoracoideum being broken off, it is clear the tubercle for lig. acrocoraco-acromion is a prominent bulge extending sternally over the sulcus supracoracoideus and was separated from the more ventral tubercle by a notch, as in Nestor. The facies clavicularis markedly overhangs a large pneumatic foramen extending mainly from the dorsal side of the sulcus supracoracoideus. TarsometatarsusThe partial tarsometatarsus NMNZ S.51398 (Fig. 1N) is nearly twice as big as Nelepsittacus donmertoni (minimum shaft width 4.0 mm vs. 2.4 mm), but is otherwise similar in that the fossa metatarsi I is shallow and entirely on the facies medialis and at right angles to the facies dorsalis. Apart from much greater size, N. daphneleeae differs from N. minimus in the form of the fossa metatarsi I: in the latter species the oor of the fossa is visible in dorsal view, and the ridge lateral of the foramen vasculare distale has a shorter proximal extent, so ends distal to the fossa metatarsi I, rather than overlapping the fossa. This species is only tentatively referred to Nelepsittacus because there is inadequate representation of the tarsometatarsus. However, it shares with previous described Nelepsittacus species a humerus in which the caudal surface of the epicondylus ventralis is wide adjacent the sulcus humerotricipitalis and a coracoid in which the attachment for the lig. acrocoraco-acromion is prominent sternally over the sulcus supracoracoideus and is separated from the more ventral tubercle for the lig. acrocoracoprocoracoideum by a notch, features otherwise seen only in Nestor. A Fourth Psittaciform Species A proximal L scapula NMNZ S.42246, from Croc Site Layer 1, noted by Worthy et al. (2007:g. 14G), represents a larger species than those described above. It is worn, but is provisionally identied as a psittaciform, and if so would be from a bird similar in size to the kea Nestor notabilis. A proximal L ulna NMNZ S.52411 from Bed HH1a (Fig. 2P), with dorsoventral cotylar width of ca. 10 mm, represents a similarly large psittaciform species. It is worn, with both the prominences of the olecranon and tuber. ligamentosa collateralis ventralis lost. It is referred to psittaciforms based on its general similarity to, for example, the ulnae of N. notabilis (Fig. 2R). Specically, it shares an identical cotylar arrangement and form of the tuber. bicipitale ulnae with two distinct ligament attachments distal to the cotyla ventralis in the incisura radialis that are aligned distodorsally towards a prominent crista traversing the cranial facies distoventrally from under the proc. cotyla dorsalis. It differs from N. notabilis by a slightly shallower impressio brachialis and that the more proximal ligament attachment in the incisura radialis undercuts the lip of the cotyla ventralis to a greater extent. DISCUSSION Comparisons of Nelepsittacus Species with Neogene Taxa Australian TaxaThe only described psittaciform fossils from Australia are an early Miocene premaxilla of a Cacatua senus lato species from the RSO Site, Faunal Zone B, Riversleigh World Heritage Area, Queensland (Boles, 1993), and bones of Pliocene age referred to Melopsittacus undulatus (Boles, 1998). Although sparse, these fossils indicate the denite presence of Cacatua in the early Miocene, and support molecular data indicating that most Australasian genera originated before the Oligocene (Wright et al., 2008). Nelepsittacus shows no afnity to cacatuids, Melopsittacus, or any other Australian taxon and so a postOligocene dispersal of this genus from Australia is not supported. European TaxaNelepsittacus minimus can be compared to the ve known crown-group fossil psittaciforms from Europe (Mayr and Gohlrich, 2004; Mayr, 2010a), as four are based on the

anterior dorsal section supporting the rhamphotheca, but this section medially is planar with, or steeper than, the more caudal part of the symphysis. NELEPSITTACUS DAPHNELEEAE, sp. nov. (Figs. 1N, 2KM) HolotypeNMNZ S.50826 (Fig. 2K, L), distal R humerus. DiagnosisHumerus about 24% larger than that of Nelepsittacus donmertoni, differing by (1) the dorsal origin of m. extensor carpi radialis being more prominent cranially; and (2) the impression of brachialis anticus in the fossa m. brachialis being aligned more transversely on the shaft, extending just slightly proximad of the impressio of the palmar branch of m. extensor carpi radialis, so is wider than long. EtymologyAfter Daphne Lee, geologist at Otago University, for her contribution to knowledge of Miocene terrestrial ecosystems in New Zealand. Type LocalityBed HH1b, Manuherikia River section, for details see above. Stratigraphy/Age/FaunaBannockburn Formation, Manuherikia Group, early Miocene (Altonian); 1916 Ma; St Bathans Fauna. Measurements of Holotype (mm)Distal width 10.9, depth through dorsal condyle 6.3. ParatypesNMNZ S.42550, cranial and sternal ends of a R coracoid; NMNZ S.51398 (Fig. 2N), a part R tarsometatarsus preserving the bases of the trochleae, foramen vasculare distale, and shaft. Measurements of Paratypes (mm)NMNZ S.42550, width sternal articulation is minimally 10.2, as it is worn. NMNZ S.51398, minimum shaft width 4.0. Paratype LocalityBoth NMNZ S.42550 and S.51398 are from Bed HH1a, Manuherikia River section, for details see above. Referred Material (Three Specimens)NMNZ S.44210, L scapula, proximal width 8.8 mm, Bed HH1a; NMNZ S.50609 (Fig. 2M), distal L ulna, preserved length 37.1 mm, dorsoventral width cond. dorsalis ulnaris 5.9 mm, maximum distal width 6.6 mm, Bed HH1a; NMNZ S.52026, L quadrate, based on a height of 10.5 mm, this specimen is tentatively referred to this taxon, Bed HH1b, Trench Excavation. Description HumerusThe humerus NMNZ S.50826 (Fig. 2K, L) is a wellpreserved distal fragment, stained black, preserving all psittaciform features described above, except the caudal surface of the epicondylus ventralis, precluding determination of its height above the sulcus humerotricipitalis; however, it was relatively dorsoventrally wide adjacent to the sulcus humerotricipitalis, as in the other smaller Nelepsittacus species. It is bigger than humeri of Alisterus scapularis, but slightly smaller than those of Eolophus roseicapillus. UlnaThe distal ulna NMNZ S. 50609 (Fig. 2M) preserves slightly more than half of total length, with the nutrient foramen 5.7 mm from the break, is stained black, and is slightly worn especially on the tuber. carpale. It is referred to parrots because of morphological similarity, for example, the lack of papillae remigalis ventralis, a stout proximally broad and attened tuber. carpale whose distal end is directed ventrally as a attened facies forming the proximal side of a broad incisura tuber. carpale, a deep sulcus intercondylaris, and the cond. dorsalis ulnaris is on the caudal facies distinctly swollen adjacent to the distal margin between the incisura tendinosa and the ventral side of the condyle. The specimen is referred to this species by its appropriate size, i.e., it is bigger than ulnae of Alisterus, but smaller than those of Eolophus roseicapillus. CoracoidNMNZ S.42550 preserves the cranial and sternal ends of a single R coracoid (see Worthy et al., 2007:g. 14F).

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WORTHY ET AL.MIOCENE PARROTS OF NEW ZEALAND tarsometatarsus and the fth includes a described tarsometatarsus. The oldest fossil parrot is Mogontiacopsitta miocaena Mayr, 2010a, from the late Oligocene/early Miocene of the Mainz Basin, Germany (Mayr, 2010a). It differs from Nelepsittacus by having a much shorter robust tarsometatarsus, with the lateral rim of trochlea metatarsi II relatively much shorter than the medial rim in plantar view (length of the lateral rim greater than half that of the medial rim in Nelepsittacus). The distal tibiotarsi associated with Mogontiacopsitta both differ from Nelepsittacus by the medial tuber. retinaculi m. bularis combining with the distal tuberositas retinaculi extensoris to form a single tubercle that is relatively closer to the condylus lateralis, being separated by a distance much less than the proximodistal height of that condyle. The tarsometatarsus of Archaeopsittacus verreauxi, from the early Miocene of France (Milne-Edwards, 18671871), differs by the presence of a distinct medial vascular foramen plantarly, the shaft not being convex adjacent to the sulcus for the tendon of m. extensor digitorum longus, and the fossa metatarsi I being deeper in dorsal view, and in so far as breakage allows interpreta tion of the hypotarsus (Mayr and Gohlrich, 2004:g. 1J), the canal pII was not enclosed and not medial to fdl. Lastly, the depth of the proximal tarsometatarsus appears similar through the cotylae lateralis et medialis, rather than deeper laterally as in Nelepsittacus. Despite damage limiting comparisons, the tarsometatarsus of Xenopsitta fejfari Mlkovsky, 1998, of early Miocene age, is relatively more robust, has an unusual convex prole to the proximomedial shaft, and lacks a medial vascular foramen. The taxon Pararallus dispar (Milne-Edwards, 18671871 [1869]) from Sansan, middle Miocene, refers to a psittaciform following the selection of a distal left humerus as a lectotype by Cracraft (1973a:33), unfortunately from among the syntypical series of Psittacus lartetianus Milne-Edwards, 1872 (not of Rallus dispar Milne-Edwards, 18671871, as Cracraft thought). Psittacus lartetianus is thus now a junior synonym of Pararallus dispar, and is known by the material listed by Mlkovsky (2002) under the taxon Psittacus lartetianus, which includes a poorly preserved tar sometatarsus. This was gured by Mayr and Gohlich (2004:g. 1E, F) and is distinguished from the St Bathans taxa by the presence of a large medial vascular foramen. Additionally, in each of Mogontiacopsitta miocaena, Archaeopsittacus verreauxi, Xenopsitta fejfari, and Pararallus dispar, the foramen distale is not preceded on the dorsal shaft surface by a long sulcus bound by a distinct ridge laterally as seen in Nelepsittacus. The fth taxon, Bavaripsitta ballmanni Mayr and Gohlich, 2004, was described from a well-preserved tarsometatarsus from the middle Miocene of Germany. It is more slender than Nelepsittacus, has a distinct medial vascular foramen, lacks a convex medial shaft adjacent to the sulcus for the tendon of m. extensor digitorum longus, and the fossa metatarsi I does not cause a notch in the shaft medial prole, nor does it extend onto the facies visible in dorsal view. In addition to the observed morphological differences, it is unlikely that any of these ve taxa would be closely related to fossils from the early Miocene of New Zealand because of their distant geographical origin. North and South American TaxaOnly two New World fossil psittaciforms older than the Pleistocene are described. Conuropsis fratercula Wetmore, 1926, is known only from a humerus of middle Miocene age from Nebraska and its afnities are uncertain despite having been named in an extant genus (Wetmore, 1926; Waterhouse, 2006). The illustration (Wetmore, 1926:g. 6) suggests that this fossil is like all examined psittacids in having caudally the sulcus humerotricipitalis bound ventrally by a proximally narrow proc. Flexorius that abruptly arises caudally above the sulcus, and so is distinguished from Nelepsittacus, in which the proc. exorius is broad and low relative to the sulcus. Biogeographic considerations suggest that it would be rather unlikely that a fossil psittaciform from New Zealand would be related to this bird.

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Described fossil parrots of pre-Quaternary age in South America are restricted to Nandayus vorohuensis Tonni and Noriega, 1996, from the late Pliocene of Argentina (Waterhouse, 2006). There is no reason to think that this fossil, named in an extant genus, tribe Arini, would in any way be related to Nelepsittacus from New Zealand. The Nestorine Lineage These New Zealand fossil taxa differ in several ways from Strigops, so they shed no light on the origin of this New Zealand endemic parrot. As the St Bathans Fauna is the only Tertiary fauna sampling terrestrial faunas from Zealandia, this absence almost certainly does not mean that the Strigops lineage was absent in Zealandia at this time. The taxa Sphenodon, Leiopelma, and moas (Dinornthiformes) have long been considered archaic components of vicariant origin in New Zealand (e.g., Fleming, 1979). However, only Oliver (1930, 1955) considered Strigops and Nestor among such archaic taxa before Cracraft (1973b) promoted the vicariant origin of these and many other taxa from Gondwana. Phylogenetic support for this hypothesis had to wait until the analysis of molecular data, which conclusively showed that Strigops and Nestor were the sister group of all remaining parrots (de Kloet and de Kloet, 2005; Tokita et al., 2007; Wright et al., 2008). This then has led researchers to use the age of the vicariant split of New Zealand from Gondwana (85 or 82 Ma) to calibrate their phylogenies. For example, noting that the fossil record then provided little calibration for the timing of divergences within psittaciforms, Wright et al. (2008) calibrated their phylogeny assuming either 82 Ma (based on the vicariant split of New Zealand from Gondwana) or 52 Ma (based on a hypothesized divergence between modern parrots and fossil forms found in Europe of lower Eocene age) for the basal psittaciform node. However, even their youngest calibration date generated Nestor-Strigops splits older than the fossils we report here. Our interpretation of Nelepsittacus as a member of Nestorinae, rather than a taxon on the stem for Strigops + Nestor, supports Wright et al.s conclusion that these taxa diverged prior to the Miocene, but does not enable renement of their dates. Fleming (1979:90) considered that a proto-kaka was present in the late Miocene and gave rise to kea and kaka in the Pleistocene. Our data show that this ancestor, better described as a proto-Nestor in todays parlance, was at least present in Zealandia by the early Miocene as suggested by some authors (e.g., Oliver, 1930, 1955; Cracraft, 1973b; Diamond and Bond, 1999; Gibbs, 2006). Importantly, a range of features spread across several elements provides strong support for the referral of the suite of Nelepsittacus species to Nestorinae and show that the extant Nestor species represent a lineage that dates back to the early Miocene in Zealandia. These features of Nelepsittacus species include synapomorphies in the coracoid, humerus, tibiotarsus, and tarsometatarsus. On the coracoid, the facies artic. clavicularis has a prominent ventrally located projection, extending over the sulcus m. supracoracoidei to which the lig. acrocoraco- procoracoideum was attached. The attachment of lig. acrocoraco- acromion is rounded and prominent dorsally on the dorsal side of the facies artic. clavicularis. In the smallest species, N. minimus, this prominence is smallest and does not extend over the sulcus m. supracoracoidei, but in the larger N. donmertoni and N. daphneleeae, this prominence is larger, with signicant sternal projection over the sulcus, creating a deep notch between it and the ventral prominence. The presence of prominent tuberosities for each of these ligamental attachment points appears to be a synapomorphy of strigopids, and that the tuberosities are separated by a deep notch is more similar to Nestor than Strigops: the notch is partly obliterated by the large and autapomorphic development of the

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JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 31, NO. 5, 2011 associated with the Plio-Pleistocene glacial interglacial cycles (Newnham et al., 1999; Billups and Schrag, 2002) must have further impacted on the biota. Clearly, it was the more tropical components of the biota that were eliminated (eucalypts, cycads, casuarinas, the diversity in laurels, crocodilians, and collocaliine swiftlets), so the depression in temperature and its associated impact on reducing the complexity of the ora almost certainly winnowed the fauna. In Recent New Zealand faunas, nestorine parrots were associated with Strigops and platycercine Cyanoramphus parakeets. These parakeets are part of a recent radiation in the southwest Pacic (Kearvell et al., 2003), which therefore may have beneted from a reduced diversity of parrots in New Zealand. It seems feasible that, following extreme reduction in forest communities during one of the Pleistocene glacial periods, parrots were reduced to a minimum of a Strigops lineage and a Nestor lineage, perhaps only three species in all on the mainland. Subsequent expanded forests would have then had vacant niches facilitating the colonization of New Zealand by the Cyanoramphus lineage. ACKNOWLEDGMENTS The authors are indebted to the unstinting efforts of many eld assistants in the St Bathans Fauna project that have amassed the fossils between 2001 and 2010. Many people helped over this period, but we are especially grateful to the efforts of J. P. Worthy, S. J. Hand, M. Archer, J. M. T. Nguyen, J. R. Wood, and A. B. Camens who helped in multiple years of excavations to reveal the fossils described here. We thank J.-C. Stahl, NMNZ, for some images. We are particularly thankful for the continued generous support to this project by the landowners A. and E. Johnstone. This research was part of the project DP0770660 funded by the Australian Research Council, and DP1094411 funded by a UNSW 2010 Goldstar award. R.P.S. was funded by a grant from The Brian Mason Technical Fund. We thank W. Boles for comments on a draft of the text and comments by G. Mayr and another referee that helped improve the text. LITERATURE CITED
Ballmann, P. 1969. Die vogel aus de altburdigalen Spaltenfullung von Wintershof (West) bei Eichstatt in Bayern. Zittelliana 1:560. Baumel, J. J., and L. M. Witmer. 1993. Osteologia; pp. 45132 in J. J. Baumel, A. S. King, J. E. Breazile, H. E. Evans, and J. C. Vanden Berge (eds.), Handbook of Avian Anatomy: Nomina Anatomica Avium, Second Edition. Publications of the Nuttall Ornithological Club, 23, Cambridge, Massachusetts. Billups, K., and D. P. Schrag. 2002. Paleotemperatures and ice volume of the past 27 Myr revisited with paired Mg/Ca and 18O/16O measurements on benthic foraminifera. Paleoceanography 17:111. Boles, W. E. 1993. 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: 818. Boles, W. E. 1998. A budgerigar Melopsittacus undulatus from the Pliocene of Riversleigh, North-western Queensland. Emu 98: 3235. Boles, W. E. 2006. The avian fossil record of Australia: an overview; pp. 387411 in J. R. Merrick, M. Archer, G. M. Hickey, and M. S. Y. Lee (eds.), Evolution and Biogeography of Australasian Vertebrates. Auscipub, Sydney. Bonaparte, C. L. 1849 [December]. Conspectus Systematis Ornithologiae. Editio altera reformata additis synonymis Grayanis (a) Classis II Aves. M. Westerman and Fil, Amsterdam. Boon, W. M., C. H. Daugherty, and G. K. Chambers. 2001. The Norfolk Island green parrot and New Caledonian red-crowned parakeet are distinct species. Emu 101:113121. Campbell, H., and G. Hutching. 2007. In Search of Ancient New Zealand. Penguin and GNS Science, Auckland, 239 pp. Christidis, L., and W. E. Boles. 2008. Systematics and Taxonomy of Australian Birds. CSIRO Publishing, Collingwood, Australia, x + 277 pp.

ventral tuberosity in Strigops. The proc. exorius on the humerus is broad and little elevated caudally above the adjacent sulcus humerotricipitalis, as in strigopids, but unlike in all psittacids and cacatuids where it is narrow and more elevated. The tibiotarsus, as in Nestor and unlike all other psittaciforms, has the retinaculum m. bularis medialis et extensorium tibiotarsi separated from the cond. lateralis by a distance greater than, or equal to, the proximodistal length of that condyle (rather than a markedly shorter separation), and the retinaculum m. bularis lateralis is a low crest that lies adjacent to its counterpart. The most diagnostic element for establishing internal psittaciform relationships is the tarsometatarsus. Nelepsittacus shares with Nestor a relatively elongate tarsometatarsus, distal width about a third of total length, a similar pattern of hypotarsal canals, a minute foramen vasculare medialis plantarly, a marked convexity to the medial shaft surface adjacent to the tuber. m. tibialis cranialis, and the synapomorphy of an elongate groove preceding the foramen distale on the dorsal facies. Signicantly, although a phylogenetic analysis of parrots based on osteological characters was outside the scope of this work, our observations based even on a limited few skeletal elements show that osteology will offer a rich potential source of phylogenetic signal in Psittaciformes and that apomorphies to dene the various well-supported molecular-based clades will be forthcoming, as Mayr (2010b) has intimated is possible. The diversity of nestorines in the St Bathans Fauna (minimally three species, plus perhaps one other) was greater than that present in modern New Zealand faunas, in which a maximum of two species are sympatric. But even then, kaka and kea, although both existing in South Island, are largely separated by ecological preferences, with kea mainly in upland, including subalpine, habitats and kaka in forests below about 1000 m (Higgins, 1999). The greater early Miocene diversity in parrots compared to modern faunas is mirrored by pigeons, with Rupephaps taketake and at least one other smaller species coexisting in the St Bathans Fauna (Worthy et al., 2009), when now there is only one pigeon in New Zealand. Both parrots and pigeons are herbivores, feeding on leaves, fruit, and seeds of trees and other plants. The oral communities in the early Miocene environs of paleolake Manuherikia comprised diverse subtropical rainforests including, for example, at least 10 genera of conifer, Arecaceae palms similar to extant nikau Rhopalostylis sapida, at least one cycad, and more than 100 taxa of angiosperms, including 22 species in Lauraceae alone and eight species of Proteaceae (Pole and Douglas, 1998; Pole et al., 2003; Pole, 2007). In the lower Bannockburn Formation contemporaneous with the St Bathans Fauna, the ubiquitous presence of charcoal indicates a seasonally dry climate and the pollen record reveals that Casuarinaceae became dominant, forming a relatively dry Casuarinaceae woodland around the lake, but with nearby mosaics of rainforest and woodlands with palms and Myrtaceae (Metrosideros, Eucalyptus) present (Pole and Douglas, 1998; Pole et al., 2003). This ora was far more complex than any shrubland-forest ora from the Recent period in New Zealand and is undoubtedly what allowed a variety of parrots and pigeons to exist. The subsequent reduction in diversity of nestorine parrots in Zealandia (to just two species at present in mainland New Zealand) mirrors a trend seen across the whole biota. Of the taxa in the St Bathans Fauna, the crocodilians, a turtle, a family of frogs, and many birds (including the families Palaelodidae and Apodidae) became extinct in addition to losses in diversity of pigeons and parrots (Worthy et al., 2007, 2010b, 2011). The fossil record is too incomplete to inform on when these losses occurred, but it is notable that the middle Miocene transition with its marked drop in global temperature followed the deposition of the St Bathans Fauna (Billups and Schrag, 2002; Shevenell et al., 2004). After that, the dramatic oscillations in temperature

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