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An early Miocene diversity of parrots (Aves,Strigopidae, Nestorinae) from New ZealandTrevor H. Worthy a , Alan J. D. Tennyson b & R. Paul Scofield ca School of Biological, Earth and Environmental Sciences, University of New South Wales,New South Wales, 2052, Australiab Museum of New Zealand Te Papa Tongarewa, P.O. Box 467, Wellington, New Zealandc 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

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ARTICLE

AN EARLY MIOCENE DIVERSITY OF PARROTS (AVES, STRIGOPIDAE, NESTORINAE) FROMNEW ZEALAND

TREVOR H. WORTHY,*,1 ALAN J. D. TENNYSON,2 and R. PAUL SCOFIELD3

1School of Biological, Earth and Environmental Sciences, University of New South Wales, New South Wales 2052, Australia,t.worthy@unsw.edu.au;

2Museum of New Zealand Te Papa Tongarewa, P.O. Box 467, Wellington, New Zealand, alant@tepapa.govt.nz;3Canterbury Museum, Rolleston Ave., Christchurch 8001, New Zealand, pscofield@canterburymuseum.com

ABSTRACT—A new genus and three species of parrot (Psittaciformes, Strigopidae, Nestorinae) are described from theearly Miocene (19–16 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. Thesetaxa 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 closerelationship with Strigops, the other endemic genus assumed to have had a long history in New Zealand. With only nestorineparrots represented, the St Bathans Fauna has nothing in common with the Australian psittaciform fauna, in which cacatuidsand a diversity of psittacid genera exist. These data add to the growing body of evidence that the New Zealand terrestrialvertebrate fauna, at a time minimally 3 Ma after the maximal marine inundation of Zealandia in the late Oligocene, washighly endemic, with no close relationship to the closest faunas in Australia. This high degree of endemism strongly suggeststhat the Zealandian terrestrial biota persisted, at least in part, through the Oligocene highstand in sea level.

INTRODUCTION

The diversity of parrots (Aves, Psittaciformes) in Australiais considerable, with approximately 55 indigenous species (in-cluding three recently extinct) in three families and 29 genera(Higgins, 1999; Christidis and Boles, 2008). In contrast, the NewZealand parrot fauna is depauperate, with just nine speciesin three genera in the whole archipelago (Gill et al., 2010).However, these few taxa include the unique kakapo Strigopshabroptilus, which with the genus Nestor, also endemic to theNew Zealand biogeographic region (i.e., New Zealand, NorfolkIsland, and Chatham Island), forms the sister group to all otherpsittaciforms globally (de Kloet and de Kloet, 2005; Tavareset al., 2006; Tokita et al., 2007; Wright et al., 2008). The kakapois the world’s heaviest and only flightless parrot; it is also noc-turnal, 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 existedon the Chatham Islands (Millener, 1999). The remaining NewZealand taxa all form part of a recent radiation of Cyanoramphusspecies, which diversified within the last million years (Boonet al., 2001; Kearvell et al., 2003) and are probably an example ofa recent arrival from the Pacific. The fossil history of psittaciformsin New Zealand has until this study been restricted to late Pleis-tocene and Holocene deposits (Worthy and Holdaway, 2002).

The upper early Miocene (19–16 Ma) Bannockburn Forma-tion, Manuherikia Group, in Central Otago, New Zealand, hasrecently been shown to contain the diverse St Bathans Fauna(Worthy et al., 2007). The fossil beds are lacustrine and weredeposited near a river delta on one side of a paleolake thatextended over some 5600 km2 (Douglas, 1986; Pole et al., 2003).

*Corresponding author.

The avifauna of the St Bathans Fauna is abundant and dominatedby waterfowl (Anseriformes), with a minimum of eight taxa infive genera (Worthy et al., 2007, 2008). The avifauna also com-prises such diverse taxa as moas (Dinornithiformes), a tubenose(Procellariiformes), birds of prey (Accipitriformes), rails (Ral-lidae), an endemic gruiform (Aptornithidae), a gull (Laridae)and other charadriiforms with features of typical waders, aheron (Ardeidae), a palaelodid (Phoenicopteriformes), pigeons(Columbidae), parrots (Psittaciformes), a swiftlet (Apodidae),an owlet-nightjar (Aegothelidae), and passerine birds (Passeri-formes) (Scofield et al., 2010; Tennyson et al., 2010; Worthy et al.,2007, 2009, 2010a, 2010b). Fish, frogs, reptiles, and mammals arealso 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 recordof 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 Mioceneor younger in age (Waterhouse, 2006). Exclusive of Pleistoceneand younger taxa, these crown-group fossil psittaciforms includefive Miocene taxa from Europe, one Miocene species from NorthAmerica, and one from the Pliocene of Argentina (Mayr andGohlich, 2004; Waterhouse, 2006; Mayr, 2010a). Despite a richNeogene 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. fromthe early-middle Miocene (Boles, 1993), and several fossils ofthe extant Melopsittacus undulatus from a Pliocene site (Boles,1998), both sites in the Riversleigh World Heritage Property innorthwest Queensland.

The identity and relationships of the psittaciforms from theearly Miocene St Bathans Fauna, already noted as including atleast three taxa (Worthy et al., 2007), are therefore importantin understanding the evolution of parrots in Australasia and

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the Pacific. If these parrots were recently derived from Papua-Australian faunas by dispersal, as required by the hypothesisof complete inundation of Zealandia (the India-sized continen-tal fragment encompassing New Caledonia and New Zealandand its sub-Antarctic outliers: Campbell and Hutching, 2007;Landis et al., 2008), various Australian genera might be expectedin the New Zealand early Miocene fauna, such as the widely dis-persed cacatuids (Forshaw, 1989). The presence of endemic NewZealand taxa or stem-group taxa for the Australian radiation inthe early Miocene would support either a vicariant origin fornestorine and strigopine parrots in New Zealand, as suggestedby, for example, Oliver (1930, 1955) and Wright et al. (2008) anddating perhaps as recently as ca. 55 million years ago (Gaina et al.,1998; Schellart et al., 2006), or a subsequent, but pre-Oligocenedispersal event. The purpose of this contribution is to describethe parrots from the St Bathans Fauna, make an assessment oftheir relationships, and thus inform the history of New Zealandbiota.

Institutional Abbreviations—AM, Australian Museum, Syd-ney, New South Wales, Australia; CM, Canterbury Museum,Christchurch, New Zealand; NMNZ, Museum of New ZealandTe Papa Tongarewa, Wellington (formerly National Museum ofNew Zealand, Dominion Museum, and Colonial Museum), NewZealand; UNSW, University of New South Wales, Sydney, NewSouth Wales, Australia.

Anatomical Abbreviations—Hypotarsal tendons run in canalsor sulci and are identified as follows: fhl, tendon of m. flexor hal-lucis longus; fdl, m. flexor digitorum longus; pII, m. flexor per-foratus digiti II; ppII, m. flexor perforans et perforatus digiti II;pIII/IV and ppIII, musculi flexores perforati digitorum III et IVand m. flexor perforans digiti III. Common anatomical terms areabbreviated as follows: artic, articularis; cond, condylus; L, left;lig, ligamentum; m, musculus; proc, processus; R, right; tuber, tu-berculum.

METHODS

Names for specific bone landmarks generally follow Baumeland Witmer (1993). In addition, we follow the definitions for fea-tures of the distal tarsometatarsus employed by Mayr (2002): forexample, trochlea accessoria for the plantarly rotated lateral rimof trochlea metatarsi IV. Interpretation of hypotarsus morphol-ogy is after Mayr and Gohlich (2004) and Mayr (2010b). Mea-surements were made with Tesa dial callipers and rounded to0.1 mm.

We follow Gill et al. (2010) in accepting three families amongRecent parrots, Strigopidae Bonaparte, 1849, with NestorinaeBonaparte, 1849, as a subfamily, Cacatuidae Gray, 1840, andPsittacidae Illiger, 1811. However, we note that Gill et al. (2010)attributed Strigopidae to Gray (1848) in error, when it is infact correctly attributed to Bonaparte (1849), wherein Nestori-nae was listed as a subfamily of Strigopidae. On the sheetthat is Bonaparte’s (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.” WithinPsittacidae, we follow the taxonomic nomenclature advocated byChristidis and Boles (2008) and Gill et al. (2010).

Analyses of molecular data have recently produced a stronglysupported topology of relationships within Psittacidae, with thatof Schweizer et al. (2010) corroborating and expanding on pre-vious studies of molecular data (de Kloet and de Kloet, 2005;Tavares et al., 2006; Tokita et al., 2007; Wright et al., 2008). Herewe assess observed morphological structure against the phyloge-netic arrangement of taxa inferred from the topology found bySchweizer et al. (2010) and employ clade names used therein.

Comparative Material

The fossils were compared to the following Recent psittaci-form taxa, with an emphasis on Australasian and southwest Pa-cific species, grouped according to Schweizer et al. (2010). Allare in the Australian Museum collection (and which have the ex-tra prefix AM), except a few from the Canterbury Museum, asfollows:

Strigopidae, Strigopinae: Strigops habroptilus A.1987; CMAv36986 (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. funereusO.54157, O.71387; Callocephalon fimbriatum O.71406;Lophochroa leadbeateri O.70456; Eolophus roseicapillusO.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 macaoO.65776; Amazona ochrocephala S.652, O.71410; Brotogerischiriri O.65100; Myiopsitta monachus O.59846; Pyrrhuramolinae 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; Ge-

offroyus geoffroyi O.59374; Alisterus scapularis O.71194,O.72078; Aprosmictus erythropterus O.60403, O.66369;Polytelis swainsonii O.60062, O.66383; P. anthopeplusO.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 discolorO.68484; Psephotus haematonotus O.58124, O.60422; P.varius O.58524; P. dissimilis O.65325; Prosopeia tabuensisS.1581; Cyanoramphus novaezelandiae O.64715, O.65091;Neopsephotus (= Neophema) bourkii O.66620; Neophemaelegans O.64696; N splendida O.58019; Neophema pulchellaO.54385, O.60918; Pezoporus wallicus O.60945, O.65346.

Loricoloriinae: Agapornis roseicollis O.58461; A. lileanaeO. 57155; Lorius lory or L. hypoinochrous S.708; Char-mosyna placentis S.811; Trichoglossus haematodus O.54381;T. chlorolepidotus O.67628; Glossopsitta concinna O.71416,72391; G. pusilla O.65469; G. porphyrocephala O.70783; Cy-clopsitta diophthalma O.65125, O.71166; Melopsittacus un-dulatus O.64782, O.71326; Chalcopsitta cardinalis S.822.

Fossil Sites

The locations of the fossil sites from which the St BathansFauna has been collected are described by Worthy et al. (2007).Most of the fossils described here derive from a stratigraphic sec-tion exposed on the banks of the Manuherikia River, Otago, NewZealand (figured in Schwarzhans et al., in press). In this section,about 30 m of the base of the Bannockburn Formation is exposedand several fossil-bearing beds have been identified, with thosecontaining parrot bones as follows:

Bed HH1a, 6.88–7.0 m above the base of the Bannock-burn Formation, main quarry at 44.907944◦S 169.858222◦E,Manuherikia River section; ca. 5–10 cm thick sand and cob-ble layer with abundant bone fragments, mud rip-up clastsand stromatolite fragments, and rare fragments of hyriid bi-valves. Fossil Record Number in the archival Fossil RecordFile of the Geological Society of New Zealand H41/f88.

Bed HH1b, 9.5–9.58 m above the base of the Bannock-burn Formation, Trench Excavation, foot of hill 50 macross terrace from river bank at 44.90780◦S 169.85844◦E,

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Manuherikia River section; ca. 10–15 cm thick sand and cob-ble layer with bones abundant, but more worn than thosefrom Bed HH1a. Fossil Record Number is H41/f0103.

Bed HH2b, 21.02–21.31 m above the base of the Bannock-burn Formation, at 44.907861◦S 169.857389◦E, ManuherikiaRiver section; a ca. 3–5 cm thick clay layer with abundantcalcified root casts, common fish bones, rare molluscs, andinfrequent bird bones, overlain by about 17 cm of fine sand.Fossil Record Number is H41/f87.

Bed HH4, 25.63–25.83 m above the base of the BannockburnFormation, and 2.15–2.35 m above the top of a prominentbed of oncolites. Manuherikia River section, 44.907861◦S169.857233◦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 cob-ble layer 3.5 m above the base of the Bannockburn For-mation, in a 3-m cliff on north side of small hill on trueleft of Mata Creek, Otago, 44.889500◦S 169.837833◦E. FossilRecord Number is H41/f84.

Mata Creek, Site 2b, fish bone bed 2 m above an oncolite bedin a bluff about 200 m downstream of Croc Site, 44.890450◦S169.838400◦E. Fossil Record Number is H41/f80.

SYSTEMATIC PALEONTOLOGY

PSITTACIFORMES Wagler, 1830STRIGOPIDAE Bonaparte, 1849 (Kakapo and allied parrots)

NESTORINAE Bonaparte, 1849 (Kaka and Kea)NELEPSITTACUS, gen. nov.

Type Species—Nelepsittacus minimus, sp. nov.Diagnosis—A parrot with a tarsometatarsus with the follow-

ing unique combination of characteristics. (1) A plesiomorphicpattern of hypotarsal canals, where fhl and fdl are well sepa-rated and fully enclosed plantarly, with fdl largest, pII positionedmesad and plantar of fdl and enclosed on its sides but open plan-tarly, and all other tendons run over the plantar surface of the hy-potarsus. (2) The depth of the proximal tarsometatarsus is greaterthrough the cotyla lateralis to the plantar extremity of the cristalateralis hypotarsi than through the cotyla medialis to the tip ofthe crista medialis hypotarsi where it bounds pII. (3) It has asmall foramen vasculare proximale mediale opening plantarly atthe distal end of fdl on the ridge bounding that canal. (4) Thetuberositas m. tibialis cranialis makes a convex protuberance onthe medial profile. (5) It is relatively elongate, with distal widthabout 30% of total length. (6) The trochlea accessoria in distalaspect expands dorsally towards its medial end and is roundedmedially. (7) The foramen vasculare distale is bound on the dor-sal facies by a ridge extending proximal of it, creating a shal-low groove extending proximad of the foramen. (8) The fossametatarsi I is large and deep, and the facies within it is exposedin dorsal view, not just as a notch in the profile. (9) Trochleametatarsi II is grooved distally and plantarly, and its plantar ar-ticular surface indicates a plane of rotation for the articulatingphalanx towards the trochlea accessoria. Character 7 is a synapo-morphy with Nestor alone of living genera, and characters 8 and9 are considered autapomorphies of the new genus.

Etymology—To denote the fossil as a parrot on the ancestrallineage of Nestor. From the Greek mythological figure, Neleus(Greek) father of Nestor and the word psittakos (Greek), a par-rot; masculine nominative.

NELEPSITTACUS MINIMUS, sp. nov.(Figs. 1A–I, L; 2A, B, E, I, J, N, Q)

Holotype—NMNZ S.52404 (Fig. 1A–D, I), a complete Ltarsometatarsus missing only the medial plantar projection

of trochlea metatarsi II. Collected 1 March 2010 by theUNSW/CM/NMNZ expedition.

Diagnosis—As for genus.Etymology—Latin, adjective for smallest, to reflect this is the

smallest species in the genus.Type Locality—Bed HH2b, Manuherikia River section,

Otago, New Zealand: details as above.Stratigraphy/Age/Fauna—Bannockburn Formation, Manu-

herikia Group, early Miocene (Altonian); 19–16 Ma; St BathansFauna.

Measurements of Holotype (mm)—Length 17.8, proximalwidth 4.7, maximum proximal depth 3.6; least shaft width 2.0, dis-tal width 5.5.

Paratypes—NMNZ S.50842 (Fig. 1F–H), a proximal R tar-sometatarsus with complete hypotarsus; NMNZ S.50843 (Fig. 1E,L), a R tarsometatarsus missing the end of the trochlea metatarsiII, the plantar surface of trochlea metatarsi IV, and part of theplantar surface of crista lateralis hypotarsi.

Paratype Locality—Both from Bed HH4, Manuherikia Riversection (details above).

Measurements of Paratypes (mm)—NMNZ S.50842, proximalwidth 4.8, maximum proximal depth 3.4; NMNZ S.50843, length17.5, proximal width 4.7, least shaft width 1.9, preserved distalwidth 5.4.

Referred Material (14 Specimens)—The following specimens,mostly from the Manuherikia River section, were referredto this species rather than the one described below based ontheir 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; NMNZS.43997, distal R; NMNZ S.52223, distal R; Bed HH4: NMNZS.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 shaftR; Bed HH1a: NMNZ S.43994 (Fig. 2E), R; Bed HH1b, TrenchExcavation: NMNZ S.51801, cranial part R. Scapula, Bed HH1a:NMNZ S.43993, R, proximal width 4.9 mm. Tibiotarsus, BedHH1a: NMNZ S.52484 (Fig. 2N), distal R. Mandibles, basedon size, we also tentatively refer to this species two symphysisfragments from Bed HH1a: NMNZ S.52115 and S.52163.

Description and Comparison

Tarsometatarsus—The holotype (Fig. 1A–D, I) is stained lightbrown and was reassembled from three pieces. Its projectingedges (rims of trochleae, cotylar margins) are slightly worn andbreakage means it is missing the medial plantar projection oftrochlea metatarsi II and the medial edge of the crista medialishypotarsi. All salient features are described in the diagnosis. Thetwo paratypic tarsometatarsi reveal some variation in the plantarsurface opposite the sulcus for the tendon of m. extensor digito-rum longus. In the holotype and NMNZ S.50842 (Fig. 1F–H), thisarea of the plantar surface is only slightly convex, but in NMNZS.50843 (Fig. 1E, L), it is marked by a prominent convex surface.In all three available fossil tarsometatarsi, the fhl is not closedover its entire proximodistal length: a similar incomplete plantarclosure was noted for Bavaripsitta ballmanni and Neophema byMayr and Gohlich (2004) and for Psittacula krameri in this study,although G. Mayr (pers. comm., 2011) noted that specimens heexamined in Forschungsinstitut Senckenberg are all completelyclosed, so this character may vary intraspecifically. In Nelepsitta-cus minimus, plantar closure of fhl is incomplete proximally, cre-ating a notch in the proximal side of the bridge closing the canal.All three available tarsometatarsi have damage prohibiting inter-pretation of the structure of the external or medial member ofthe impressiones retinaculorum extensorium; however, that thelateral impression is low suggests that its counterpart will not be

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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; andI, medial view. A–D, I, Nelepsittacus minimus holotype, L, NMNZ S.52404; E, L, paratype, R, NMNZ S.50843; F–H, paratype, proximal R, NMNZS.50842; K, interpretation of proximal end; J, M, N. donmertoni, holotype, R, NMNZ S.52016; and N, N. daphneleeae, paratype, part R, NMNZS.51398. Abbreviations: clh, crista lateralis hypotarsi; fdl, m. flexor digitorum longus; fhl, tendon of m. flexor hallucis longus; fmI, fossa metatarsi I; fvd,foramen vasculare distale; fvm, foramen vasculare mediale; lire, lateral impressiones retinaculum extensorium; mire, medial impressiones retinaculumextensorium; pII, m. flexor perforatus digiti II; ppII, m. flexor perforans et perforatus digiti II; pIII/IV and ppIII, musculi flexores perforati digitorumIII et IV and m. flexor perforans digiti III; sedl, sulcus for tendon of m. extensor digitorum longus; ta, trochlea accessoria; tmtc, tuberositas m. tibialiscranialis; TII, trochlea metatarsi II. Scale bars equal 10 mm.

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FIGURE 2. Appendicular elements of Nelepsittacus species: Coracoids in A, C–E, dorsal view and B, ventral view; humeri in F, H, I, L, caudal viewand G, J, K, cranial view; ulnae in M, ventral view and P–R, cranial view; and N, O, tibiotarsi in anterior view. A, B, Nelepsittacus minimus, NMNZS.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; andQ, 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 NMNZS. 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

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robust and dorsally elevated as it is in many cacatuids or othertaxa with short tarsi. The moderately elongate tarsometatar-sus of N. minimus distinguishes it immediately from all cacatu-ids, the tribe Arini, and most members of Loricoloriinae withinpsittacids.

A hypotarsus with just two enclosed canals, for fhl and fdl, anda well-defined sulcus for pII is shared by N. minimus, strigopids,and cacatuids, but excludes many parrot genera with various apo-morphic modifications to this pattern. All members of Schweizeret al.’s (2010) platycercine group A are characterized by whatMayr (2008) termed a platycercini hypotarsus morphology orhis derived pattern A (Mayr, 2010b), in which the crista lateralishypotarsi is developed as a prominent flange to largely enclosea canal for pIII/IV and ppIII plantarly, and the crista medialishypotarsi is plantarly deepened, with the canal for pII usuallywholly enclosed, and, plantar to it, the canal for ppII is also welldefined or enclosed. Within this group there is variation withlinkages of fdl to pII (e.g., Prosopeia, Cyanoramphus, Northiella,and Lathamus), and occasionally fhl may be partly open to thecanal for pIII/IV and ppIII (e.g., Cyanoramphus). Forming thesister clade to these core platycercines in Schweizer et al.’s (2010)topology, Neophema and Neopsephotus have very small tarsi andfurther differ by lacking the large plantar-medial developmentof the crista lateralis hypotarsi. However, both taxa have adeeper crista medialis hypotarsi than the St Bathans fossils. InNeophema, this results in enclosure of the linked canals fdl, pII,and ppII, but in Neopsephotus, fdl remains discrete and pII isopen plantarly.

The sister group to platycercine group A, the Loricoloriinaeof Mayr (2008), which includes Cyclopsittacini (e.g., Cyclopsitta),Loriini (e.g., Lorius, Charmosyna, Glossopsitta), some Psittac-ulini (e.g., lovebirds, Agapornis, and hanging parrots, Loriculus),and budgerigar (Melopsittacus), are defined by the completeenclosure of all tendons within the hypotarsus, Mayr’s (2010b)derived pattern B. There is variation within this group from anextreme of all canals linked as one complex (Loriini), two canalsoffset plantarly from each other, where fdl + fhl lie dorsally andthe other tendons in a single canal more plantarly (Agapornis andLoriculus), or various other combinations of canal linkage (Mayr,2008). We found that Cyclopsitta had a single complex canal asin Loriini, but differed from genera such as Trichoglossus by thiscanal being partly open plantarly, although Mayr (2008, 2010b)reported that fhl, fdl, and pII were discrete canals. Different os-sification of the intervening septa may relate to ontogenetic age.We prefer to include Cyclopsitta in Mayr’s group characterizedby his ‘derived pattern B’ hypotarsus morphology, because it iscloser to this than the platycercini type where there is no plantar-lateral outgrowth of the crista medialis hypotarsi towards that onthe crista lateralis hypotarsi: in Cyclopsitta it appears a small stepto 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 PsittaculiniGroup A (includes Alisterus, Aprosmictus, Polytelis, Prioniturus,Eclectus, Geoffroyus, and Psittacula). Micropsitta differs fromNelepsittacus minimus by having a hypotarsal structure like thatof Melopsittacus, as noted by Mayr (2008, 2010b). Prioniturushas fusion of fhl and fdl in a single canal, with all deeper tendonspartly enclosed in a sulcus plantarly (Mayr and Gohlich, 2004).

The remaining taxa are similar to N. minimus, with a plantarlylow crista lateralis hypotarsi that lacks a medial projection.Aprosmictus and Polytelis differ from N. minimus and are likesome members of platycercine group A, with fdl, pII, and ppII alllinked and wholly enclosed. Eclectus has the canal for pII whollyenclosed and that for ppII distinct, but open. Alisterus has a canalstructure quite similar to N. minimus, with fdl and fhl discreteand pII open plantarly. However, this genus, as in Eclectus,Psittacula, Aprosmictus, Polytelis, and Prioniturus, differs fromN. minimus in the mediolateral position of pII relative to fdl.In N. minimus, pII is distinctly offset mediad of the lateralmargin of fdl, whereas in all these taxa, pII is positioned entirelyplantar of fdl. In this feature, N. minimus is like Nestor (pIIentirely 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 fdlin Calyptorhynchus), and members of Arini, suggesting that arelatively medial location for pII is plesiomorphic. In Strigops,however, although the external margin of pII is only slightlymedial to fdl, pII broadly overlaps fdl on the lateromedial plane.

All members of Arini are distinguished by a distinct patternof the hypotarsus, in which fdl and fhl are usually the onlyenclosed 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 run-ning over the plantar surface of the hypotarsus in very poorlydefined sulci. Normally fdl and fhl are close together and theircombined width is relatively narrow compared to the width ofthe tarsometatarsus. Characteristically, fdl slightly overlaps fhlon its cranial side and sometimes the septum is not ossified (e.g.,Myiopsitta, Pyrrhura). The single representative of Psittaciniexamined, 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 isalso seen in most cacatuids and is another character distinguish-ing members of this tribe from Nelepsittacus minimus. In Cacatua(galerita, tenuirostris, sanguinea, ducorpsii), Callocephalon fim-briatum, and Calyptorhynchus banksii, canal fdl was close to fhland separated by a narrow osseous septum such that fdl over-lapped fhl dorsally. In other taxa (Nymphicus, Calyptorhynchusfunereus, C. lathami, Eolophus roseicapillus, and Lophochroaleadbeateri), the canals were close but did not overlap each other.Only Probosciger among cacatuids was observed to have well-separated canals.

The greater depth of the proximal tarsometatarsus throughthe cotyla lateralis to the plantar extremity of the crista later-alis hypotarsi compared to that through the cotyla medialis tothe tip of the crista medialis hypotarsi distinguishes Nelepsitta-cus minimus from several of the taxa with a plesiomorphic canalpattern. Strigops has the least well-developed crista lateralis hy-potarsi among extant psittaciforms, and its greatest proximaldepth is through the cotyla medialis to a point beside a partlyenclosed pII. A relatively shallow lateral side of the proximaltarsometatarsus also characterizes all cacatuids (e.g., Nymphi-cus, Probosciger, Cacatua, Calyptorhynchus, Eolophus, and Cal-locephalon), Psittacus, most examined taxa in Psittaculini groupA (e.g., Eclectus, Alisterus, Psittacula, Aprosmictus, and Poly-telis), and many of the platycercine group A taxa.

← 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. fibularis; pbmr, impression of the palmar branch of m. extensor carpi radialis; pcd, proc. cotyla dorsalis; pf, proc. flexorius;pp, proc. procoracoideus; sh, sulcus humerotricipitalis; tbu, tuber. bicipitale ulnae; tc, tuber. carpale; td, tuber. dorsale; tsv, tuber. supracondylareventrale. Note that in A and B the proc. procoracoideus is preserved with some sediment adhering to its ventral side to ensure its retention, but whichmakes it look thicker than it is. Scale bar equals 10 mm.

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The plantar development of the crista lateralis hypotarsi fur-ther distinguishes Nelepsittacus minimus from taxa with a moreplesiomorphic hypotarsal structure. In N. minimus, this crista ex-tends plantarly as a robust ridge, such that in proximal view, itis seen to laterally enclose and so define a sulcus for ppIII/IVand ppIII that is deeper than in all cacatuids. In lateral view, thiscrista is seen to extend proximad with increased plantar depth sothat its most plantar point is proximal to the cotyla lateralis. Inplantar aspect, this crista projects proximolaterad, and is not un-dercut either distally or laterally. In this, it differs markedly fromall cacatuids, including Nymphicus, in which the plantar surfaceof the crista lateralis hypotarsi markedly overhangs, or is under-cut by, the shaft both distally and laterally. Nelepsittacus minimusis, however, like Nestor and Strigops in this respect.

A small medial vascular foramen is shared with Nestor. Alarger, more obvious foramen is seen in Strigops, Psittacus, mostLoricoloriinae, including Melopsittacus, and many members ofPsittaculini group A (including Eclectus, Alisterus, Aprosmictus,and Polytelis). A few platycercines also have a foramen: it may besmall or absent in Platycercus elegans and Psephotus haematono-tus, but is large in Northiella and distinct in Lathamus.

That the tuberositas m. tibialis cranialis causes a marked con-vexity on the medial margin of the shaft distinguishes Nelep-sittacus minimus from all cacatuids and Loricoloriinae, andmany other taxa (including Neophema, Neopsephotus, Eclectus,Prosopeia, Cyanoramphus, Psephotus, Barnardius, and Platycer-cus). The medial profile in Strigops is only slightly convex besidethe tuberositas m. tibialis cranialis. The elongate groove extend-ing proximad from the foramen vasculare distale on the dorsalfacies, formed by the presence of a ridge extending up the shaftfrom the trochlea metatarsi IV, is a feature of Nelepsittacus min-imus observed only in Nestor among examined modern taxa.

The form of the trochlea accessoria is relatively simple inNelepsittacus minimus. In distal view, it expands significantly dor-sally towards its medial end, so partly occluding the furrow sep-arating it from trochlea metatarsi IV, and at its medial end isrounded, not bearing a groove aligned parallel to its dorsal andventral 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 thewhole trochlea accessoria being nearly parallel. Nestor has an au-tapomorphic condition, not shared with the fossil, of the medialend of the trochlea accessoria being greatly expanded dorsallyand deeply grooved on its medial tip giving it a bifid appearance.

Nelepsittacus minimus differs from all psittaciforms examinedby the form of the trochlea metatarsi II, which in plantar viewis seen to be well grooved over its distal end, and which grooveindicates that the plane of rotation of the articulating phalanxis towards the trochlea accessoria. In all other psittaciforms ex-amined, this plane of rotation is towards a point proximal of thetrochlea accessoria.

Nelepsittacus minimus shares with Nestor a relatively elongatetarsometatarsus with a distal width about a third of total length,a similar pattern of hypotarsal canals, a minute foramen vascu-lare medialis plantarly, a marked convexity to the medial shaftsurface adjacent to the tuber. m. tibialis cranialis, and the ap-parent synapomorphy of a groove preceding the foramen distaleon the dorsal facies. It is distinguished from Nestor by three fea-tures, in addition to the form of the trochlea accessoria: (1) thefossa metatarsi I is larger and deeper, and impinges on the medialprofile, extending (autapomorphically) onto the dorsal facies;(2) the crista lateralis hypotarsi is lateromedially thinner in thearea bounding the fhl canal, so that the distal exit of fhl is rela-tively closer to the lateral facies; and (3) the distal exit of canalpII lies ventral to the bounding crista laterad of fdl and is passedon its medial side by a shallow sulcus extending from the plan-tar shaft facies up the medial facies to near the lip of the cotylamedialis. In Nestor, a similar sulcus undercuts the exit of pII, but

does not extend onto the medial facies. In Strigops, there is noevidence of such a sulcus.

In summary, Nelepsittacus minimus shows greatest similarity toNestor (both species), sharing one exclusive (among extant taxa)synapomorphy, a distinct groove extending up the shaft from theforamen vasculare distale, but is distinguished from it by severalfeatures that support its generic distinction. This similarity withNestor and the lack of shared apomorphies with members respec-tively of Loricoloriinae, platycercini group A, Psittaculini groupA (including Micropsitta), Arini, Cacatuidae, and its distinctionfrom Strigops support Nelepsittacus minimus being provisionallyclassified in Nestorinae.

Humerus—The humerus NMNZ S.50171 (Fig. 2I, J) is referredto Nelepsittacus minimus because it exhibits most of the followingpsittaciform features seen in distal humeri (breakage precludesassessing characters 1 and 2, included here for completenessbecause of their relevance to assessing other taxa hereafter): (1)the lack of a proc. supracondylaris dorsalis, with the dorsal originof m. extensor carpi radialis (Vanden Berge and Zweers, 1993:annotation 87) instead of a variable-shaped rugosity on thedorsocranial margin proximal of the cond. dorsalis; (2) a distinctrounded scar, sometimes a tuberosity, immediately ventral tothe dorsal origin of m. extensor carpi radialis and proximal tothe cond. dorsalis, termed tuber. origii m. pronator superficialisby Livezey and Zusi (2006:character 1463), or a ‘tubercle abovedorsal condyle’ by Mayr (2004:fig. 4), or the impressio of the‘palmar’ branch of m. extensor metacarpi radialis as describedfor passerines (Hamon, 1964; Noriega and Chiappe, 1993),but which is the ventral head of origin for m. extensor carpiradialis (Vanden Berge and Zweers, 1993); (3) a shallow fossabrachialis; (4) a large, elevated tuber. supracondylare ventrale;(5) proc. flexorius bears a pair of deep pits for the origin ofm. extensor ulnaris (cranially) and the origin of m. pronatorprofundus (caudally) on the ventral facies, which in cranialaspect presents a notch in the ventral profile due to prominenceof the epicondylus ventralis on the proximal side of these pits;(6) the condyli dorsalis et ventralis and proc. flexorius have equaldistal extent; (7) a narrow, well-defined sulcus scapulotricipitaliscaudally; 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 largerand more numerous taxon described below (Table 1). It pre-serves the condyli dorsalis et ventralis and proc. flexorius and isbroken across the proximal side of tuber. supracondylare ven-trale, but has the above features 3–8. The fossa m. brachialissharply undercuts the tuber. supracondylare ventrale. In disto-caudal view, the proc. flexorius is relatively dorsoventrally wide,little raised above the adjacent sulcus humerotricipitalis, and fea-tures a distinct groove traversing its width, which forms the distalside of the ligamental attachment point. A wide proc. flexorius isshared with Nestor and cacatuids, to the exclusion of other taxa,especially those of Psittaculini group A, where it is very narrow.That the proc. flexorius is little elevated caudally above the ad-jacent sulcus humerotricipitalis distinguishes N. minimus from allpsittacids and cacatuids, and is an apomorphy shared only withNestor and Strigops among compared taxa.

Ulna—These ulnae are distinguished from the followingspecies by their smaller size (Table 2). The proximal ulna (NMNZS.50870; Fig. 2Q) of Nelepsittacus minimus is missing the ole-cranon and is broken through the shaft distal to the impressiobrachialis. The impressio brachialis is deep and undercuts its cau-dal margin. The tuber. bicipitale ulnae (Livezey and Zusi, 2006),or tuber. musculi bicipitis (Ballmann, 1969), is complex with twomain parts: (1) an elongate ridge-like tuberosity starting prox-imally just below (distal of) the cotyla dorsalis and extends c.2.5 mm distally and ventrally; and (2) from below the lip of thecotyla ventralis, another elongate ridge, aligned roughly at rightangles to the first, runs distad and dorsad to end close to the

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TABLE 1. Measurements (mm) for humeri of Nelepsittacus speciesfrom the St Bathans Fauna.

SpeciesCatalognumber

Proximalwidth

Shaftwidth

Distalwidth

maximum

Depthdorsal

condyle

N. minimus S.50171 — — 7.3 4.3N. donmertoni S.42252 — 3.3 — —N. donmertoni S.43995 11.9 3.3 — —N. donmertoni S.43996 — 3.6 7.7 4.5N. donmertoni S.50190 — — 8.3 —N. donmertoni S.50396 — — 8.1 4.8N. donmertoni S.50864 — — 7.9 4.6N. donmertoni S.51422 — 3.4 — —N. daphneleeae S.50826 — — 10.9 6.3

proximal end of the tuberosity. This second ridge has proximaland distal parts that are contiguous, whereas in many psittaci-forms, the parts are separate, as in Platycercus elegans and Psit-tacula. Distal ulnae of parrots vary little and the referred speci-mens exhibit no special features.

Coracoid—Measurements (mm): NMNZ S.52683 (Fig. 2A,B), length from the sternal side of the humeral facet to theend of the proc. acrocoracoideus 6.5; width of humeral facet2.6; NMNZ S.43994 (Fig. 2E), medial length 23.7, width ster-nal facet 6.5. The fossil coracoids referred to Nelepsittacusminimus have the general form of psittaciforms and sharewith all parrots a prominent bulge dorsally between thecranial end of the facies artic. humeralis and the sulcus m.supracoracoidei. The better-preserved NMNZ S.52683, fromthe same bed as the holotype, differs from nearly all coracoidsof extant psittaciforms (slight hollow seen in Psittacula), by thecotyla scapularis having a shallow, but distinct central depres-sion, rather than forming a transverse groove across the axis.The facies artic. clavicularis has a prominent ventrally locatedprojection, extending over the sulcus m. supracoracoidei, towhich the lig. acrocoraco-procoracoideum was attached. Theattachment of lig. acrocoraco-acromion forms a rounded dorsallyprominent bulge on the dorsal side of the facies clavicularis, butdoes not overhang the sulcus m. supracoracoidei. The facies artic.clavicularis markedly overhangs a pneumatic fossa that deeplypenetrates 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 fromthe St Bathans Fauna.

SpeciesCatalognumber

Proximalwidth

Shaftwidth

Distalwidth

maximum

Distalwidthdorsal

condyle

N. minimus S.52223 — 2.4 4.7 3.8N. minimus S.50870 5.1 — — —N. minimus S.42671 — — 4.2 3.6N. minimus S.43997 — — 4.6 3.6N. donmertoni S.42615 — — 5.1+ 4.1+N. donmertoni S.42832 — — 5.1 4.4N. donmertoni S.44211 — — — 4.2N. donmertoni S.51407 — 2.7 5.3 3.9N. donmertoni S.51136 — — 5.2 4.5N. donmertoni S.51915 — — 5.3 4.3N. donmertoni S.51695 — 2.8 5.2+ 4.3N. donmertoni S.51501 — — 5.2 4.5N. donmertoni S.51317 — — 5.2 4.3N. donmertoni S.52455 — 2.9 5.2 4.1N. daphneleeae S.50609 — 3.8 6.6 5.9+

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

proc. procoracoideus is complete on NMNZ S.52683 and showsthat it extends craniomediad from the shaft a distance equivalentto 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 strigopidswith a dorsally prominent bulge for the lig. acrocoraco-acromion,and so differ from other psittaciforms, which lack such a dorsalprominence. However, they differ from coracoids of strigopids,because the same tuberosity for the lig. acrocoraco-acromion isnot prominent sternally over the sulcus m. supracoracoidei. Thislack of a sternally directed prominence may relate to the smallsize 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 hav-ing greater pneumaticity of the proc. acrocoracoideus: in Nestor,pneumatic foramina are usually restricted to small area under thedorsal part of the facies artic. clavicularis; in Strigops, a small areais variably present under the ventral process extending from thefacies artic. clavicularis. Coracoids of N. minimus also differ fromstrigopids in the form of the proc. procoracoideus. Strigops of-ten has the proc. procoracoideus extending well cranial to thecotyla scapulae, and occasionally it is linked to the proc. acroco-racoideus via the ventral process on the facies artic. clavicularis.Nestor is characterized by a reduced proc. procoracoideus, suchthat in N. meridionalis it barely protrudes from the shaft and inN. notabilis, although extending mediad, it lacks any cranial pro-jection. That the facies clavicularis markedly overhangs the sul-cus supracoracoideus distinguishes Nelepsittacus from membersof Arini, in which the overhang is slight.

Tibiotarsus—The tibiotarsus NMNZ S.52484 (Fig. 2N) ofNelepsittacus minimus is worn, and with a distal width of 4.5 mm,is of appropriate size for the holotype tarsometatarsus. Althoughworn, it displays typical psittaciform features as follows: distalend broad, caudally flattened, and craniocaudally shallow; dis-tal condyli of similar size, their proximodistal length about halfof the distal width, and well separated from each other; a shortpons supratendineus; and the distal or medial tuber. retinaculi m.fibularis is confluent with, or occupies the same crista as, the dis-tal tuberositas retinaculi extensoris. The specimen presents thefollowing features: the tuberculi retinaculi m. fibularis medialiset retinaculi extensoris is an elongate, very robust, and craniallyprominent tuberosity; the tuber. retinaculi m. fibularis lateralisforms a low elongate crest on the lateral shaft margin adjacentto its counterpart and separated from it by a flat groove alignedwith the shaft; these tuberculi are separated from the cond. later-alis by a distance slightly greater than the proximodistal length ofthe cond. lateralis; the sulcus extensorius is deep adjacent to thesetuberculi and lies entirely within the medial half of the shaft; thepons supratendineus was probably completely ossified, with ero-sion breaking it in NMNZ S.52484; the incisura intercondylaris isbroad and secondarily excavated into the cond. lateralis; and thecond. lateralis extends slightly distal to the cond. medialis.

Nelepsittacus minimus is similar to Nestor and unlike all otherpsittaciforms in having the tuberculi retinaculi m. fibularis me-dialis et retinaculi extensoris separated from the cond. lateralisby a distance greater than or equal to the proximodistal lengthof that condyle: in all other psittaciforms, the separation is muchless than the length of the condyle. In N. minimus, the tuber.retinaculi m. fibularis lateralis forms a low crest on the lateralmargin adjacent to the tuber. retinaculi m. fibularis medialis ettuberositas retinaculi extensoris, as seen in Strigops, Nestor, andmany psittacids. However, cacatuids (except Nymphicus) aredistinguished by the tuber. retinaculi m. fibularis lateralis beingoffset distally from its medial counterpart and enlarged, withgreatest development seen in Cacatua galerita, C. ducorpsii, andEolophus roseicapillus. In the last two taxa, the crest is veryprominent laterally and extends distad onto the lateral faciesof the cond. lateralis. Similarly, Micropsitta and all members of

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Loricoloriinae, including Melopsitticus, have a distally displacedand enlarged tuber. retinaculi m. fibularis lateralis developed intoa large crest. Micropsitta and all lories and kin are floral feeders,and often climb about upside down while feeding. Eolophus,like many cacatuids, is primarily a ground feeder, foraging onseeds; however, it likes to play and while doing so frequentlyhangs about upside down, and this species also performs a ‘raindance’ involving it hanging upside down (Higgins, 1999). Cacatuagalerita also spends considerable time hanging upside downwhile foraging. Cacatua ducorpsii is one of the tropical species ofCacatua and is primarily a canopy feeder (Forshaw, 1989), thus islikely to spend considerable time clambering about upside downon terminal branches. Thus we interpret the development of thetuber. retinaculi m. fibularis lateralis into a large crest as a homo-plasy related to, and facilitating, the arboreal agility of these taxa.

The likely presence of a completely ossified pons supra-tendineus (shown for N. donmertoni below), shared with Nestor,distinguishes it from taxa in which the osseous bridge is incom-plete, including Strigops, several genera within Arini (we noteMayr’s [2010b] survey indicates several taxa that have a bridge),the cacatuids Probosciger, Cacatua, Callocephalon, Eolophus,and usually in Calyptorhynchus (but ossified in C. funereus AMO.54157, O.71387), but not Nymphicus. The medial locationof the sulcus extensorius in Nelepsittacus distinguishes it fromNestor and Strigops, wherein the sulcus is centred on the shaft.

Mandible—The two mandible fragments tentatively referredto N. minimus, based on smaller size than those described here-after, reveal that the symphysal zone is dorsally concave alongthe axial plane in the part bearing the rhamphotheca. Dorsally,the caudal part of the symphysal zone has a rounded ridge run-ning parallel to the caudal margin of the mandible that separatesthe rhamphotheca from the caudal edge of the mandible. Imme-diately anterior to this ridge and in a median position is a shallowfossa with three foramina entering the corpus. Ventrally, the sur-face is crenulated and on each side a marked groove passes fromthe caudal end anteriorly along the sides and onto the flattenedventral symphysis to end in a foramen.

NELEPSITTACUS DONMERTONI, sp. nov.(Figs. 1J, M; 2C, D, F–H, O)

Holotype—NMNZ S.52016 (Fig. 1J, M), a R tarsometatarsusmissing the cotyla lateralis, most of the hypotarsus, the trochleametatarsi II, and the end of the trochlea accessoria. Collected be-tween 8 and 16 January 2008, by the UNSW/CM/NMNZ expedi-tion.

Diagnosis—Tarsometatarsus larger (22% longer) than Nelep-sittacus minimus, plantar opening of medial vascular foramenlarger, sulcus for the tendon of m. extensor digitorum longus shal-lower as it passes the impressiones retinaculi extensorii medialis.

Etymology—For the late Don Merton (1939–2011), in recogni-tion of his major contributions over many years in the conserva-tion of New Zealand birds, and in particular of kakapo Strigopshabroptilus.

Type Locality—Bed HH1b, Manuherikia River section,Otago, New Zealand (details above).

Stratigraphy/Age/Fauna—Bannockburn Formation, Manu-herikia Group, early Miocene (Altonian); 19–16 Ma; St BathansFauna.

Measurements of Holotype (mm)—Length (actual) 21.8, esti-mated total length 22.0, least shaft width 2.4.

Paratypes—NMNZ S.50261, a proximal end and shaft of R tar-sometatarsus preserving fdl and fhl, but more plantar parts of hy-potarsus lost; NMNZ S.50311, a R tarsometatarsus part proximalend and shaft, with part cotyla medialis and dorsal parts fdl andfhl; NMNZ S.52049, part L tarsometatarsus, shaft, and trochleaaccessoria.

Paratype Locality—NMNZ S.50261 and S.50311 are from BedHH1a; NMNZ S.52049 is from Bed HH1b Trench Excavation inthe 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:fig. 14A, D); NMNZS.43995 (Fig. 2H), proximal R; NMNZ S.43996, distal L; NMNZS.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; NMNZS.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 width3.7; NMNZ S.52622, distal R, distal width 3.8; NMNZ S.52623,distal L, distal width ca. 3.6. Femora: NMNZ S.42723, prox-imal R, proximal width 5.4; NMNZ S.44310, distal L femur,distal width 5.9. Tibiotarsi: NMNZ S.42784 (Fig. 2O), distalR, preserved distal width 5.7 (estimated 6.4); NMNZ S.42853,worn distal L; NMNZ S.50129, distal L, preserved distal width5.6; NMNZ S.50301, shaft L; NMNZ S.52226, distal R. Tar-sometatarsi: NMNZ S.42614, trochlea metatarsi III of R; NMNZS.42856, trochlea III and part of distal end L; NMNZ S.44309,proximal R, minimum shaft width 2.3; NMNZ S.51720, fragmentof a R, with trochlea metatarsi IV and trochlea accessoria. Cora-coids: NMNZ S.40458 sternal end L; NMNZ S.42663, sternal endR, width sternal facet 6.7; NMNZ S.42855, sternal end R, widthsternal facet 7.0 (Worthy et al., 2007:fig. 14J); NMNZ S.50277,imperfect L; NMNZ S.51409, cranial part L. Scapulae: NMNZS.42833, L, proximal width 6.2 (Worthy et al., 2007:fig. 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; NMNZS.51803 (Fig. 2C), cranial part L, length from the sternal side ofthe humeral facet to the end of the proc. acrocoracoideus 6.8,width of humeral facet 2.7; NMNZ S.51900, cranial part L, lengthfrom the sternal side of the humeral facet to the end of the proc.acrocoracoideus 7.9; width of humeral facet 2.7. Scapula: NMNZS.51625, L, proximal width 5.8. Tibiotarsus: NMNZ S.51972, dis-tal part shaft R. Mandible: Based on size, we tentatively referthree 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:fig. 14B, E). Car-pometacarpi: NMNZ S.42409, proximal L, proximal width 5.1+;NMNZ S.42480, R, length 24.7.

Description

Tarsometatarsus—The holotype NMNZ S.52016 (Fig. 1J, M) isstained black and has been reassembled from two fragments suchthat the trochlea metatarsi III is not correctly oriented, being di-rected too much in line with the shaft rather than with a medialinclination. Apart from size and the other features listed in thediagnosis, Nelepsittacus donmertoni differs from N. minimus inhaving a shallower fossa metatarsi I, which is entirely on the fa-cies medialis and at right angles to the facies dorsalis, so the floorof the fossa is not visible in dorsal view. As in N. minimus, theridge lateral of the foramen vasculare distale extends proximadto proximodistally overlap the fossa metatarsi I.

Humerus—The fossil humeri attributed to Nelepsittacus don-mertoni, as best exemplified by NMNZ S.50396 (Fig. 2F, G) andS.50864, exhibit all the above listed psittaciform features and areapproximately the same size as Platycercus elegans (Table 1). The

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origin of the dorsal branch of m. extensor carpi radialis straddlesthe dorsal margin and is not prominent cranially and narrowlyseparated from the more ventrally located impressio of the pal-mar branch of m. extensor carpi radialis, which is also not promi-nent. This latter is separated from the cond. dorsalis by an arcedsulcus whose curve is proximally convex. The fossa m. brachialisis flat, with the impression of brachialis anticus within it extend-ing to the ventral margin and abutting the tuber. supracondylareventrale in a gentle curve (rather than abruptly), and having anelliptical shape aligned along the shaft. As in most psittaciforms,but unlike Nestor, these fossils exhibit a shaft that is not markedlyconvex cranially at the proximal end of the fossa m. brachialis andthe pits on the proc. flexorius are distinct and clearly divided by amedium ridge.

The flat fossa m. brachialis distinguishes Nelepsittacus donmer-toni from all cacatuids, in which taxa it is deeper than in allpsittacids. Strigops and Nestor species share with N. donmertoni avery shallow fossa m. brachialis, but Nestor differs with greater in-flation/convexity of the cranial shaft facies at the level of the fossathan observed in the fossil and other psittaciforms. Nestor humeridiffer more trenchantly from that of N. donmertoni in that on theproc. flexorius, the pits for the origins of m. extensor ulnaris andm. pronator profundus are less distinctly separated by a ridge; thepits are clearly separated in Strigops, cacatuids, and all psittacids.

The proximal humerus is best exemplified by NMNZ S.43995(Fig. 2H), which is complete except for loss of the proximal sideof the tuber. ventrale; however, the entire margin of the fossapneumotricipitalis is preserved. It is similar to humeri of Nestormeridionalis, for example NMNZ OR.28161, in that the capitalshaft ridge connects to the caput humeri just dorsal to a distincttubercle; this ridge bears two prominent ligamental attachmentscars, one overlapping with, and extending distad of, the cristam. supracoracoidei, and a second farther distally and directed to-wards the cranial facies; crista m. supracoracoidei elongate andproximally overhung caudally by the tuber. dorsale; incisura capi-tis with sulcus in distal half closed distally by a low ridge; fossapneumotricipitalis with few pneumatic foramina penetrating itsinternal wall; impressio coracobrachialis deep and well definedventrally, extending distad to a point level with the distal marginof the crista bicipitalis; and the cranial part of the crista deltopec-toralis is worn so its height in indeterminate; however, it extendswell distad of the crista bicipitalis and in its distal half rises fromthe cranial facies at a wide angle, not nearly at a right angle asin Platycercus, for example. Humeri of Strigops differ from thoseof 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, Scapulae—The ulnae (Ta-ble 2) referred to Nelepsittacus donmertoni exhibit no special fea-tures and differ little from those of Platycercus. No significantdifferences in ulnae were found to distinguish nestorines frompsittacids. Similarly, the fossil radii, os carpi ulnare, and scapulawere referred to N. donmertoni based on their size being similarto that of Platycercus elegans; no attempt was made to detect dif-ferences among psittaciforms for these elements. However, theacromion on the referred scapula is less cranially prominent thanit is in Platycercus.

Carpometacarpus—The referred carpometacarpi are rela-tively worn and are referred by their general similarity to those ofparrots. The proximal end is relatively smaller than in Platycer-cus, but has similar proportions to that of Nestor carpometacarpi.Also, as in Nestor, the foramen in the fossa m. infratrochlearis isrelatively large.

Coracoid—The fossil coracoids referred to Nelepsittacus don-mertoni, as best exemplified by NMNZ S.51803 (Fig. 2C),S.51900, and S.51291 (Fig. 2D), are slightly larger than those re-ferred to N. minimus. As in N. minimus, these coracoids have ashallow and distinct central depression in the cotyla scapularis

and a prominent ventrally located projection on the facies ar-tic. clavicularis extending over the sulcus m. supracoracoidei towhich the lig. acrocoraco-procoracoideum was attached. How-ever, they differ from N. minimus in that the attachment of lig.acrocoraco-acromion on the dorsal side of the facies artic. clav-icularis is larger, forming a distinct projection that is separatedby a deep notch from its ventral counterpart. The presence ofprominent tuberosities for each of these ligamental attachmentpoints is an apomorphy shared with strigopids, in contrast toall cacatuids and psittacids in which the dorsal ligament attach-ment point is linear and not developed as a projection both dor-sally and sternally. The structure is most similar to Nestor, whichhas near-equal-sized projections separated by a deep notch forthese ligament attachment points. Strigops has a rounded dor-sal tuberosity with a weak and variable depth notch separating itfrom a markedly more elongate ventral tuberosity, which is occa-sionally linked to the proc. procoracoideus by ossification of thelig. acrocoraco- procoracoideum. The fossils have greater pneu-maticity of the acrocoracoid than does Nestor, in which pneu-maticity is mainly restricted to a small area in the dorsal half ofthe sulcus m. supracoracoidei, under the projection for the attach-ment of lig. acrocoraco-acromion. This pneumatic fossa in cora-coids of N. donmertoni is relatively smaller than in the coracoidsreferred to N. minimus, because it does not undercut the ventralligament attachment point to the same extent.

The fragment NMNZ S.42663 reveals that the sternal end ofthe coracoid has a prominent crista medialis extending craniadof the medial angle a distance equivalent to about half the widthof the sternal end and that the impressio m. sternocoracoidei isdeep, and so is similar to N. minimus (Fig. 2E).

Tibiotarsus—The 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. Thesetibiotarsi are larger than, but otherwise have identical morphol-ogy to, that of Nelepsittacus minimus. The fragment NMNZS.42784 reveals that the medial tuber. retinaculi m. fibularis isdistinct from, and located proximal to, the distal tuberositas reti-naculi extensoris and is at the same level as the tuber. retinaculim. fibularis lateralis and the proximal tuberositas retinaculiextensoris. The fragment NMNZ S.42853 shows that an ossifiedpons supratendineus characterizes the species, with wear openingthe pons in other available specimens.

Mandible—The mandible fragments, apart from being largerthan those referred tentatively to N. minimus, have a differentform. As in Nestor, the median plane through the symphysis re-veals an anterior zone that supports the rhamphotheca, which isanterocaudally planar (not concave) and at a markedly shallowerangle to the caudal section of the symphysis, which rises steeplyfrom the caudal margin. This caudal section also features a shal-low median sulcus and is less than half as long as the anteriorsection. Ventrally, the fragments referred to N. donmertoni lackthe well-marked grooves terminating anteriorly in foramina onthe median flattened anterior symphysis; instead, as best shownby NMNZ S.51624, these grooves remain on the side of the sym-physal area. These specimens are most similar to mandibles ofStrigops in that the anterior rhamphothecal covered zone is ata relatively shallow angle to the plane of the ventral margin ofthe mandible and more than twice the length as the more caudalzone: in Nestor, the length of the caudal zone exceeds the ante-rior part. In no fossil specimen is the dorsal margin of the anteriormandible preserved, precluding ascertaining whether the char-acteristic notch seen in Strigops is present. Examined psittacidssuch as Platycercus, Prosopeia, Eclectus, and Alisterus have thefull length of the symphysis forming a single axial plane at a rel-atively steep angle to the ventral surface of the mandible, withthe rhamphothecal zone extending close to the caudal margin ofthe symphysis. Cacatuids have highly derived mandibles differingfrom the fossils in several ways, including having a foreshortened

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anterior dorsal section supporting the rhamphotheca, but this sec-tion medially is planar with, or steeper than, the more caudal partof the symphysis.

NELEPSITTACUS DAPHNELEEAE, sp. nov.(Figs. 1N, 2K–M)

Holotype—NMNZ S.50826 (Fig. 2K, L), distal R humerus.Diagnosis—Humerus about 24% larger than that of Nelepsit-

tacus donmertoni, differing by (1) the dorsal origin of m. extensorcarpi radialis being more prominent cranially; and (2) the impres-sion of brachialis anticus in the fossa m. brachialis being alignedmore transversely on the shaft, extending just slightly proximadof the impressio of the palmar branch of m. extensor carpi radi-alis, so is wider than long.

Etymology—After Daphne Lee, geologist at Otago Univer-sity, for her contribution to knowledge of Miocene terrestrialecosystems in New Zealand.

Type Locality—Bed HH1b, Manuherikia River section, for de-tails see above.

Stratigraphy/Age/Fauna—Bannockburn Formation, Manu-herikia Group, early Miocene (Altonian); 19–16 Ma; St BathansFauna.

Measurements of Holotype (mm)—Distal width 10.9, depththrough dorsal condyle 6.3.

Paratypes—NMNZ S.42550, cranial and sternal ends of a Rcoracoid; NMNZ S.51398 (Fig. 2N), a part R tarsometatarsus pre-serving the bases of the trochleae, foramen vasculare distale, andshaft.

Measurements of Paratypes (mm)—NMNZ S.42550, widthsternal articulation is minimally 10.2, as it is worn. NMNZS.51398, minimum shaft width 4.0.

Paratype Locality—Both NMNZ S.42550 and S.51398 are fromBed HH1a, Manuherikia River section, for details see above.

Referred Material (Three Specimens)—NMNZ S.44210, Lscapula, proximal width 8.8 mm, Bed HH1a; NMNZ S.50609(Fig. 2M), distal L ulna, preserved length 37.1 mm, dorsoven-tral width cond. dorsalis ulnaris 5.9 mm, maximum distal width6.6 mm, Bed HH1a; NMNZ S.52026, L quadrate, based on aheight of 10.5 mm, this specimen is tentatively referred to thistaxon, Bed HH1b, Trench Excavation.

Description

Humerus—The humerus NMNZ S.50826 (Fig. 2K, L) is a well-preserved distal fragment, stained black, preserving all psittaci-form features described above, except the caudal surface ofthe epicondylus ventralis, precluding determination of its heightabove the sulcus humerotricipitalis; however, it was relativelydorsoventrally wide adjacent to the sulcus humerotricipitalis, asin the other smaller Nelepsittacus species. It is bigger than humeriof Alisterus scapularis, but slightly smaller than those of Eolophusroseicapillus.

Ulna—The distal ulna NMNZ S. 50609 (Fig. 2M) preservesslightly more than half of total length, with the nutrient foramen5.7 mm from the break, is stained black, and is slightly worn es-pecially on the tuber. carpale. It is referred to parrots becauseof morphological similarity, for example, the lack of papillaeremigalis ventralis, a stout proximally broad and flattened tuber.carpale whose distal end is directed ventrally as a flattened faciesforming the proximal side of a broad incisura tuber. carpale, adeep sulcus intercondylaris, and the cond. dorsalis ulnaris is onthe caudal facies distinctly swollen adjacent to the distal mar-gin between the incisura tendinosa and the ventral side of thecondyle. The specimen is referred to this species by its appropri-ate size, i.e., it is bigger than ulnae of Alisterus, but smaller thanthose of Eolophus roseicapillus.

Coracoid—NMNZ S.42550 preserves the cranial and sternalends of a single R coracoid (see Worthy et al., 2007:fig. 14F).

Despite the bone having been worn before fossilization and thetubercle for the lig. acrocoraco-procoracoideum being brokenoff, it is clear the tubercle for lig. acrocoraco-acromion is aprominent bulge extending sternally over the sulcus supracora-coideus and was separated from the more ventral tubercle by anotch, as in Nestor. The facies clavicularis markedly overhangs alarge pneumatic foramen extending mainly from the dorsal sideof the sulcus supracoracoideus.

Tarsometatarsus—The partial tarsometatarsus NMNZ S.51398(Fig. 1N) is nearly twice as big as Nelepsittacus donmertoni (min-imum shaft width 4.0 mm vs. 2.4 mm), but is otherwise similar inthat the fossa metatarsi I is shallow and entirely on the facies me-dialis and at right angles to the facies dorsalis. Apart from muchgreater size, N. daphneleeae differs from N. minimus in the formof the fossa metatarsi I: in the latter species the floor of the fossais visible in dorsal view, and the ridge lateral of the foramen vas-culare distale has a shorter proximal extent, so ends distal to thefossa metatarsi I, rather than overlapping the fossa.

This species is only tentatively referred to Nelepsittacus be-cause there is inadequate representation of the tarsometatarsus.However, it shares with previous described Nelepsittacus speciesa humerus in which the caudal surface of the epicondylus ven-tralis is wide adjacent the sulcus humerotricipitalis and a cora-coid in which the attachment for the lig. acrocoraco-acromion isprominent sternally over the sulcus supracoracoideus and is sep-arated from the more ventral tubercle for the lig. acrocoraco-procoracoideum by a notch, features otherwise seen only inNestor.

A Fourth Psittaciform Species

A proximal L scapula NMNZ S.42246, from Croc Site Layer 1,noted by Worthy et al. (2007:fig. 14G), represents a larger speciesthan those described above. It is worn, but is provisionally identi-fied as a psittaciform, and if so would be from a bird similar in sizeto the kea Nestor notabilis. A proximal L ulna NMNZ S.52411from Bed HH1a (Fig. 2P), with dorsoventral cotylar width ofca. 10 mm, represents a similarly large psittaciform species. It isworn, with both the prominences of the olecranon and tuber. lig-amentosa collateralis ventralis lost. It is referred to psittaciformsbased on its general similarity to, for example, the ulnae of N.notabilis (Fig. 2R). Specifically, it shares an identical cotylar ar-rangement and form of the tuber. bicipitale ulnae with two dis-tinct ligament attachments distal to the cotyla ventralis in the in-cisura radialis that are aligned distodorsally towards a prominentcrista traversing the cranial facies distoventrally from under theproc. cotyla dorsalis. It differs from N. notabilis by a slightly shal-lower impressio brachialis and that the more proximal ligamentattachment in the incisura radialis undercuts the lip of the cotylaventralis to a greater extent.

DISCUSSION

Comparisons of Nelepsittacus Species with Neogene Taxa

Australian Taxa—The only described psittaciform fossils fromAustralia are an early Miocene premaxilla of a Cacatua senus latospecies from the RSO Site, Faunal Zone B, Riversleigh WorldHeritage Area, Queensland (Boles, 1993), and bones of Plioceneage referred to Melopsittacus undulatus (Boles, 1998). Althoughsparse, these fossils indicate the definite presence of Cacatuain the early Miocene, and support molecular data indicatingthat most Australasian genera originated before the Oligocene(Wright et al., 2008). Nelepsittacus shows no affinity to cacatu-ids, Melopsittacus, or any other Australian taxon and so a post-Oligocene dispersal of this genus from Australia is not supported.

European Taxa—Nelepsittacus minimus can be compared tothe five known crown-group fossil psittaciforms from Europe(Mayr and Gohlrich, 2004; Mayr, 2010a), as four are based on the

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tarsometatarsus and the fifth includes a described tarsometatar-sus. 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 hav-ing a much shorter robust tarsometatarsus, with the lateral rimof trochlea metatarsi II relatively much shorter than the me-dial rim in plantar view (length of the lateral rim greater thanhalf that of the medial rim in Nelepsittacus). The distal tibiotarsiassociated with Mogontiacopsitta both differ from Nelepsittacusby the medial tuber. retinaculi m. fibularis combining with thedistal tuberositas retinaculi extensoris to form a single tuberclethat is relatively closer to the condylus lateralis, being separatedby a distance much less than the proximodistal height of thatcondyle. The tarsometatarsus of Archaeopsittacus verreauxi, fromthe early Miocene of France (Milne-Edwards, 1867–1871), differsby the presence of a distinct medial vascular foramen plantarly,the shaft not being convex adjacent to the sulcus for the tendonof m. extensor digitorum longus, and the fossa metatarsi I beingdeeper in dorsal view, and in so far as breakage allows interpreta-tion of the hypotarsus (Mayr and Gohlrich, 2004:fig. 1J), the canalpII was not enclosed and not medial to fdl. Lastly, the depth ofthe proximal tarsometatarsus appears similar through the coty-lae lateralis et medialis, rather than deeper laterally as in Nelep-sittacus. Despite damage limiting comparisons, the tarsometatar-sus of Xenopsitta fejfari Mlıkovsky, 1998, of early Miocene age,is relatively more robust, has an unusual convex profile to theproximomedial shaft, and lacks a medial vascular foramen. Thetaxon Pararallus dispar (Milne-Edwards, 1867–1871 [1869]) fromSansan, middle Miocene, refers to a psittaciform following theselection of a distal left humerus as a lectotype by Cracraft(1973a:33), unfortunately from among the syntypical series ofPsittacus lartetianus Milne-Edwards, 1872 (not of Rallus disparMilne-Edwards, 1867–1871, as Cracraft thought). Psittacus larte-tianus is thus now a junior synonym of Pararallus dispar, andis known by the material listed by Mlıkovsky (2002) under thetaxon Psittacus lartetianus, which includes a poorly preserved tar-sometatarsus. This was figured by Mayr and Gohlich (2004:fig.1E, F) and is distinguished from the St Bathans taxa by the pres-ence of a large medial vascular foramen. Additionally, in each ofMogontiacopsitta miocaena, Archaeopsittacus verreauxi, Xenop-sitta fejfari, and Pararallus dispar, the foramen distale is not pre-ceded on the dorsal shaft surface by a long sulcus bound by adistinct ridge laterally as seen in Nelepsittacus.

The fifth taxon, Bavaripsitta ballmanni Mayr and Gohlich,2004, was described from a well-preserved tarsometatarsus fromthe middle Miocene of Germany. It is more slender than Nelep-sittacus, has a distinct medial vascular foramen, lacks a convexmedial shaft adjacent to the sulcus for the tendon of m. extensordigitorum longus, and the fossa metatarsi I does not cause a notchin the shaft medial profile, nor does it extend onto the facies visi-ble in dorsal view. In addition to the observed morphological dif-ferences, it is unlikely that any of these five taxa would be closelyrelated to fossils from the early Miocene of New Zealand becauseof their distant geographical origin.

North and South American Taxa—Only two New World fossilpsittaciforms older than the Pleistocene are described. Conurop-sis fratercula Wetmore, 1926, is known only from a humerus ofmiddle Miocene age from Nebraska and its affinities are uncer-tain despite having been named in an extant genus (Wetmore,1926; Waterhouse, 2006). The illustration (Wetmore, 1926:fig. 6)suggests that this fossil is like all examined psittacids in havingcaudally the sulcus humerotricipitalis bound ventrally by a proxi-mally narrow proc. Flexorius that abruptly arises caudally abovethe sulcus, and so is distinguished from Nelepsittacus, in whichthe proc. flexorius is broad and low relative to the sulcus. Biogeo-graphic considerations suggest that it would be rather unlikelythat a fossil psittaciform from New Zealand would be related tothis bird.

Described fossil parrots of pre-Quaternary age in South Amer-ica 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 extantgenus, tribe Arini, would in any way be related to Nelepsittacusfrom New Zealand.

The Nestorine Lineage

These New Zealand fossil taxa differ in several ways fromStrigops, so they shed no light on the origin of this New Zealandendemic parrot. As the St Bathans Fauna is the only Tertiaryfauna sampling terrestrial faunas from Zealandia, this absence al-most certainly does not mean that the Strigops lineage was absentin Zealandia at this time.

The taxa Sphenodon, Leiopelma, and moas (Dinornthiformes)have long been considered archaic components of vicariantorigin in New Zealand (e.g., Fleming, 1979). However, onlyOliver (1930, 1955) considered Strigops and Nestor among sucharchaic taxa before Cracraft (1973b) promoted the vicariantorigin of these and many other taxa from Gondwana. Phyloge-netic support for this hypothesis had to wait until the analysisof molecular data, which conclusively showed that Strigops andNestor were the sister group of all remaining parrots (de Kloetand de Kloet, 2005; Tokita et al., 2007; Wright et al., 2008). Thisthen has led researchers to use the age of the vicariant split ofNew Zealand from Gondwana (85 or 82 Ma) to calibrate theirphylogenies. For example, noting that the fossil record thenprovided little calibration for the timing of divergences withinpsittaciforms, Wright et al. (2008) calibrated their phylogenyassuming either 82 Ma (based on the vicariant split of NewZealand from Gondwana) or 52 Ma (based on a hypothesizeddivergence between modern parrots and fossil forms foundin Europe of lower Eocene age) for the basal psittaciformnode. However, even their youngest calibration date generatedNestor-Strigops splits older than the fossils we report here. Ourinterpretation of Nelepsittacus as a member of Nestorinae, ratherthan a taxon on the stem for Strigops + Nestor, supports Wrightet al.’s conclusion that these taxa diverged prior to the Miocene,but does not enable refinement of their dates. Fleming (1979:90)considered that a ‘proto-kaka’ was present in the late Mioceneand gave rise to kea and kaka in the Pleistocene. Our data showthat this ancestor, better described as a proto-Nestor in today’sparlance, was at least present in Zealandia by the early Mioceneas 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 ele-ments provides strong support for the referral of the suite ofNelepsittacus species to Nestorinae and show that the extantNestor species represent a lineage that dates back to the earlyMiocene in Zealandia. These features of Nelepsittacus species in-clude synapomorphies in the coracoid, humerus, tibiotarsus, andtarsometatarsus.

On the coracoid, the facies artic. clavicularis has a prominentventrally located projection, extending over the sulcus m. supra-coracoidei to which the lig. acrocoraco- procoracoideum wasattached. The attachment of lig. acrocoraco- acromion is roundedand prominent dorsally on the dorsal side of the facies artic.clavicularis. In the smallest species, N. minimus, this prominenceis smallest and does not extend over the sulcus m. supracora-coidei, but in the larger N. donmertoni and N. daphneleeae, thisprominence is larger, with significant sternal projection over thesulcus, creating a deep notch between it and the ventral promi-nence. The presence of prominent tuberosities for each of theseligamental attachment points appears to be a synapomorphyof strigopids, and that the tuberosities are separated by a deepnotch is more similar to Nestor than Strigops: the notch is partlyobliterated by the large and autapomorphic development of the

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ventral tuberosity in Strigops. The proc. flexorius on the humerusis broad and little elevated caudally above the adjacent sulcushumerotricipitalis, as in strigopids, but unlike in all psittacids andcacatuids where it is narrow and more elevated.

The tibiotarsus, as in Nestor and unlike all other psittaciforms,has the retinaculum m. fibularis medialis et extensorium tibiotarsiseparated from the cond. lateralis by a distance greater than, orequal to, the proximodistal length of that condyle (rather thana markedly shorter separation), and the retinaculum m. fibularislateralis is a low crest that lies adjacent to its counterpart. Themost diagnostic element for establishing internal psittaciformrelationships is the tarsometatarsus. Nelepsittacus shares withNestor a relatively elongate tarsometatarsus, distal width abouta third of total length, a similar pattern of hypotarsal canals, aminute foramen vasculare medialis plantarly, a marked convex-ity to the medial shaft surface adjacent to the tuber. m. tibialiscranialis, and the synapomorphy of an elongate groove preced-ing the foramen distale on the dorsal facies.

Significantly, although a phylogenetic analysis of parrots basedon osteological characters was outside the scope of this work, ourobservations based even on a limited few skeletal elements showthat osteology will offer a rich potential source of phylogeneticsignal in Psittaciformes and that apomorphies to define the var-ious well-supported molecular-based clades will be forthcoming,as Mayr (2010b) has intimated is possible.

The diversity of nestorines in the St Bathans Fauna (mini-mally three species, plus perhaps one other) was greater than thatpresent in modern New Zealand faunas, in which a maximum oftwo species are sympatric. But even then, kaka and kea, althoughboth existing in South Island, are largely separated by ecologi-cal 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 mod-ern faunas is mirrored by pigeons, with Rupephaps taketake andat least one other smaller species coexisting in the St BathansFauna (Worthy et al., 2009), when now there is only one pigeonin New Zealand.

Both parrots and pigeons are herbivores, feeding on leaves,fruit, and seeds of trees and other plants. The floral commu-nities in the early Miocene environs of paleolake Manuherikiacomprised diverse subtropical rainforests including, for example,at least 10 genera of conifer, Arecaceae palms similar to extantnikau Rhopalostylis sapida, at least one cycad, and more than 100taxa of angiosperms, including 22 species in Lauraceae alone andeight species of Proteaceae (Pole and Douglas, 1998; Pole et al.,2003; Pole, 2007). In the lower Bannockburn Formation con-temporaneous with the St Bathans Fauna, the ubiquitous pres-ence of charcoal indicates a seasonally dry climate and the pollenrecord reveals that Casuarinaceae became dominant, forming arelatively dry Casuarinaceae woodland around the lake, but withnearby mosaics of rainforest and woodlands with palms and Myr-taceae (Metrosideros, Eucalyptus) present (Pole and Douglas,1998; Pole et al., 2003). This flora was far more complex than anyshrubland-forest flora from the Recent period in New Zealandand is undoubtedly what allowed a variety of parrots and pigeonsto exist.

The subsequent reduction in diversity of nestorine parrotsin Zealandia (to just two species at present in mainland NewZealand) mirrors a trend seen across the whole biota. Of thetaxa in the St Bathans Fauna, the crocodilians, a turtle, a familyof frogs, and many birds (including the families Palaelodidaeand Apodidae) became extinct in addition to losses in diversityof pigeons and parrots (Worthy et al., 2007, 2010b, 2011). Thefossil record is too incomplete to inform on when these losses oc-curred, but it is notable that the middle Miocene transition withits marked drop in global temperature followed the depositionof the St Bathans Fauna (Billups and Schrag, 2002; Shevenellet al., 2004). After that, the dramatic oscillations in temperature

associated with the Plio-Pleistocene glacial interglacial cycles(Newnham et al., 1999; Billups and Schrag, 2002) must havefurther impacted on the biota. Clearly, it was the more tropicalcomponents of the biota that were eliminated (eucalypts, cycads,casuarinas, the diversity in laurels, crocodilians, and collocaliineswiftlets), so the depression in temperature and its associatedimpact on reducing the complexity of the flora almost certainlywinnowed the fauna.

In Recent New Zealand faunas, nestorine parrots were asso-ciated with Strigops and platycercine Cyanoramphus parakeets.These parakeets are part of a recent radiation in the southwestPacific (Kearvell et al., 2003), which therefore may have benefitedfrom a reduced diversity of parrots in New Zealand. It seems fea-sible that, following extreme reduction in forest communities dur-ing one of the Pleistocene glacial periods, parrots were reducedto a minimum of a Strigops lineage and a Nestor lineage, perhapsonly three species in all on the mainland. Subsequent expandedforests would have then had vacant niches facilitating the colo-nization of New Zealand by the Cyanoramphus lineage.

ACKNOWLEDGMENTS

The authors are indebted to the unstinting efforts of many fieldassistants in the St Bathans Fauna project that have amassedthe fossils between 2001 and 2010. Many people helped overthis 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, andA. B. Camens who helped in multiple years of excavations to re-veal the fossils described here. We thank J.-C. Stahl, NMNZ, forsome images. We are particularly thankful for the continued gen-erous support to this project by the landowners A. and E. John-stone. This research was part of the project DP0770660 fundedby the Australian Research Council, and DP1094411 funded by aUNSW 2010 Goldstar award. R.P.S. was funded by a grant fromThe Brian Mason Technical Fund. We thank W. Boles for com-ments on a draft of the text and comments by G. Mayr and an-other referee that helped improve the text.

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