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IBIS 135: 8-18

A new (Psittaciformes: Cacatuidae) from the Tertiary of Riversleigh, northwestern Queensland, and an evaluation of rostral characters in the systematics of

W AL TER E. BOLES Division of Vertebrate Zoology (), Australian Museum. 6-8 College Street. Sydney, N.S. W. 2000, Australia

: The first Tertiary cockatoo or any psittaciform from Australia is recorded from an incom- plete rostrum from the Riversleigh deposits of northwestern Queensland.Australia. It is placed in the extant Cacatuabut cannot be diagnosed at the specific level. This cockatoo was probably similar in size to the modern C. roseicapilla.with a short. unspe- cialized and probably white plumage. The rostrum is useful in studies of the system- atics of parrots. Rostral characters support sometraditional subdivisionsof the order. while suggestingthat others warrant re-examination.

Parrots comprise one of the most distinctive orders of birds. Milne- Edwards (1867 -1871 ) described Psittacus verrauxi from The approximately 340 living speciesof Psittaciformeshave the Early Miocene of France. for which Lambrecht (1933) a pan-tropical and southern distribution. Of these birds, the later proposed the new generic name Archaeopsittacus. Wet- (Cacatuidae),with 18-21 speciesin five genera, more (1926) named Conuropsisfratercula. the only pre-Pleis- centred on the Australo-Papuan region, form one of the tocene record of this order from the New World. No bills or most characteristic groups. With five generaand 10-12 spe- skulls of these Tertiary forms have thus been reported. and cies, Australia is' the centre of cockatoo diversity, although it is not possible to determine at what stage these early speciesextend as far as the Philippines in the north, Sulawesi parrots were in the evolution of the distinctive cranial mor- and the Sunda Islands in the west and the Solomon Islands phology that characterizes parrots. in the east (Forshaw 1981). Thesebirds form one of the most All previous Australian records of the Psittaciformes are distinctive elementsof the Australian avifauna. Despite this of Quaternary age and are indistinguishable at specific It:vel wide distribution, there is only a very limited fossil history from Recent taxa (Baird 1991). Smaller, non-cacatuine forms of parrots (Olson 1985). At present the known record con- have been reported from several sites (e.g. Hope et al. 1977 , tains only a handful of Tertiary taxa, none of which is a Rich et al. 1983. Baird 1985; summarized in Baird 1991); cockatoo. Fro!D Australia, with 55 extant ,no pre- other psittacine material has not yet been evaluated. Cock- Pleistocenematerial has been reported (Vickers-Rich 1991). atoos have only recently been reported from the fossil rec- The only cockatoos are of Pleistoceneage and can be as- ord-none was listed by Brodkorb (1971)-and these have signed to extant species(Baird 1985, 1991). been only from southeastern Australian sites. The identified Describedhere is a fossil cockatoofrom the Early to Middle species are all referable to living taxa: Red-tailed Cockatoo Miocene of northwestern Queensland,constituting the old- magnificus, Glossy Cockatoo C. lathami, Black est fossil record of the order for Australia and of the Caca- Cockatoo C. funereus. Gang-gang Callocephalon fimbriatum. tuidae for the world. Long-billed tenuirostris and C. rosei- capilla. Other cockatoo material. as yet unpublished. has been recovered from a number of other Pleistocene sites (R.F. Baird. pers. comm.). FOSSIL HISTORY

The fossil record of the Psittaciformes has been summarized by Olson (1985). The earliest report is Palaeopsittacusgeorgei GEOGRAPHIC AND GEOLOGIC SETTING Harrison (1982) from the Lower Eocene of England. al- though there have been doubts expressed about the iden- The Riversleigh depositsare located on Riversleigh Station, tification (Olson 1985). Mourer-Chauvire (1982) noted re- approximately 200 kID north of Mount Isa, northwestern mains from the Upper Eocene of France. which she assigned Queensland.More than 100 sites and local faunas are now to the Psittacidae without elaboration. known from this area but few of these have been described Only two Miocene records have so far been published. in the literature. Preliminary stratigraphic relationshipshave although P. Ballman (pers. comm. to Olson 1985) has drawn been determined by Archer et al. (1989), but these, as well attention to the recovery of some as yet undescribed as the interpreted ages,must be regarded as tentative. remains from the Middle Miocene of Germany and France. The fossil subject of this paper came from the RSOLocal 1993 A NEW COCKATOO FROM QUEENSLAND 9

Fauna from RSO Site. This site is at the lower end (180 m some characters. Some taxa do not fit comfortably into the a.s.I.) of the Godthelp Hill Sequence. which forms part of categoriesadopted, but some limitation has been made on System B of Archer et al. (1989). System B. a collection of the number of categoriesused to accommodateunique ex- regionally clustered sites differing in age but not significantly amples. The rhamphotheca often bears little resemblanceto in position and/or spatially isolated but approximately con- the underlying bone and, together with the cere, prevents temporaneous. is considered to be ?Early to Middle Miocene. any external assessmentof the rostral characters. The taxa It appears to represent fossil assemblages accumulated in examined are listed in Appendix II. Rostral characters for pools or shallow lakes (Archer et al. 1989). many Australo-Papuan and selected extralimital psittaci- Based primarily on information derived from the rich and form genera are given in Appendix III. These should be diverse mammal remains, Archer et al. (1989) have inter- useful in the identification of unknown rostra. preted the vegetation of Riversleigh at this time to have been "dense. species-rich gallery rainforests probably similar to those that persist today in mid-montane New Guinea.'. SYSTEMA TIC P ALAEONTOLOGY The Riversleigh rostrum clearly belongs to the Cacatuidae. A suite of character statesserves to identify any member of MA TERIALS AND METHODS this family. Most of the character states are found in other Measurementswere made with vernier calipers accurate to taxa; however. the shape of the naso-frontal hinge and the 0.05 mrn and rounded to the nearest 0.1 mm. specializedconfiguration of the cere region are diagnostic. Names of psittaciform taxa follow Forshaw (1981) except Five genera of cockatoos are currently recognized: Pro- for changesbased on more recent studies. For convenience bosciger(. 1 species).Calyptorhynchus (black of reference. three families of Psittaciformesare recognized cockatoos. 3-5 species). Callocephalon(Gang-gang. 1 spe- in the discussionof this fossil (after Forshaw 1981): Loriidae cies). Cacatua(white cockatoos. 12-13 species)and Nym- {lorikeetsand lories). Cacatuidae(cockatoos) and Psittacidae phicus (. 1 species).The character states, particu- (all other parrots). larly of the nostrils, and the shape and size of the rostrum, The fossil material referred to herein is currently held in indicate that the fossil parrot from Riversleighis most similar the collections of the Vertebrate PalaeontologyLaboratory . to. and indeed inseparable from. Cacatua. University of New South Wales but will be transferred to The other cacatuine genera can be distinguished from the Queensland Museum at the completion of this study; Cacatuaby the following rostral characters: prefixesto registration numbers for theseinstitutions are AR Callocephalon. Cere region saddle-shapedwith thin centre; and QM. respectively. nostrils oriented to the side (300from midline of culmen); length: width ratio smaller than in all cacatuid speciesex- cept for some individuals of C. sulphurea. ANAL YSIS OF CHARACTERS Calyptorhynchus. Cere region saddle-shapedwith thin cen- The decurved upper mandible of parrots is distinctive. con- tre; internarial septum wider; nares small. directed to side; size large (compared with small Cacatuaspecies). stituting one of the major external charactersby which these birds are recognized. A few other orders of raptorial birds Probosciger. lnternarial septum wider; nares small. direct- have superficially similar hooked bills, but each group can ed to side; size very large; shape diagnostic. be easily distinguished from parrots by a combination of Nymphicus. Cereregion spade-shaped(Fig. 18); internarial characters, not all applicable to each group, e.g. presence and shape of the naso-frontal hinge, shape of the nostrils, septum narrow; size small. presenceof a nasal tubercule, absenceof a bifurcated groove on the culmen and degreeof lateral compressionof the ros- trum. Order PSITTACIFORMES Wagler. 1830 The most easily recognized feature of parrot bills is the Family CACA TUIDAE groove on the upper surface that runs from the cere region down the internarial septum and then bifurcates. continuing (Rostrum) Naso-frontal outline with centre raised and along each side of the culmen towards the tip. The presence rounded. extending above point of attachment with skull; of this groove is diagnostic for the Psittaciformes. cere region smoothed and raised as saddle-shapedplatform There are about 340 speciesof living or recently extinct (shapedifferent in one species);corners of naso-frontal hinge speciesof parrot placed in 81 genera (Forshaw 1981). In entended into acutely angled horns (approximately 90" in order to assessthe relationship of the fossil rostrum, 10 one species);internarial septum medium to very wide (nar- characters were evaluated from skulls. The characters and row in one species); nares small to large. facing directly the states used are given in Appendix I and illustrated in forward or towards the side; tomium sigmoid shaped; size Figure 1. There is a degreeof arbitrariness to the scoring of medium or large (one speciessmall. one very large). 10 W.E.BOLES IBIS 135

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MEASUREMENTS

Culmen length (tip broken) 20.6 mm; width of naso-frontal hinge from caudolateral corner to midpoint 9.3 mm; greatest dorsoventral distance 17.2 mm; greatest rostrocaudal dis- tance 15.1 mm; width of internarial septum at midpoint 5.4 mm.

DESCRIPTION Rostrum with damageto the right side, caudal end, tip, and some areas on the left side above and below the left orbit, along the tomium and caudoventral corner. In anterior view, lateral side slightly concave; top margin extending laterally then curving medially at about 60- forming a distinct acute 'horn'; cere region raised and flattened, distal side concave, other sides not iliscernable owing to damage; nostrils me- ilium, forward facing, set on side of rostrum about equiilis- tant between culmen and cranial border; internarial septum wide; groove along midline of culmen, curving right just distal of line connecting midpoints of nares, at which point another groove curves left, both continuing along the re- spective sidesof the culmen towards the tip. In lateral view, tomium deep at posterior, curving concavely dorsad below Figure 2. Rostrum of fossil cockatoo Cacatua species indeterminate nostril, then ventrad towards tip; caudalmostsection of tom- (QM F16866) from Tertiary of Riversleigh. northwestern Queens- ium lost through abrasion. In ventral view, premaxilla con- land. cave with a narrow slit-like opening along caudal end of rostrocaudal midline; maxilla more dorsal, rostral end un- Genus Cacatua Vieillot, 1817 dercutting caudal end of premaxilla, slightly concave, thin ridge running along rostrocaudal midline with a slit-like (Rostrum) Cereregion saddle-shapedwith wide centre. In- opening on either side of rostral end. In caudal view, many temarial septum wide; nares medium, directed forward; surfacesabraded, no obvious features present. overall shape'normal' ; overall sizemedium. The upper man- Based on size and form, the Riversleigh cannot be dible with Cacatuaexhibits a range of variation in size and distinguished from some similarly sizedextant species,such shape,excee

Cacatua species indeterminate DISCUSSION Two major subdivisions can be recognized within Cacatua MA TERIAL (Adams et al. 1984). One group is characterizedby the pos- sessionof heavy bills, up curving coloured crests and round OM F16866 (AR 10542), incomplete rostrum, damaged on wings (alba,9alerita, leadbeateri, moluccensis, ophthalmica, sul- right side (Fig. 2). phurea);within this group leadbeateriand alba have substan- tially smaller bills than the other speciesbut are alike in the shapes of the crests and wings. The other subdivision has LOCALITY smaller bills, short uncoloured crestsand more slenderwings RSO Site, Godthelp Hill Sequence (System B of Archer et al. (ducorpsi,9offini, haematuropy9ia,pastinator, roseicapilla1, ten- 1989), Riversleigh Station, Queensland. uirostris). Large individuals of the larger species(e.g. pastin-

1 The Galah C. roseicapilla has frequently been placed in the mono- LOCAL FAUNA AND AGE typic genus Eolophus on the basis of larger zygomatic process, smaller RSO Local Fauna, (Early to Middle Miocene (Archer et al. temporal fossa, crescentic rather than semicircular auditory meatus, 1989). plumage colouration and behavioural traits (Holyoak 1970, Forshaw 12 w. E. BOLES IBIS 135

ator) approach and possibly slightly overlap smaller exam- mosaically distributed among these to the south (Tanimbar ples of leadbeateriand alba in bill size. Islands-goffini) and west (central and northern Moluccas- On the basis of bill size. the Riversleigh bird most likely alba) of New Guinea, and on the periphery of the genus's belongs to the small-billed group of white cockatoos. It is range to the north (Philippines-haematuropygia) and east smaller than the smallestindividuals of the large-billedgroup. (Solomon Islands-ducorpsi). Other than on the Australian Assuming that the correlation of the other charactersholds, mainland, no more than a single speciesoccurs at any lo- this form would have been a relatively small cockatoo with cality. a short. unspecialized,uniformly coloured crest. Following As an environmental indicator, the Riversleigh bird nei- the trend of modern species.its plumage would have been ther corroborates nor contradicts current ideas on the cli- predominantly white. mate and vegetation at Riversleigh during the Early to Mid- Although the tip is missing. it is almost certain that the dle Miocene. In Australia. the smaller members of Cacatua bill of the Riversleigh bird was not elongated as in c. ten- (roseicapilla.tenuirostris. pastinator) are birds of the open me- uirostris. Given the small basal cross-sectionof the broken sic, semi-arid and arid zones.On the periphery of the range tiP. the tiP. if attenuated, would have to have been so to a of the genus. ducorpsi.goffini and haematuropygiaoccur in degree in excessof any living form. The inward curvature both primary rainforest and secondarygrowth, also entering of the distal end eliminates this as a possibility. agricultural areasfor crops (Forshaw1981). Therefore,while Although it is possible to determine to which section of the presence of the Riversleigh bird is consistent with a the genus the Riversleigh bird probably belongs, the fossil rainforest environment. it is not contra-indicative of drier, giveslittle information about the phylogenetic history of the more open habitats. genus. Its morphology is as modern as any extant member of the species.indicating that the genus was present in an essentiallymodern form by the early Miocene.There is noth- SYSTEMATIC USE OF THE ROSTRUM IN THE ing to rule out that large-billed specieswere not also con- PSITTACIFORMES temporaneous. Both groups have representativeson the Australian main- Parrots are an intensively studied group, with a wide range land (large-billed: galerita, leadbeateri:small-billed: pastin- of morphological, anatomical, pterylographic and behav- ator, tenuirostris, roseicapilla).The large-billed species are ioural characters having been used by numerous authors centred on New Guinea (galerita).with speciesoccurring on (for a summary of previous work and characters, seeSmith the islands directly to the west (moluccensis-southern Mo- 1975). Despitethis attention, parrot systematicsare far from luccas; sulphurea-Sulawesi. Sunda Islands) and east settled. (ophthalmica-Bismarck Archipelago).Small-billed forms are Although previously ignored, the rostrum of parrots has some value in the systematic study of this group. Features of the external bill have been used in systematicstudies but

1981). The validity of this assignment was addressed by the bio- have almost invariably referred to aspectsof the external chemical studi.es of Adams et al. (1984), which showed that rosei- rhamphothecal covering, not to the underlying bone. The capilla belongs in Cacatua and that it, tenuirostris and pastinator "are correlation between the external features and their appear- monophyletic to the exclusion of all other [Australian] species. This ance on the bone is not high, and, indeed, some of the implies that the colour pattern of C. roseicapilla, and its crescentic characteristic bill morphologiesthat are apparent when the auditory meatus, small temporal fossa and pink natal down, are rhamphotheca is present are reduced to considerable uni- unique derived traits of minor taxonomic significance" (p. 120). Yet formity in its absence.Some characters of the rostrum ap- later, one of the authors of that paper (Schodde 1989) resurrected pear more related to peculiarities of feeding and food choice Eolophus because of the "colour pattern, pink-downed nestlings, and than as clues to a taxon's phylogenetic background. Thus distinctive skull, with well-developed maxillary processes and re- rostrum shape, length: width ratio and size may be impor- duced temporal fossae", the same characters previously disregard- ed. This implies that, of extant taxa, roseicapilla is the sister-group tant in diagnosing certain taxa but have only limited use in of all Australian white cockatoos. However, to accept both sets of suggestingrelationships with other psittacine groups. Some conclusions (i.e. the tree of Adams et al. 1984 and the generic as- other character statesare at least somewhat related to over- signment of Schodde (1989) results in a paraphyletic phylogeny. all bill size. On the other hand, several characters are par- The only solution that permits retaining Eolophus within the tree ticularly valuable in indicating where relationships lie. In topography produced by Adams et al. (1984) is to separate the cor- various combinations, the naso-frontal hinge outline and ellas from the remaining Australian white cockatoos at generic level comers, cere region configurations,intemarial septumwidth, as well. This still does not address the relationship to Eolophus of tomium shape,nostril size and orientation, and culmen cur- non-Australian species not included in the study of Adams et al. vature can aid in the diagnosisof various subgroupsof the (1984). There is no feature by which the rostrum of Eolophus can be parrots. diagnosed from those of Cacatua; some individuals cannot be distin- This survey was not intended to be a thorough systematic guished from other species. If Eolophus were accepted as valid, it study of the rostra acrossall taxa in the Psittaciformes.Ad- would not be possible to assign the Riversleigh specimen to one ditionally, there are other characters that could be scored genus or the other. in a more detailed study, such as bill curvature perpendic- 1993 A NEW COCKATOO FROM QUEENSLAND 13

ular to the culmen. the configuration of the narial region. ered it allied with the fig-parrots. Verheyen (1956) and Glen- shape of the tip. etc. Nonetheless.the limited examination ny (1957) created a separate subfamily for it. Generally, it of the rostra performed here allows some comments to be has been agreedthat, despitesome superficial similarities to made on aspectsof parrot systematics.Even where no useful other groups, Lathamusbelongs with the platycercines. Ex- indication of relationships is obtained. the scores of each amination of the rostrum supports this conclusion. Not only taxon should assist in the identification of unknown taxa. does it have the characteristic culmen curvature and pro- The cockatoos are easily the most distinctive group of jection of the platycercines,it has a wider internarial septum parrots on the basis of rostral morphology. These charac- and tomium shape than do the lorikeets. teristic features were discussed above. Several subgroups The remaining Old World taxa are often separated into can be recognized:Probosciger is distinctive. but its oversized two subgroups:the large African forms (Psittacus,Coracopsis rostrum is so extensivelymodified that many of its character and Poicephalus:) and the primarily Asian, Aus- statesare unique; Callocephalonand Calyptorhynchusare sim- tralo-Papuan and South Pacific forms (Agapomis,Geoffroyus, ilar in having similar cere region configurations and lateral ., ,, Alisterus, Apro- orientations of the nares; Cacatuais set apart by the cere smictusand Prosopeia:Palaeornithinae). The latter is some- region configuration and the size and orientation of the nos- times divided, segregating the last four Australo-Papuan trils. and South Pacific genera in the Polytelinae. The Cockatiel Nymphicushollandicus has had a chequered Divisions within these Old World forms can be discerned taxonomic history; some authors placed it with the Polyte- but show some inconsistency with the traditional bound- linae or Platycercinae(Verheyen 1956. Brereton 1963. Con- aries. Agapomis,Psittacula and Poicephalusare the only gen- don 1975) and others assignedit to the Cacatuidae.usually era surveyed that have large, upwards directed nostrils; the as a sister taxon to the other genera of cockatoos(Salvadori nostrils of Tanygnathusare also directed upwards but are 1891. Thompson 1900. Reichenow 1913. Peters1937. Glen- proportionally smaller (probably a consequenceof the over- ny 1957. Holyoak 1972. Smith 1975. Wolters 1975-1982. sized bill.) Eclectusstands apart on a combination of septum Forshaw 1981. Adams et al. 1984). The rostrum demon- width, size of the nares and the unique configuration of the strates unequivocably that Nymphicusis a cockatoo. It has cere region. The other generado not differ substantially from the states for the outline and corners of the naso-frontal each other, and all appear relatively generalized in their hinge. configuration of the cere region and tomium shape character states, based on distribution of these features that diagnosethis group. Its unique form of the raised cere through the parrots. region indicates that it is separatefrom the other cockatoos Peters (1937), Glenny (1957), Smith (1975) and others at a significant level. have included the New World taxa in a singletribe/subfamily Another major sectionof the Psittaciformesis the lorikeets without further internal division, whereas Salvadori (1891), and lones. Loriidae. While not as easily diagnosedon rostral Thompson (1900), Reichenow (1913), Verheyen (1956), characters as the cockatoos.this group can neverthelessbe Brereton (1963) and Wolters (1975-1982) distributed these segregatedon a suite of character states.These include the among at least two subgroups(although with differing com- cranially flaftened culmen. relatively straight tomium. binations of component taxa). There is an indication that rounded naso-frontal outline and correlated obtuse comer there are indeed several diagnosablesubgroups. angles and the very thin intemanal septum. Although there are differencesbetween them, Anodorhyn- Three of these groups can be identified on the basis of chus, Ara, Aratinga, Conuropsis,Cyanoliseus, Enicognathus, rostral shape. but there are few other clues to where their Myiopsitta.Pionites, Pyrrhura and Rhynchopsittaall sharesome relationships lie. The fig parrots (Opopsitta.Psittaculirostris) character stateswith restricted distribution: nostrils that are have noticeably broad rostra. the width approaching the small (markedly so in Myiopsitta) and directed upwards or length more closely than for any other similarly sizedforms. sideways.Ara manilata differs from all other examined taxa Other than their very different rostral shapes.Nestor and of this assemblage,including other membersof its genus, in Strigopsare superficially similar .The character states by both thesecharacters. (Anodorhynchusand, to a lesserextent, which they resembleone another are alsowidely distributed Ara approach Proboscigerin some characters. This homo- in other generaand appear generalized:therefore. they may plasic expression seemsdirectly related to their dispropor- have little significance.The more definedcere region appears tionately large size.) All these genera are frequently segre- to characterize many larger forms that are otherwise not gated in the Conurinae. Brotogerisis usually also included closely related. here but it differs in its larger, forward-facing nostrils. ~ost of the features of the platycercine rostrum seem The other New World generaexamined, Amazona, Pionus, unspecializedand undiagnostic. Only the curvature of the Touit, Forpusand Bolborhynchus,share the widespread and culmen. with its flattened cranial end and projection on the apparently generalizedlarge, forward-facing nares.Theyare septum. servesto identify the component taxa of this Aus- sometimesplaced in the samesubfamily, sometimesin sep- tralasian group. Lathamushas long been problematic to par- arate ones (the first three in Amazoninae, the last two in rot systematists.On the basisof its tongue modified for nectar Forpinae). None has obviously unusual features, although feeding. some authors (Peters 1937. Wolters 1975-1982) they do differ from each other in several states. Without have associatedit with the lorikeets. Forbes (1879) consid- additional representativesof each assemblage,it is not pos- 14 w. E. BOLES IBIS 135

sible to assessthe importance of this variation. On the basis Condon, H.T. 1975. Checklist of the Birds of Australia, Part 1, of purported similarities with the platycercines. Brereton Non-passerines. Melbourne: Royal Australasian Ornithologists (1963) placed the Forpidae in a different superfamily from Union. the other New World genera. near the Platycercidae.Forpus Forbes, W.A. 1879. On the systematic position of the genus La- and Bolborhynchusboth lack the platycercine culmen pro- thamusof Lesson.Proc. Zool. Soc. Land. 1879: 166-174. jection but otherwise agree with the Australian group in its Forshaw, J.M. 1981. Parrots of the World, 2nd ed. Sydney:Lands- downe. generalizedcharacters. Glenny, F.H. 1957. A revised classification of the Psittaciformes This cursory survey of rostral characters was initially un- basedon the carotid artery arrangement patterns. Ann. Zool. 11: dertaken to make possiblethe identification of isolatedparrot 47-56. rostra. and it has been shown that. for some groups of these Harrison, C.J.O. 1982. The earliest parrot: a new speciesfrom the birds. identification can be made even to specieslevel. It British Eocene.Ibis 124: 203-210. was also revealed that the rostrum provides useful infor- Holyoak, D.T .1970. Structural characters supporting the recog- mation towards the systematic study of the Psittaciformes. nition of the genus Eolophusfor CacatuaroseicapiIla. Emu 70: 200. Some traditional groupings are strongly supported. Some Holyoak, D.T. 1972. The relation of Nymphicusto the . others are brought into question and deserve further ex- Emu 72: 77-78. Hope, J.W., Lampert, R.J., Edmondson,E., Smith. M.J. & Van Tets, amination. Rostral characters should not by themselvesbe G.F. 1977. Late Pleistocene faunal remains from Seton rock the basis for classifying parrots. but they can serve as valu- shelter, Kangaroo Island. South Australia. I. Biogeog.4: 363-385. able adjuncts to other character systemsin broad-basedsys- Lambrecht, K. 1933. Handbuch der Paiaeornithologie.Berlin: Ge- tematic work on these birds. briider Borntraeger. MiIne-Edwards, A. 1867-1871. RecherchesAnatomiques et Pa- For the use of specimens under their care, I thank the curators of leontologiquespour Servir a I'Histoire des OiseauxFossiles de la the American Museum of Natural History, Australian Museum, France (4 vols). Paris: Victor Masson et Fils. Australian National Wildlife Collection, Museum of Victoria, Mourer-Chauvire, C. 1982. Les oiseaux fossilesdes Phorphorites Queensland Museum, South Australian Museum, United States Na- du Quercy (EoceneSuperieur a OligoceneSuperieur): implications tional Museum of Natural History and University of Kansas Museum paieobiogeographiques.Geobios, mem. spec. 6: 413-426. of Natural History. For discussion of this study and comments on Olson, S.L. 1985. The fossil record of birds. In Farner, D.S., King, the manuscript, I am grateful to Dr M. Archer, Dr R. F. Baird, Mr J.R. & Parkes, K.C. (eds)Avian Biology, VIII: 79-238. New York: H. Godthelp, Dr S. Hand, Dr P. Vickers-Rich and Dr T. F. Flannery. Academic Press. Ms L. Albertson drew the figures. Mr A. Farr took the photographs. Peters, J.L. 1937. Check-list of Birds of the World. 3. Cambridge, The Australian Museum provided avenue in which to work and Mass.: Harvard University Press. funds to visit other relevant collections. The Riversleigh material Reichenow, A. 1913. Die Vogel. Handbuch der Systematischen was collected via an ARC Programme Grant to M. Archer; a grant Ornithologie, 1. Stuttgart: F. Enke. from the Department of Arts. Sport, the Environment. Tourism and Rich. P.V., Van Tets, G.F., Orth, K., Meredith, C. & Davidson, P. Territories to M. Archer. S. Hand and H. Godthelp; a grant from the 1983. Prehistory of the Norfolk Island biota. In Schodde, R., National Estate Programme Grants Scheme to M. Archer and Dr A. Fullagar, P. & Hermes, N. (eds)A review of Norfolk Island Birds: Bartholomai; a!ld grants in aid to the Riversleigh Research Project pastand present: 7-29. Aust.Nat. ParksWildl. Serv.Spec. Publ. 8. from Wang Australia Pty Ltd.. ICI Australia and the Australian Salvadori, T. 1891. Catalogue of the Psittaci, or Parrots, in the Geographic Society. Collection of the British Museum. London: British Museum (Nat- ural History). Schodde,R. 1989. New subspeciesof Australian Birds. Canberra Bird Notes 13: 119-122. REFERENCES Smith, G.A. 1975. Systematicsof parrots. Ibis 117: 18-68. Thompson. D.W. 1900. On characteristic points in the cranial Adams, M., Baverstock, P.R., Saunders, D.A., Schodde, R. & Smith, osteologyof the parrots. Proc. Zool. Soc. Lond. 1899: 9-46. G.T .1984. Biochemical systematics of the Australian cockatoos Verheyen. R. 1956. Analyse du potentiel morphologique et projet (psittacifonnes: Cacatuidae). Aust. I. Zool. 32: 363-377. d'une nouvelle classification des Psittaciformes.Inst. R. Sci. Nat. Archer, M., Godthelp, H., Hand, S.I. & Megirian, D. 1989. Fossil Belgique 55: 1-54. mammals of Riversleigh, northwestern Queensland: preliminary Vickers-Rich, P. 1991. The Mesozoicand Tertiary history of birds overview of biostratigraphy. correlation and environmental change. on the Australian plate. In Rich, P.V., Baird, R.F., Monaghan,J.M. Aust. Zool. 25: 29-65. & Rich, T.H. (eds) The Vertebrate Palaeontologyof Australasia: Baird, R.F. 1985. Avian fossils from Quaternary deposits in 'Green 721-808. Melbourne: Nelson. Waterhole Cave', south-eastern South Australia. Rec. Aust. Mus. Wetmore, A. 1926. Descriptionsof additional fossilsfrom the Mio- 37: 353-370. cene of Nebraska. Am. Mus. Novit. 211: 1-5. Baird, R.F. 1991. Avian fossils from the Quaternary of Australia. Wolters.H. 1975-1982. Die VogelderErde.Hamburg: Paul Parey. In Rich, P.V., Baird, R.F., Monaghan, I.M. & Rich, T.H. (eds) The Vertebrate Palaeontology of Australasia: 809-870. Melbourne: Submitted 25 July 1991; accepted 15 October 1991 Nelson. Brereton, I.L. 1963. Evolution within the Psittacifonnes. Proc. Int. Om. Congr. XIII: 499-517. Brodkorb, P. 1971. Catalogue of fossil birds. part 4 (Columbiformes through Picifonnes). Bull. Florida State Mus. 15: 163-266. 993 A NEW COCKATOO FROM QUEENSLAND

APPENDIX I with faint indication that it resemblescondition 3a. very wide lateral profile (Probosciger). Characters and character states used in identifying rostra of parrots (C) Lateral angle of naso-frontal hinge (Fig. IC) SeeFigures lA-J for illustrations of each character state. The angle formed by the junction of the naso-frontal hinge and the lateral margin of the rostrum and the configuration of this corner (A) Outline of naso-frontal hinge (Fig. IA) region.

The shape of the naso-frontBl hinge when viewed from directly in I End of naso-frontal hinge rounded. lateral margin vertical, form- front. ing obtuse angle, c. 12O" (e.g. , Glossopsitta);

1 Slightly but smoothly rounded convexly over entire. length with 2 End of naso-frontal hinge more or less straight, lateral margin no indentation at the midpoint (e.g. rrichoglossus); vertical, forming approximately 9O" angle (e.g. Alisterus, Cyanoram- 2 Slightly indented at the centre, the remainder of the hinge being phus); more or lessstraight (e.g.most Psittacidae),slightly elevatedthrough 3 End of naso-frontal hinge more or less straight, lateral margin the centre region (e.g. , Agapomis,Eclectus, Nestor) or slightly angled slightly towards midline of rostrum, forming a slightly acute convexly rounded (e.g. Melopsittacus.Psittaculirostris); angle, c. 60-75° (e.g. Eclectus, Platycercus, Psittacus);

3 Slightly to moderately elevated through the centre region with a 4 End of naso-frontal hinge more or less straight, lateral margin gradual angle at the commencement of the rise and without any slightly acute at junction then becoming more vertical, forming small indentation (e.g.Amazona, Psittacus); but nonprominent 'horn' at corner (e.g. Prosopeia, Strigops);

4 Considerablyelevated through the centre region with a marked 5 End of naso-frontal hinge more or less straight, lateral margin angle at the commencementof the rise and no indentation (e.g.most sharply acute at junction then becoming more vertical, forming large Cacatuidae);a variation on the last condition is seen in Probosciger. and prominent 'horn' at corner (e.g. Cacatua, Ca/1ocephalon,Calyp- which has the centre region markedly swollen obscuring a definite torhynchus, Nymphicus). sharp angle at the commencementof the rise. In lateral view, in the first three conditions the highest level of the naso-frontal hinge is evenwith or just lower than or slightly abovethe point of attachment (D) Width of internarial septum (Fig. ID) with the skull; in the last the raised centre extendsabove this point, This is a somewhat subjective combination of the overall width of sometimesmarkedly so. the septum and its size in relation to the entire upper mandible. The width of the septum is through the midpoint and that of the rostrum is the length of the arc taken from one side of the tomium across (B) Configuration of cere region (Fig. lB) the dorsal surface. passingthrough this midpoint and continuing to the other side. Somewhat cotTelated with the previous character is the configu- ration of the area circumscribed by the rear edge of the nares, the 1 Verythin. Septum :$1 mmorbetween 5% and 10% of the rostrum (usually) proximal end of the internarial septum and the naso-frontal arc length (e.g. Trichoglossus.Glossopsitta. Psephotus); hinge-the region covered by the fleshy cere. 2 Thin. Septum 1-2 mm or about 10% of the rostrum arc length 1 Extending across the base of the rostrum and depressed below (e.g. Nymphicus,Bamardius. Polytelis); the level of the hinge and the remainder of the rostrum (only ob- 3 Medium. Septum about 3-5 mm or between 10% and 20% of the served in Eclectus among taxa examined); rostrum arc length (e.g. Amazona.Prosopeia. Strigops); 2 At the same level as the naso-frontal hinge and either (a) with a 4 Medium wide. Septum about 7 mm or about 25% of the rostrum slight indentation in the centre (e.g. Polytelis. Bamardius, Platycercus) arc length (septum width about the same as that of lateral section or (b) without an indentation and more or less level or even slightly of rostrum between the naris and tomium) (Cacatua.Eclectus); rounded (e.g. Pezoporus. Lorius. Strigops); some taxa, particularly larger forms and platycercines, exhibit a more clearly defined distal 5 Wide. Septum >10 mm or >30% of the rostrum arc (septum border to this area (.) (e.g. Nestor. Strigops, Platycercus); width greater than that of lateral section of rostrum between the naris and tomium) (Probosciger,Calyptorhynchus. Callocephalon. Ca- 3 Most or part of the region raised in a discrete area: (a) saddle- catuasulphurea). shaped broadly across entire base of septum, distal border concave, wide in lateral profile (Cacatua); (b) saddle-shaped but very thin across base of septum, widely concave distally, narrow in lateral (E) Size of nares (Fig. 1£) profile (CalIocephalon. Calyptorhynchus); (c) across entire central cere Two measuresof nostril size have been made: one is based on the region, distal border straight and extending further distally along opening viewed facing them from their direction of orientation. the internarial septum, wider in lateral profile than Cacatua (Nymphicus); other is the extent of the opening along the side of the nostril in 4 Raised in non-discrete area, very bulbous, no distinct borders but lateral and dorsal views. Both assessmentsare somewhatsubjective. 16 w. E. BOLES JS

basedin part on the absolute size and the proportional size relative 1 Smoothly cuT\led.Culmen rounded over entire length without any to the rest of the rostrum. Narial width is that acrossthe diameter marked flattening (e.g. Probosciger. Prosopeia. Anodorhynchus); ( + ) of one naris; the proportional width is basedon both nostrils and is curve arched (Callocephalon, Calyptorhynchus); ( -) curve depressed measured in a manner similar to proportional septum width. In (e.g. Amazona. Opopsitta); forward aspectthe states are: 2 Cranially flattened-smooth. Culmen flattened on craniall13-1f2 be- I Small. One naris <3 mm or both constitute less than 20% of fore curving downwards without obvious angle at transition (e.g. rostrum arc length, narial diameter is less than the width of the lorikeets, Psittacus. Psittacula); septum and of the lateral edge between the naris and tomium 3 Cranially flattened-angled. Culmen flattened on cranial 'I], before (Myiopsitta.Calyptorhynchus lathami); curving downwards with obvious angle and a small projection at 2 Medium.One naris 5-6 mm or both constitute between 20% and the point of transition (e.g. Barnardius. Platycercus, Melopsittacus); 35% of rostrum arc length, narial diameter is always less than the 4 Distally flattened. Culmen flattened on distal 'I]-1f2 and curved width of the septum and usually less than that of the lateral edge cranially (Myiopsitta. Cacatua tenuirostris). between the naris and tomium (e.g. Eclectus.Probosciger, Calypto- rhynchus.Callocephalon); (H) Shape of tomium (Fig. lH) 3 Large.One naris >6 mm or both constitute more than 35% of rostrum arc length, narial diameter is greater than or equal to the The outline of the tomium is assessed from the lower rear (cranial) width of the septum and of the lateral edgebetween the naris and comer forwards. tomium (e.g. most psittaciforms examined). 1 Sigmoid. The basal section is deep, before curving dorsally then The size of the nares in lateral and dorsal aspect seemsto some ventrally towards tip (all Cacatuidae except Probosciger); extent to be related to the overall sizeof the rostrum, with the larger speciesexhibiting the greater lateral/dorsal extent; this doesnot hold 2 Smooth. The basal section is not much deeper than rest of tomium, for the Cacatuidae.This character is not as easilyscored as the width which is more or less smooth (Strigops), or with a slight, gradual in forward aspect. The states for lateral/dorsal extent are: concavity towards tip (e.g. Trichoglossus. Polytelis, Nestor. Myiopsitta);

Small. More or less equal (I x) to the width in forward aspect, in 3 Notched. The basal section is not much deeper than rest of tomium, lateral view round naris in shape (e.g.Cacatua. Callocephalon. Nym- and there is a concavity before the tip. In contrast to the previous phicus,Eclectus, Platycercus, Myiopsitta); state, the cranial border of the concavity is abrupt, often creating a notched or 'toothed' appearance (e.g. Platycercus, Bamardius, Eclec- Moderate.Between I x and 1.5 x the width in forward aspect, in tus. Amazona). It is sometimes difficult in some taxa to distinguish lateral view naris somewhat oval in shape (e.g. Trichoglossus,Alis- this state from the previous one (e.g. Melopsittacus. Geoffroyus); terns,Prosopeia. Bamardius); 4 Tomium is concave overall except for a small rise near the centre Marked.Between 1.5 x and 2 x the width in forward aspect,in lateral view naris oval and elongated in shape (Nestor,Strigops. Amazona). (Probosciger).

(F) Orientation of nares (Fig. IF) (1) Width: length ratio (Fig. II)

The angle of orientation is judged relative to the horizontal and to Most parrots have rostra in which the length is more than 1 x but a vertical plane passing through the distocranial midline. Within the less than 2 x the width. There are a few exceptions in both directions. general categories. several substates are discernable. 1 Length = width (Cacatua sulphurea. Calyptorhynchus lathami, ap- 1 Forward. Nares oriented laterally less than 100 from forward and proached by Psittaculirostris and Opopsitta); less than 100 above vertical (most psittaciforms examined); 2 Width < length < 2 x width (most psittaciforms examined); 2 Sideways. Nares oriented laterally more than 300 from forward 3 Length > 2 x width ( papou, Probosciger. Nestor. Pioni- and less than 60" above vertical; (a) 00 above horizontal. 45-60" to tes. Purpureicephalus. Stri9oPS). side (Probosciger): (b) 300 above horizontal. 450 to side (Calyptorhyn- chus);

3 Upward. Nares oriented laterally less than 300 from forward and 0) Overall size (Fig. 1!) more than 60" above vertical: (a) 750 above horizontal. 300 to side Scoring of overall size is a somewhat subjective assessment of the (e.g. Agapomis); (b) greater than 750 above horizontal. 15-200 to side overall bulk of the rostrum, allowing for variation in shape. Speci- (Myiopsitta). mens were placed into categories of similarly sized rostra that had some discernable discontinuities between them. (G) Distocranial curvature (Fig. lG) Representatives of each category were used for comparison with newly examined taxa. No quantification of these categories was In lateral view. the curve of the culmen may be smoothly rounded. made. or it may have various degrees of flattening through either the cra- niaI or distal portion. 1 Very small (Melopsittacus, Neophema); 1993 A NEW COCKATOO FROM QUEENSLAND 17

2 Small (e.g. Glossopsitta. Nymphicus. Psephotus. Platycercus [smaller (K) Overallshape species and females of larger ones]); The bill of parrots is instantly identifiable as an ordinal character and is relatively uniform in gross appearancefor most taxa. Some 3 Small medium (e.g. Trichoglossus. Barnardius zonarius bamardi. species.however. have bills that are so substantially different that Platycercus caledonicus. Polytelis); these are one of the primary characters on which some genera are 4 Medium (e.g. Lorius. Cacatua [small species]. Alisterus. Myiopsitta); defined. This results from a combination of characters acting in concert. Other potentially useful charactersfor separatingtaxa have 5 Large medium (e.g. Callocephalon. Eclectus. Amazona. Nestor); not been evaluated becausethese are missing on the fossil. No at- 6 Large (Calyptorhynchus. Cacatua [large species]. Strigops); tempt has been made to score overall shape. but it is useful as a means to eliminate certain taxa from comparisons. 7 Very large (Probosciger, Anodorhynchus. Ara).

APPEN )IX II

Taxa for which the rostrum was examined Lathamus (discolor); Lonus (9arrulus. hypoinochrous); Melopsittacus (undulatus); Myiopsitta (monachus); Neophema (chrysostoma. ele9ans. Agapomis (fischerii. lilianae. personata. roseicollis); Alisterus (scapular- petrophila. pulchella. splendida); Nestor (mendionalis. nobilis); Northiel- is); Amazona (aestiva. albifrons. amazonica. autumnalis. Jarinosa, fin- la (haemato9aster); Nymphicus (hollandicus); Opopsitta (diophthalma); schi, leucocephala. ochrocephala, viridigenalis): Anodorhynchus (hyacin- Pezoporus (wallicus); Pionites (leuco9aster. melanocephala); Pionus (fus- thinus): (erythropterus): Ara (ambigua. ararauna, macao, cus. maximiliana. menstruus. senilis. seniloides); Platycercus (adscitus. aurico1lis, chloroptera, manilata. nobilis); Aratinga (aurea): Barnardius caledonicus. e. ele9ans. e. jlaveolus. eximius. icterotis. venustus); Poice- (z. bamardi. z. zonarius): Bolborhynchus (lineola): Brotogeris (jugularis. phalus (9ulielmi. meyeri. sene9alus). Polytelis (alexandrae. anthopeplus. versicoluris): Cacatua (alba, ducolpsii. galerita. haematuropygia, lead- swainsonii); Probosciger (aterrimus) Prosopeia (personata. tabuensis); beateri. moluccensis, ophthalmica. pastinator, roseicapi1la. sulphurea, Psephotus (haematonotos. varius); (fuscata); Psittacula (kra- tenuirostris): Ca1locephalon (fimbriatum): Calyptorhynchus (funereus. men); Psittaculirostns (demarestii. quilielmitertii); Psittacus (enthacus); lathami): (cardinalis. duivenbodei): Charmosyna (papou. (versicolor); Pu1pureicephalus (spunus); Pyrrhura (frontalis. pulche1la): Conuropsis (carolinensis): Coracopsis (vasa): Cyanoliseus (pa- picta); Rhynchopsitta (pachyrhyncha); Stn9opS (habroptilus); Tany- tagonus): Cyanoramphus (novaezelandiae cooki); Eclectus (roratus): En- 9nathus (lucionensis. me9alorhynchus); Touit (huetii); Tncho9lossus icognathus (ferrugineus); (bomea): FolpuS (cyanopygius, passerinus): (chlorolepidotus. 9oldei. haematodus moluccensis. h. rubntorquis). Geoffroyus (geoJfroyi); Glossopsitta (concinna, polphyrocephala. pusi1la):

APPENDIX III

Rostral characters of Australo-Papuan and selected occurs in only one species, the other members of the genus having extralimital genera of parrots the first score. A slash between two scores (e.g. 3/4) indicates that the state is intermediate between them and difficult to distinguish SeeAppendix I and Figure 1 for explanation of charactersand char- with confidence. A plus ( + ) or minus ( -) sign following a score acter states..Although some states are different stagesof a mor- indicates that it is at the respective end of the state, approaching phocline, overall no implication of relative polarity is intended or another. Square brackets ([ ]) indicate that the character could not should be inferred. When two scoresfor a single character are sep- be evaluated on the material available. arated by a dash (e.g. 2-3) or a comma (e.g. 4,6), it indicates that Australasian taxa not examined are Neopsittacus, Oreopsittacus, more than one state occurs among members of the genus. Paren- Eunymphicus, Geopsittacus. Loriculus, Micropsitta, Psittacella and Psit- thesesaround the secondscore (e.g.2(1» indicate that the condition trichas.

Rostral character

Genus A B c D E F G H

WRllDAE

Chalcopsitta 3 2B 1,3 I' 3B 1 2 2? 2 4

Chamlosyna 1 2B 2 I 3A 1 2 2+ 2/3 1-2 Eos 3 2B 3 I 3B 1 2 2 2+ 4

Glossopsitta 2B 1 3B 1 2 2 2 2 Lonus 1-2 2B 2/3 1 3B 1 2 2 2-3 4 Pseudeos 2 2A 2 1 3B 1 2- 2 2 3 Psitteuteles 2 2A 1 1 3A 2 2 2 2

Tnchoglossus I 2B 1 1 3B 1 2 2 2 3 (T. goldei) 2 2A 1 3B 1 1. 2 2 2 18 w. E. BOLES IBIS 135

Rostral character

) Genus A B c D E F Gj H

CACATUIDAE Cacatua 4 3A 5 3+-5 2A 1 1-2(4) 2(1) 4.6 Callocephalon 4 3B 5 5 lA 2B 1+ 2- 5 Calyptorhynchus 4 3B 5 4 1-2A 2B 1+ 2-(1) 6 Nymphicus 4 3C.o 5 2 3A 1 2 2 2 Probosciger 4 4* 2 5 lA 2A 1 4 3 7

PSITfACIDAE

Agapomis 2 2A 3 3 3A 3A 2 3 2 2 AIisterus 2 2A 2 3 3B 2 3 2 4 Amazona 3 2B 314-5 3C 1 1- 3 2 S Anodorhynchus 3 2A 1 3 lA 2 I 2 3 7 Aprosmictus 2 2A 3 2 3A+/B- 1 2 2/3? 2 3 Ara 3 1-2A. 2 3-4 lA 2 2-3 2-3 7 (A. manilata) 3 2A 3 3 2B 1 1- 1/2 2 4 Aratinga 3 2B 2 2 lA 2 2 3 2 2 Bamardius 2 2A. 314 2 3B 3 3 2 3.5 Bolborhynchus 2 2A 2 3 3A 1 I 2? 2 2 Brotogeris 2 2B 2 2 3A 1 3 3 2 Conuropsis 2 2B 3 4 2B 2A 1 3 2 4 Coracopsis 1+ 2B 2 3 3C 1 1 2 2 S CyanoIiseus 1 2A 3 3 lA lA 1- 3 2- 4 Cyanoramphus 2 2A. 2 2 3B 1 3 3 2 3-4 Eclectus 2 3 4 2A 1 1 3 2 S Enicognathus 2 2A 3 4 lA lA 1- 3 2 4 Fo1pus 1 2B 3 2 3B 1 1- 3 2 112 Geoffroyus 2 2A 4 3 3A 1 2 2-3? 2 4 Lathamus 2 2A* 3 2 3B 1 3 2 2 2 Melopsittacus 2 2A* 1 1 3A 1 3 2+ 2 1 Myiopsitta 2 2A 2-3 3+/4- lA 3A 1-/4 3 2 4 Neophema 2 2A* 4 1 3B 1 3 3 2 1 Nestor 2 2B* 3 3 3C 1 1 2 3 S Nolthiella 2 2A* 3 2 3A 1 3 3 2 2 Opopsitta 3 2A 2 1-2 3A 1 3 2- 2 Pezoporus 2 2B* 3 2 3B 1 3 3 2 2 Pionites 2 2B 3 3 1+A/2- 2 1-/2 3 2/3 4 Pionus 2 2A 3 2 3C 1 1 3 2 4 Platycercus 2 2B* 3 2 3B 1 3 3 2 2-3 Poicephalus 2 2A 3 3- 3 3A 2 3- 2 4/5 PolyteIis 2 2A 4 2 3B 1 1- 2 2 .l Prosopeia 2 2A* 4 3 3B 1 1 3 2 Psephotus 2 2A* 2 1 3A 3 3 2 Psittacula 2 2A 4 3 3A 3 2 3 2 4 PsittacuIirostris 2 2A 1 2 3-A 1 3 2- 214 Psittacus 3 2A* 3 3 3A 1 2 3 2 5 Pu1pureicephalus 2 2A* 3 2 3B 1 1- 3 3 Pyrrhura 3 2B 2 3 lA 2 2 3 2 Rhynchopsitta 3 4 3 4 lA 2A 2 3 2 Strigops 2 2B* 4 3 3C 1 2 2 3 Tanygnathus 2 2A 3 1.3 2A 3A 1 2 2 Touit 2 2A 3 3 3A 1 [] [] 2