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Revision of Dromaius Gidju Patterson and Rich 1987 Fr Om Riverslei G H, Nor Thwestern Q Ueensland , Australia, with a Reassessment of Its Generic Position

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Revision of Dromaius Gidju Patterson and Rich 1987 Fr Om Riverslei G H, Nor Thwestern Q Ueensland , Australia, with a Reassessment of Its Generic Position s~ REVISION OF DROMAIUS GIDJU PATTERSON AND RICH 1987 FR OM RIVERSLEI G H, NOR THWESTERN Q UEENSLAND , AUSTRALIA, WITH A REASSESSMENT OF ITS GENERIC POSITION Walter E. Boles ABSTRACT. Dromaiusgid;u Patterson and Rich 1987, from the Mio- representation in the fossil record, thus presenting virtually no cene Kutjamarpu local fauna of central Australia, was described on the clues about the divergence of the subfamilies. basis of associated hindlimb elements. New hindlimb material from the Based on fossil and Recent material, the Casuariinae and Dro- type locality and from Oligocene/Miocene deposits in Riversleigh, maiinae represent graviportal and cursorial birds, respectively, nonhwestem Queensland, are refened to this taxon. Tibiotarsal and with concomitant preferences for more densely vegetated and tarsometatarsal remains of D. gid;u are more similar to those of emus more open habitats. The differences listed in Table 1, taken than those of cassowaries, whereas the femora are more similar to femora of cassowaries. All hindlimb elemenrs of D. gid;u show some from Patterson and Rich (1987) and Rich et al. (1988), distinguish affinity to both cassowaries and emus. The combination of hindlimb cassowaries from emus for the elements available. The inter- characters indicates an earlier stage in the development of increased mediate, mosaic nature of the material described in this paper cursoriality, so I propose anew genus for this ta~on, Emuarius, which blurs some of these distinctions, with some elements exhibiting is interpreted to be nearer the mutual ancestor of the emu and cassowary more features characteristic of the Casuariinae, whereas others lineages than any other described fonD. are more similar to those of the Dromaiinae. The recently described Dromaius gidju Patterson and Rich Key words: fossil, emu, cassowary, Casuariidae, Dromaius, Casuarius, 1987 from central Australia has some characters that indicate Emuarius, Riversleigh, Australia. it was less cursorial than other emus, suggesting that it may h.ave been close to a mutual emu-cassowary ancestor. Patterson and INTRODUCTION Rich (1987:99) placed Dromaius gidju with the emus on the Emus and cassowaries {Casuariidae: Casuariiformes) comprise basis of available material, but they noted that ". ..as our one of the most distinctive and characteristic elements of the records of this fonn increase, there may be sufficient reason to Australo-Papuan avifauna {Fig. 1). I follow the practice of pat- separate it from both genera within this family as an early, quite terson and Rich {1987), Rich et al. {1988), and others of con- unspecialized fonn." New finds from Riversleigh, northwestern sidering Casuarius Brisson 1760 and Dromaius Vieillot 1816 Queensland, extend the knQwn geographic range of D. gidju as representing two subfamilies within the Casuariidae, rather and pennit a reassessment of the generic placement of this taxon. than as comprising separate families. Emus {Dromaius; Dro- maiinae) are represented by a single living species, D. novae- MATERIALS AND METHODS hollandiae {Latham 1790), which is restricted to the Australian mainland. Two dwad, insular species have become extinct in Names of species taxa for Recent Casuarius and Dromaius recent times: D. ater Vieillot 1817 [=D. minor Spencer 1906 follow Mayr (1979), except for the use of D. ater as the name auct. ] on King Island, in Bass Strait between Tasmania and the for the King Island population and recognition of the distinction mainland, and D. baudinianus Parker 1984, on Kangaroo of D. baudinianus, which follow Parker (1984). Measurements Island, off the South Australian coast {Parker, 1984). The tax- were made with vernier calipers accurate to 0.05 mm and round- onomic status of the extinct Tasmanian population of D. no- ed to the nearest 0.1 mm. Terminology primarily follows that vaehollandiae diemenensis Le Souef 1907 is unresolved. Cas- of Howard (1929, 1980). The holotype of D. gidju is in the sowaries {Casuarius; Casuariinae) are represented by three extant South Australian Museum. Other material referred to this taxon species: the Southern or Double-wattled Cassowary, C. cas- herein is currently held in the collections of the Vertebrate uarius Linnaeus 1758 {northeastern Australia, New Guinea, Ser- Paleontology Laboratory, University of New South Wales, but am-introduced?), Single-wattled Cassowary, C. unappendicu- it will be transferred to either the Australian Museum or the latus Blyth 1860 {New Guinea), and Bennett's or Dwad Queensland Museum. Initial citations of fossil material give the Cassowary, C. bennetti Gould 1857 {New Guinea, New Brit- registration numbers of both the Vertebrate Paleontology Lab- ain-introduced?) {Mayr, 1979). Despite forming a distinctive oratory and respective museum; thereafter, only the museum and endemic or near endemic part of the Australo-Papuan avi- number is cited. The following abbreviations are used: AM, fauna, the family Casuariidae has had, until recently, only a small Australian Museum; AR, Vertebrate Palaeontology Laboratory, ~C. CASUARIUS 0 ~ ~ the genus. Additionally, digits II and IV were less reduced. These .C. UNAPPENDICULATUS ~ authors interpreted this as an indication that D. gidju was the least cursorial member of the genus. ~ C. BENNETTI 6 A small emu from the Late Miocene is represented in the m D. NOVAEHOLLANDIAE Alcoota local fauna, Northern Territory (Patterson and Rich, 1987). It is similar in size to Dromaius gidju and possibly re- E] D N. DIEMENENsiS ferable to that taxon, but because the remains are scanty (just .D. ATER fragmentary tarsometatarsi and phalanges) Patterson and Rich ID D. BAUDINIANUS (1987) assign.ed it to Dromaius sp. indet. pending discovery of additional material. CASUARIINAE. Lydekker (1891) reported a small cassowary from the Pleistocene of New South Wales based on the distal end of aright tibiotarsus; it was subsequently named Casuarius lydekkeri Rothschild 1911. This fossil was presumed to have come from the Wellington Caves. However, the manner of preservation is not matched by other material from these de- posits, and several workers have expressed doubt abo~t the presumed locality (Rich et al., 1988). Plane (1967) reported three phalanges from the Pliocene Awe local fauna, Papua New Guinea, without referring them to a species. Although closest to Casuarius bennetti among living .Riversleigh species, there were sufficient differences in the shape and pro- .Lake Ngapakaldi (Kutjamarpu) , pOrtions of the bones to doubt that they were from this species. Rich et al. (1988) described a pygmy cassowary from Pleis- .Pureni tocene bog deposits at Pureni, Papua New Guinea, that cannot ...Awe be separated from Casuarius lydekkeri on the basis of the distal ..Wellington Caves end of the tibiotarsus, the only shared element. Consequently, those authors referred all the Pureni material to this species, noting that, although it shares several features with C. benneUi, Figure 1. Distribution of Recent species of Dromaius and Casuarius in other ways it more closely resembles C. casuarius. Lack of and casuariid fossil-bearing localities mentioned in the text. common elements precluded comparison with the Awe casso- wary. NEW CASUARIID MATERIAL. New material from Riv- University of New South Wales; CPC, Commonwealth Palaeon- ersleigh includes fragments of lower limb bones (femur, tibio- tological Collections, Bureau of Mineral Resources, Canberra; tarsus, tarsometatarsus), a scapulocoracoid, and a partial maxilla, QM, Queensland Museum; SAM, South Australian Museum; all referred to Dromaius gidju. A small femur referred herein and UCMP, University of California, Museum of Paleontology, to this family was collected from the Leaf Locality, the type locality for D. gid;u. This new material and its bearing on the Berkeley. relationships of D. gid;u form the subject of this paper. FOSSIL HISTORY GEOGRAPHIC AND GEOLOGICAL SE1TING The fossil histories of the emus and cassowaries have recently been reviewed by Patterson and Rich (1987) and Rich et al. The material described as Dromaius gidju, and an additional (1988), respectively, thus necessitating only brief synopses here bone reported herein, came from the Leaf Locality {UCMP (see also Rich and Baird, 1986). V-6213), a site on the eastern shore of Lake Ngapakaldi in the DROMAIINAE. Of the nominal forms of emus known from eastern Lake Eyre sub-basin, South Australia. The Leaf Locality fossil remains, Patterson and Rich (1987) maintained two. After faunal assemblage including this taxon is the Kutjamarpu local eliminating several fossils incorrectly assigned to the subfamily fauna. The age of this fauna is suggested by its diprotodontid Dromaiinae, these authors recognized Dromaius ocypus Miller marsupials, whose closest affinities lie with those from the Riv- 1963 from the Pliocene Palankarinna local fauna of the Lake ersleigh and Bullock Creek local faunas. The latter are under- Eyre basin, South Australia, and referred all other determinable stood to be between Late Oligocene and Middle Miocene in mainland material, including D. patricius De Vis 1888 and D. age {Archer et al., 1989). gracilipes De Vis 1892, to the modern D. novaehollandiae. They The Riversleigh deposits are located on Riversleigh Station, noted that there may have been greater variability in body size approximately 200 km north of Mount Isa, northwestern in Pleistocene emus than at present, but confirmation awaits Queensland. Only one of the sedimentary units has been given further material of both fossil and Recent
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