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Warkaiania, Anew Meiolaniid Turtle from the Tertiary Riversleigh Deposits of Queensland, Australia

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Warkaiania, Anew Meiolaniid Turtle from the Tertiary Riversleigh Deposits of Queensland, Australia The Beagle, Records of the Northern Territory Museum of Arts and Sciences, 19929(1):35-48 WARKAIANIA, ANEW MEIOLANIID TURTLE FROM THE TERTIARY RIVERSLEIGH DEPOSITS OF QUEENSLAND, AUSTRALIA. EUGENE s. GAFFNEYI, MICHAEL ARCHER2 AND ARTHUR WHlTE2 IDepartment of Vertebrate Paleontology, American Museum of Natural History, Central Park West at 79th St, New York, NY, USA 10024-5192. 2University of New South Wales, P.O. Box 1, Kensington, NSW 2033, Australia. ABSTRACT Warkalania carinaminor sp. nov., is based on posterior skull elements from the (?) Oligocene -early Miocene Carl Creek Limestone, Pancake Site of Riversleigh Station, northwest Queensland. Warkalania is a meiolaniid cryptodire because it has the synapomorphies of the squamosal produced into posteriorly and posterolaterally directed processes and an extensive squamosal-quadratojugal contact beneath the cavum tympani. Within meiolaniids, Warkalania is unique in having the squamosal processes formed into low, horizontal ridges, small and roughly similar to each other in size, rather than having one or more of these scale areas extending prominently away from the skull, as in all other meiolaniids. KEYWORDS:Warkalania carinaminor gen et sp. nov., new genus, new species, Meiolaniidae, horned turtle, Oligocene, Miocene, Carl Creek Limestone, Queensland. INTRODUCnON The reader should refer to Figures 5- 7 and Gaffney (1983) for scale terminology of Turtles and other vertebrates were discovered meiolaniid skulls. Descriptions of the skull of in the Carl Creek Limestone at Riversleigh Sta- Meiolania platyceps Owen can be found in tion in northwestern Queensland by Tedford in Gaffney (1983). Institutional prefixes to cata- 1968. Further work by teams led by M. Archer, logue numbers are as follows: AMF, Australian S. Hand, and H. Godthelp resulted in the discov- Museum, Sydney; BMNH, British Museum of ery of nearly 100 additional sites spanning in Natural History, London; QMF, Queensland time the Oligocene to Pleistocene, and the recog- Museum, Brisbane. nition of a relatively rich turtle fauna (Gaffney et al. 1989). A review of the Riversleigh localities and an introduction to the growing literature on SYSTEMA TICS Riversleigh and its fossils can be found in Archer et al. ( 1989). The first turtles described from Order Testudines Riversleigh Tertiary sites were three genera of Megaorder Cryptodira chelids based on fragmentary material (Gaffney Capaxorder Selmacryptodira et al. 1989). Meiolaniids in the Riversleigh Hyperorder Daiocryptodira Tertiary were first recognized by A. Ritchie Parvorder Eucryptodira (pers. comm. ) on the basis of tail ring fragments Suborder Meiolanoidea (seebelow) in 1987. One of us(ESG)found more Family Meiolaniidae meiolaniid fragments in Riversleigh collections in 1989 but it was not until Neville Pledge's Type Genus. Meiolania Owen, 1886b. discovery of a partial skull, that diagnosable Known Distribution. Pre-Oligocene ("Cre- meiolaniid material was recovered from taceous or Eocene", Simpson 1938) of Argen- Riversleigh Tertiary deposits. This material rep- tina, Miocene to Pleistocene of Australia, resents anew genus of meiolaniid (Fig. 1) and is Pleistocene (or younger) of Lord Howe Island, described here. Walpole Island, and New Caledonia. 35 E.S. Gaffney, M. Archer and A. White Previous work. Gaffney (1983) relates the surrounded by dennal ossifications; tail club long and complex history of work on the fonned by fusion of tenninal caudal vertebrae meiolaniids, beginning with Owen's ( 1881) iden- and osteodenns (at least in Ninjemys oweni and tification of the first known meiolaniid as a giant Meiolania platyceps); cervical formula is homed lizard. Recent papers are Megirian ( 1989, (2«3«4»5»6»7»8); free cervical ribs present 1992), Gaffney and McNamara (1990), and on cervicals 2-6; caudal vertebrae opisthocoelus Gaffney (1992). with well developed haemal spines; plastron Revised Diagnosis. Eucryptodiran turtles with lacking axillary and inguinal buttresses; no the squamosal and supraoccipital uniquely pro- mesoplastra present; plastron with fontanelles duced into posteriorly and posterolaterally di- on midline; first thoracic of the carapace facing rected processes, three scale areas (A,B,C in anteriorly, and first thoracic rib long and reach- Gaffney, 1983) being most prominent; temporal ing plastron laterally; posterior peripherals scal- emargination completely absent and related to loped; adults usually with cranial and shell su- extensive squamosal-supraoccipital contact and tures fused; carpal and tarsal fonnula 2-2-2-2-2. relatively small parietal; supraoccipital with large It should be noted that most of these characters horizontal plate on skull roof; nasal bones unu- are known only in M eiolania platyceps ( Gaffney , sually large; sinus formed from nasal and max- 1983, 1985). illa lateral to, and communicating with, apertura Discussion. The higher phylogenetic rela- narium extema (determinable only in Meiolania tionships of meiolaniids and the higher classifi- platyceps and Ninjemys oweni); broad squa- cation of cryptodires is discussed in Gaffney mosal-quadratojugal contact ventral to com- (1983), Gaffney and Mey1an (1988), Gaffney et pletely enclosed incisura columellae auris which al. (1991), and Gaffney (1992). contains both stapesand eustachian tube; medial plate of pterygoid separatedventrally from basi- sphenoid to form intrapterygoid slit; palate con- Warkalania gen. nov. cave ventrally with vomerine ridge on midline; well developed Iabial and Iingual ridges not Type Speci~ Warkalania carinaminor sp. nov. greatly expanded; tail partially or completely Known Distribution. Mid Tertiary , north- west Queensland. Diagnosis. Meiolaniid with B scale area ( en- compassing the largest horn in other meiolaniids ), delimiting a low, horizontal ridge, not a recurved horn as in Meiolania nor a large lateral projec- tion as in Niolamia (including Crossochelys) and Ninjemys oweni; A, B, and C scales formed into low, horizontal ridges, small and roughly similar in size to each other, in contrast to all other meiolaniids; A scale area not as protuberant as in Niolamia and Ninjemys oweni but more protu- berant than in Meiolania; C scale area a low ridge, higher than in Meiolania but lower than ir. the other meiolaniids; X scale small, and D scales in midline contact as in Ninjemys oweni and Meiolania. Etymology. Warka, Queensland aboriginal for turtle; lania, in reference to the usual endings for meiolaniids (Lanius is Latin for "butcher" but Owen (1886b) gave no indication of an etymol- ogy when he erected Meiolania). Warkalania carinaminor sp. nov. Fig I. Warkalania carinaminor ~n. et sp. nov. Dorsal view of skull based on QMF 22649, 22650, and 22651, partially Type material. HOLOTYPE QMF 22649, a restored by transferring side to side. See Figure 5 for scale right squamosal (Fig. 2). This fragment includes labels. the posterior margin of the cavum tympani, all of 36 Miocene homed turtle scale c, most of scale K, and parts of scales B,H, 22654 consist of the same bone elements. The and D. It is likely, but not definitely demonstra- remaining fragments, QMF22649, 22650,22651, ble, that QMF 22650-22653 and 22682 belong to 22652, 22653 and 22682, come very close to the sanle individual. actual contact, show no overlap, and can be Type locality. Pancake Site, Riversleigh sta- restored as a reasonable skull from one indi- tion, northwestQueensland, see Archer et aL ( 1989). vidual. However, the type specimen chosen, Horizon. .'? late Oligocene to early Miocene" QMF 22649, has enough diagnostic characters Archer et at. (1989:64). so that it can stand alone as a new taxon of Other material. Referred specimens, all from meiolaniid, even if the composite reconstruction the type locality: QMF 22650, left sqUanlosal is in error due to the mixing of more than one with complete scale areasA,B, and C plus part of individual and more than one species. The scale D (Fig. 3). A fragment of the posterior wall holotype QMF 22649, shows the large squa- of the antrum postoticum is preserved internally; mosal, enclosed incisura columellae auris, and QMF 22651, central section of right and left development of protuberances identifiable as B parietals, bearing on its dorsal surface scale X and C horns, diagnostic of Meiolaniidae. But the and surrounding portions of scales G and D; Band C horns of QMF 22649 differ significantly QMF 22652, nearly complete right quadrate; from all other meiolaniids. The C horn is cone QMF 22653, right exoccipital and basioccipital shaped (or flat in Meiolania} in all other lacking ventral surface; QMF 22682, part of meiolaniids, but in Warkalania it is a horizontal right supraoccipital containing semicircular ca- ridge directed anteroposteriorly. The B scale of nals, small portions of prootic and opisthotic; QMF 22649 is also a flattened ridge in strong QMF 22654, left sqUanlosal with scale areas B contrast to the cow-like, recurved B horn core and C complete portions of scale areas A and K synapomorphic for Meiolania. The B horn cores (Fig. 4). ofNinjemys oweni and Niolamia are much larger, Discussion. An important question to be dealt laterally directed spines, rather than the rela- with in the proposal of Warkalania as anew tively low, horizontal ridge of Warkalania. genus of meiolaniid, is: do the seven skull frag- The other cranial fragments identified here as ments actually belong to one species or one belonging to Warkalania allow the diagnosis to specimen? It is apparent that at least two indi- be extended to the X scale area,
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