B.P. KEAR, M. ARCHER & T.F. FLANNERY

KEAR, B.P., ARCHER, M. & FLANNERY, T.F., 2001:12:20. Postcranial morphology of bilamina Cooke, 1997 (Marsupialia: ) from the middle of Riversleigh, northwestern Queensland.Memoirs of the Association of Australasian Palaeontologists 25, 123-138. ISSN 0810-8889

Postcranial remains of the early to middle Miocene bulungamayine Ganguroo bilamina Cooke, 1997 are described. This is one ofonly two reports (see also Kear et al. this volume) of bulungamayine postcranials and describes some of the oldest postcranial material known for macropodoids. Functional analysis suggestsa hopping gait, though the well developed forelimbs and scapular morphology indicate consistent use of more quadrupedal progression. Phylogenetic analyses using postcranial characters place G. bilamina as the sister taxon to macropodids. This supports the conclusions of Case (1984), Woodburne (1984), Cooke (1997a, b), Kear (1998), Cooke & Kear (1999) and Kear & Cooke (this volume) in advocating a revised position for Bulungamayinae.

B.P. Kear {kea7:[email protected]).1:F. F/annery. South Austra/ian Museum. Ade/aide. South Austra/ia, 5000; M. ArcheI; Vertebrate Pa/aeont%gy Laboratory. Schoo/ of Bi%gica/ &iences. University of New South Wa/es.Sydney, New South Wa/es.2052 and Austra/ian Museum, 6-8 Co//ege St. Sydney.New South Wa/es.2000. Received 30 May 2000

Keywords:Marsupialia, Macropodidae, Ganguroo, Miocene, Riversleigh, Queensland

A PARTIAL juvenile skeleton and skull of the predominantly quadrupedal bulungamayine kangaroo Ganguroo bilamina moschotus (Johnson & Strahan 1982), however, Cooke, 1997was collected from AL90 Site, System suggests consistent use of more quadrupedal Cdeposits(Archeretal.1994;Archeretal.1997; modes of progression. This paper describes Black 1997).This material, along with that reported postcranial remains of G. bilomino and analyses by Kear et al. (this volume), is the only definite bulungamayine phylogeny and functional postcranial remains for bulungamayines and some anatomy using postcranial features. of the oldest known for macropodoids. Skeletal and myological terminology follows Bulungamayines are small to large extinct Elftman (1929), Hopwood (1974), Hopwood & kangaroosknown from the late Oligocene to early Butterfield(1976, 1990), Murray (1995), Wells& late Miocene. Flannery et al. (1983) placed them Tedford (1995) and Bishop (1997). The as the sister group to potoroines. Case (1984), abbreviation QMF refers to the fossil collection Woodbume(1984), Cooke (1997a,b), Kear(1998), of the Queensland Museum, Brisbane. All Cooke & Kear (1999) and Kear & Cooke (this measurements were made using digital callipers volume), however, suggestedmacropodid affinity and are in millimetres (mm). on the basis of synapomorphies present in G. bilamina, other derived bulungamayines and SYSTEMAD C pALAEO NfO LOGY plesiomorphic macropodids such asDorcopsoides jossilis, of and SuperfamilyMACROPooomEAGrey,1821 and Hadronomas puckridgi. The possibility of Family MAcRoPODmAEGrey, 1821 bulungamayine paraphyly (Cooke 1997a, b) or Subfamily BULUNGAMAYINAE Flannery, polyphyly (Kear 1998; Kear et al. this volume) Archer & Plane, 1983 has also been proposed. The postcranial anatomy of G. bilamina is GenusGanguroo Cooke, 1997 similar to that of potoroines and macropodids in exhibiting features indicative of hopping gaits. Typespecies. Gonguroo bilomino Cooke, 1997 Forelimb morphology comparable to the 124 AAP Memoir 25 (200 1)

CERVICAL VERTEBRAE METACARPAL Centrum height: 2.59 Maximum length: 5.87 Centrum width: 4.33 Maximum proximal width: 2.02 Body height (inc. neural spine): 9.34, 9.38, 9.58, Minimum shaft width: 1.47 9.87,10.48,10.71 DISTAL MANUAL pHALANGE Width across postzygapophyses: 9.47, 10.04, Maximum length: 4.16,4.08 10.46,10.67,10.87,10.91 Ungual process width (at base): 1.61, 1.54 moRACIcVERTEBRAE Ungual process depth (at base): 0.73,0.78 Centrum height: 3.16,3.22,3.25,3.35,3.57,3.63, n.IUM 4.16 Maximum ilial blade length: 28.33 Centrum width: 4.3,4.4,4.49,4.57 ,5.38, 5.39,6.18 Minimum ilial blade width: 8.05 Width across postzygapophyses: 5.53, 5.58, 6.01, Distal apex height: 10.79 6.12,6.19,6.37,6.59,7.95,8.86 Total length: 38.98 LUMBARVERTEBRAE FEMUR Centrum height: 4.08, 4.87 Maximum distal epiphysis width: 13.06 Centrum width: 6.37,6.51 Intercondylar notch width: 3.92 Maximum height: 11.66,13.62 TIBIA Width across postzygapophyses: 6.18, - Shaft circumference: 20.29 CAUDAL VERTEBRAE Proximal epiphysis width: 11.66 Centrum height: 1.43,4.29,4.96 Intercondylar eminence height: 3.1 Centrum width: 2.41,5.31,6.02 F1BULA Maximum length: 7.07,10.18,13.57 Distal epiphysis width: 6.3 Minimum width: 1.09,3.24,3.8 Distal epiphysis ant./pos. length: 3.41 SCAPULA Peroneal tendon sulcus width: 2.17 Glenoid fossa diaphysis height: 5.15 NAVICULAR Glenoid fossa diaphysis width: 3.46 Maximum height: 5.86 Glenoid neck height: 5.99 Dorsal surface length: 3.81 Maximum blade height: 14.17 Plantar surface length: 5.24 Supraspinous fossa height: 4.31 Astragalar facet height: 5.21 Infraspinous fossa height: 7.84 Astragalar facet width: 2.19 HUMERUS ECTOCUNEIFO RM Maximum distal width: 9.31 Maximum height: 7 .25 Supinator crest height: 8.86 Plantar process length: 2.77 Supinator crest width: 1.19 Dorsal surface length: 4.17 Deltoid crest length: 16.78 Navicular facet height: 4.31 Total length: 33.39 Navicular facet width: 2.77 ULNA MetatarsalIIl facet height: 3.42 Width across coronoid process: 3.46 MetatarsalIIl facet width: 1.36 Olecranon process depth: 3.91 Total length (estimated): 36.74 RADIUS Table 1. Measurements of postcranial elements included Distal facet height: 3.24 in this study. (all measurements in mm). Distal facet width: 4.8 Total length {estimated): 36.4

Age and distribution. Early to middle Miocene of vertebrae2-9, 12 and 13; lumbar vertebrae 1 and 2; northern Australia. caudal vertebrae 3 and 5; ribs left and right 1-12; left and right clavicle; manubrium and sternebra Diagnosis. As for G. bilamina until postcranial 5; fragmentary left and partial right scapula; material of other species recovered. complete left and right humerus; partial left and right ulna; partial left radius; left metacarpal II; Ganguroo bilamina Cooke, 1997 two distal manual phalanges; right ilium; right distal femoral epiphysis; left proximal epiphysis

Holotype. QMF19915, leftdentary. and distal section of tibia (lacking epiphysis); left distal fibula; complete left navicular and Additional Material. Additional material includes ectocuneiform. an associatedjuvenile partial skeleton and skull {skull not describedhere) QMF30845. Postcranial Supplemental Diagnosis. Lumbar vertebrae with elements include: cervical vertebrae 1-6; thoracic strong mid-ventral keel; scapula broadly AAP Memoir 25 (200 1) 125

view; B, posterior view. C, D, axis; C, anterior view; D, lateral view. E, F, cervical vertebra; E, anterior view; F, lateral view. G, H, thoracic vertebra; G, anterior view; H, lateral view. I, J, lumbar vertebra; I, anterior view; J, lateral view. K, L, anterior caudal vertebra; K, anterior view; L, lateral view. M, posterior view of clavicle. Scale bar is 10 Inm.

rectangular in outline with supraspinous fossa coronal depressiondeeply excavated and strongly equal in length to infraspinous; acromion process delineated; capitellum and trochlea unevenly reduced and rounded; humerus straight and sized; ulna robust with straight ventral edge; slender with reduced supinator and long, distally olecranon process well developed and bears low deltoid crests; lateral teres tuberosity poorly distinct transversedorsal crest; distal apex of ilium developed; entepicondyle closely abuts trochlea; dorsoventrally elongate and transversely 126 AAPMemoir25 (2001)

CVA SSF

CB BCP ./ AP \ -,- \, ) Fig. 2. Scapulafragment of juvenile Ganguroo bilamina Cooke, 1997 (QMF30845). Scale bar is 5 Inm.

MPI SSP ISF compressed; navicular large and crescentic with GVA well developed plantar eminence; astragalar facet TMA GB on navicular transversely broad; Fig. 3. Reconstructed scapula (lacking epiphysis) of juvenile Ganguroo bilamina Cooke, 1997 (QMF30845). Additional material locality distribution and Scale bar is 10 mm. Abbreviations: AP, acromion age. AL90 Site, southernsection Gag Site Plateau, process; BCP; base of coracoid process; CB, coracoid border; CVA, coracovertebral angle; OB, glenoid border; Riversleigh World Heritage area, northwestern OVA, glenovertebral angle; ISF, infraspinous fossa; MP, Queensland.AL90 Site is correlated with System metacromion process; SSF, supraspinous fossa; SSP, C (early middle Miocene) deposits (Black 1997). scapular spine; TMA, triceps muscle attachment. Type locality is Wayne's Wok Site (System B, early Miocene; Cooke 1997c). Paratype and additional with tall neural spines becoming shorter and more cranial/dental material is known from inclined in successive vertebrae. Transverse Bitesantennary Site, System NB and Wayne's processes robust with costal fovea shallowing Wok, Camel Sputum,Mike's Menagerieand Upper posteriorly and being lost in fourth or fifth sites, all SystemB (Archeretal. 1994). vertebra. First thoracic lacks transverse foramen. Pre/postzygapophyses prominent, becoming Description larger and more elevated in posterior vertebrae. Axial skeleton Parapophyses on fIrst thoracic subtriangular and Cervicalvertebrae(Fig.lA-F, Table I). Complete well-developed, becoming reduced in posterior axis, atlas and cervicals 3-6 preserved. Centrum vertebrae. Neural canal arched and wider than tall, from second or third cervical present. Atlas tall becoming narrower posteriorly. Centra deep and and arched in anterior view; ventral arch not slightly opisthocoelous; weak rnid-ventral keel closed by neurapophyses. Condylar cusps present on all thoracics. reniform and transversely narrow. Transverse processesshort, spatulate and anteriorly inclined. Lumbar vertebrae (Fig. li-J, Table 1). Represented Transverse foramen small and laterally opening. by lumbars 1 and 2 only. Transverse processes Axis transversely narrow and tall with massive, present only as a low crest in lumbar 1, becoming lobate and anteroposteriorly elongateneural spine. spatulate and subhorizontally oriented in lumbar Neural arch tall and transversely narrow. Prezyg- 2. Anapophyses moderately large and lobate; apophyses ovoid; odontoid process short and metapophyses very weak. Pre/postzygapophyses dorsally inclined. Postzygapophysestransversely large and laterally flaring. Prezygapophyseswith narrow and subhorizontal; transverse foramen indistinct and flat articular facets. Neural spine laterally opening. Cervical vertebrae antero- moderately tall. Sulci for ligamentum flavium very posteriorly short and dorsoventrally tall. Neural weakly developed. Centra slightly opisthocoelous canal highly arched,becoming lower in successive with distinct mid-ventral keel. vertebrae. Neural spines tall and tapering at apex. Pre/postzygapophyses subovoid and slightly Caudal vertebrae (Fig. lK-L, Table 1). concave. Centrum platycoelous. Represented by caudals 3 and 5 only. Vertebral body ovoid in cross-section and dorsoventrally Thoracic vertebrae (Fig. 1G-B, Table 1). Thoracic tall. Low neural canal present in both vertebrae. vertebrae 2-9 and 12, 13 preserved. Eight unfused Third caudal exhibits large pre/postzyg- thoracic centra present. Anterior thoracics low apophyses; fifth caudal exhibits large prezyg- AAP Memoir 25 (2001) 127 apophyses only. Transverse processes of third distal trochlea with widely spaced capitulum and caudal narrow and wing-like; more trochlea. Capitulum globular and larger than the anteroposteriorly elongate and extending the full more cylindrical trochlea. Medial and lateral length of centra in caudal five. epicondylae reduced. Olecranon depression low, wide and weakly defined dorsally. Coronoid fossa Ribs. Left and right ribs 1-12 present. Preservation deep and extensive, separated from shallower ranges from complete to fragmentary. Rib shaft trochlear depression by distinct inter-fossa ridge. anteroposteriorly compressed being thicker Entepicondylar foramen dorsoventrally long and towards vertebral extremity. Tuberculae anteroposteriorly narrow. Entepicondylar buttress subcircular and posteriorly offset in posterior ribs. robust, D-shaped in cross-section. Capitulae spatulate and ovoid in outline, being anteroposteriorly elongate. Ulna (Fig. 4D-F, Table I). Left and right ulnae present, left lacking most of shaft, right Clavicle and Sternebrae (Fig. IM, Table 1). represented by distal extremity only. Ulna large Complete left and right clavicle present; with slender shaft and straight ventral edge. manubrium and penultimate sternebrae present. Proximal extremity deep with a broad, shallow Clavicle large with anteroposteriorly compressed depression for insertion of m. flexor digitorum shaft. Sternal extremity spatulate and flattened; profundus. Olecranon process large, being external surface slightly concave. Vertebral dorsoventrally tall and antero-posteriorly short extremity is a subtriangular lobe with dorsal edge when ulna is horizontally oriented. Process base bearing an elongate oblique costal fovea facet. transversely broad, narrowing dorsally and Coracoclavicular ligament tubercle reduced to a terminating in a distinct dorsal crest. Semilunar raised crest on ventral edge of vertebral extremity notch medially pinched; capitular facet open and lobe. Manubrium (not figured) kite-shaped with separated from narrow trochlear facet by a low prominent posterior facet for articulation of ridge. Radial facet narrow and also separatedfrom sternebra. Penultimate sternebrae (not figured) trochlear facet by a low ridge. Capitular and radial short and distinctly waisted. Articular facets facets strongly laterally sloping with low, weak rectangular in outline. buttressing. Coronoid process tall and laterally flaring. Anconeal process less distinct and Appendicular skeleton anteroposteriorly thick. Subtriangular rugose Scapula (Fig. 2,3, Table I). Material consists of insertion for m. brachialis and gleno-ulnar right glenoid (lacking epiphysis) and anterior divisions of the m. biceps brachii present scapular spine, and fragmentary left scapular immediately anterior to the coronoid process. blade. Scapular blade (Fig. 3) subrectangular in Depression for radial bursa elongate and outline with obtuse glenovertebral angle and indistinct; interosseous ridge long and low. straight glenoid border; coracoid border inclined; coracovertebral angle near perpendicular; Radius (Table 1). Left radial shaft preserved.Lacks supraspinous and infraspinous fossae extensive; distal epiphysis, proximal articular facet and neck. supraspinousfossa equal to infraspinous in length Radial shaft D-shaped in cross-section over much though dorsoventrally narrower posteriorly. of its length becoming more dorsoventrally Anterior portion of scapular spine deep and flattened distally and circular proximally. The rounded; acromion process poorly developed; interosseous ligament groove is well developed metacromion process dorsoventrally deep and and extends along the flattened proximolateral semicircular in outline. M. triceps attachment surface; the crest-Iike ventral border of the groove deeply hollowed. Glenoid fossa and coracoid representsthe interosseousmembrane attachment process broken away in specimen. (whichjoins the radial and ulnar shafts together). Adjacent to the interosseous ligament groove and Humerus (Fig. 4A-C, Table I ). Both left and right situated on the dorsolateral surface is an elongate, humeri preserved, left lacking distal trochlea. arched depression for attachment of the m. Humeral shaft straight and slender; supinator crest adductor digit I longus. Distal apex of the radial weakly developed and low; deltoid crest long, low shaft is both broad and deep relative to shaft and blade-like. Terestuberosity weakly developed. length; distal facet surface trapezoidal in outline. Glenoid large and globular; epiphysis missing. Distal ulnar articulation forms shallow lateral Greater and lesser trochlea transversely narrow; groove. 128 AAPMemoir25 (2001) AAP Memoir 25 (2001) 129

Fig. 5. Hindlimb elements ofjuvenile Ganguroo bilamina Cooke, 1997 (QMF30845) A, B, ilium; A, lateral view; B, ventrolateral view. C, plantar view of distal femur epiphysis. D, dorsal view of proximal tibia epiphysis. E, anterior and posterior views of distal fibula. Scale bar is 10 mm.

Metacarpals (Table I ). Left metacarpal II present Pelvis (Fig. SA-B, Table I). Represented by only; lacks distal epiphysis. Metacarpal shaft oval complete right and distal extremity of left ilium, in cross-section, dorsoventrally compressed and both unfused. Ilium outwardly flared distinctly waisted. Proximal trapezius facet (approximately lor from vertebral plane); trapezoidal in outline, tapering plantarly and sacroiliac articulation situated far back on ramus. shallowly concave. Metacarpal I and III facets Distal ilial apex spatulate, being dorsoventrally broad and triangular; bordered dorsally by low elongate and transversely compressed.Iliac blade ridges. Proximal plantar surface raised into a broad with low anteriorly flattened iliac crest. M. rounded node for interosseous ligament gluteus medius/m. gluteus profundus portion of attachment. blade shallowly concave, becoming slightly narrower anteriorly; m. iliacus portion narrow and Distal manual phalanges (Table 1). Two complete flattened becoming broader anteriorly. M. rectus distal manual phalanges preserved. Ungual femoris scar forms prominent triangular rugosity process elongate relative to plantar process immediately anterior to acetabulum border. length. Process slightly downcurved and Iliopectineal process large and subcylindrical. dorsoventrally flattened with oval cross-section; elongate and parallel sided in dorsal view with Femur (Fig. 5C, Table I). Represented by right tapered tip. Posterior articular facet shallow and distal epiphysis only. Condyles smooth and oval in outline; borders produced into laterally separated by a deep U-shaped intercondylar flaring crests. Plantar process oval in outline and fossa. Lateral condyle transversely broad and transversely narrow. ventrally flattened; medial condyle transversely narrow and rounded ventrally. Anterior condylar crests widely spaced and low.

Fig. 4 (opposite). Forelimb elements ofjuvenile Ganguroo bilamina Cooke, 1997 (QMF30845). A-C, humerus; A, anterior view; B, lateral view; C, posterior view. D-F, ulna; D, lateral view; E, dorsal view; F, medial view. Scale bar is 10 mm. 130 AAPMemoir25 (2001)

Fig. 6. Tarsalelements of juvenile Ganguroobilamina Cooke, 1997 (QMF30845)A, B, navicular;A, lateral view; B, posteriorview. C, D, ectocuneiform;C, lateral'view; D, anteriorview. Scalebar is 5 Inm. nbia (Fig. SD, Table 1). Represented by right transversely compressed and dorsoventrally tall; distal shaft section lacking epiphysis, and left plantar tuberosity well developed and oval. proximal epiphysis. Distal shaft section oval in Navicular facet dorsoventrally elongate, renifofll1 cross-section, with flattened medial edge for and concave. Metatarsal III facet small, fibula contact. Proximal epiphysis with dorsoventrally elongate and crescentic. Cuboid transversely broad, flat and circular, lateral articulation fofll1ed by a low lateral crest. condylar surface; medial condylar surface transversely narrow and concave. Intercondylar COMPARISONS AND FUNCTIONAL eminence tall with wide transverse ligament INTERPRETAllON attachment. Depression for anterior cruciate The following are functional interpretationsand ligament shallow and indistinct. Shallow posterior comparisonsof the postcranial elementspreserved cruciate ligament depression present on medial for Ganguroo bilamina with other macropodoids. edge of intercondylar eminence. DefInitions of macropodoid gaits follow Windsor & Dagg (1971) and are as outlined in Kear et al. Fibula (Fig. SE, Table I ). Left distal fibular shaft (this volume). and epiphysis present. Shaft slender and D- shaped in cross-section. Lateral peroneal tendon Vertebrae(Fig. IA-L, Table I ). Cervical vertebrae groove shallow; bordered anteroposteriorly by in G. bilamina resemble those of sthenurines low tuberosities for attachment of superior (Wells & Tedford 1995) in the presence of a peroneal retinaculum. Peroneal tendon groove massive neural crest on the axis and tall terminates in a circular pit. Medial malleolar fossa successive neural spines. Elongation of neural of distal epiphysis shallowly inclined and slightly spines is associated with enlargement of the concave. Anterior edge of epiphysis produced nuchal ligament and epaxial muscles, increasing into a rounded node. strength and mobility in the neck (Finch & Freedman1986). Lengthening of the cervical neural Navicular and ectocuneiform (Fig. 6, Table I). spines is also associated with increased Complete left navicular and ectocuneiform attachment area for the m. trapezius (Wells & recovered. Navicular (Fig. 6A, B) transversely Tedford 1995),a major manipulator of the forelimb. compressed and crescentic in lateral view. This indicates increased strength in the forelimb Astragalar facet transversely wide and for either locomotion (providing forelimb extension dorsoventrally elongate; facet occupies entire during the propulsive stroke) or browsing posterior surface of navicular. Posteriormost facet (elevating the limb ). extremity slightly medially offset. Plantar The thoracic vertebrae of G. bilamina resemble tuberosity shallow and transversely narrow. those of other macropodoids with tall, slightly Entocuneiform facet small and oval; caudally directed neural spines and outwardly ectocuneiformfacet large,bilobed; mesocuneiform slanted prezygapophyses. This suggests contact absent. Low, broad anterior lateral crest restricted dorsoventral movement in the thoracic bears cuboid facet. Ectocuneiform (Fig. 6C, D) region, and saltating locomotion, as rigidity in the AAPMemoir25 (2001) head and thoracic region stabilizes the body (and acromion of G. bilamina and macropodids (with raises the centre of gravity) during hopping the exception of Hadronomas puckridgi; Murray (Christian & Preuchoft 1996). 1995) therefore may be related to a reduced Lumbar vertebrae in G. bilamina are similar to locomotory role for the forelimb. those of other macropodoids in possessing ventrally deflected transverse processes and Humerus (Fig. 4A-C, Table I ). The humeral shaft steeply oriented prezygapophyses. Ventral in G. bilamina resembles that of H. moschatus deflection of the transverse processes implies and the propleopine oscillans (Ride enlargement of the m. erector spinae, involved in et al. 1997) in being relatively straight and slender, flexing the lumbar region during hopping (Elftman characteristics indicative of quadrupedal cursors 1929). Orientation of the zygapophyses also (Ride et al. 1997). The deltoid crest is narrow and suggestsconsiderable lumbar flexibility necessary extends far down the shaft as in T. vulpecula, for generating power for the initial upward and species of Dendrolagus and Dorcopsis and anterior propulsive thrust (Elftman 1929). sthenurines(Murray 1995;Wells & Tedford 1995). Crest form in G. bilamina, species of Dorcopsis Scapula (Fig. 2, 3, Table 1). The scapular blade in and sthenurines, however, differs in being G. bilamina resembles that of the phalangerid uniformly low along its length. Lengthening of Trichosurus vulpecula, the hypsiprymnodontid the deltoid crest corresponds to elongation of the Hypsiprymnodon moschatus and the potoroine m. deltoideus muscle (divided into anterior Potorous tridactylus in being subrectangular in abducting and posterior retracting segments outline. Ganguroo bilamina differs from these respectively) lever arm, thus increasing the taxa, however, in possessing a more acute muscle's range of action and strength.Enlargement coracovertebral angle (a feature ofmacropodids of the clavicle in G. bilamina (with its related to reduction of the attachment area for the correspondingly lengthened m. clavodeltoid m. supraspinatus, and thus, extensor power of attachment) similarly suggests extensive m. the forelimb, see below; Murray 1995). The deltoideus musculature. Reduction of deltoid crest supraspinous and infraspinous fossae in G . height is a feature of quadrupedal cursors (Ride bilamina are large and subequalin anteroposterior et al. 1997), however, its reduction in bipedal extent suggesting powerful m. supraspinatus (an sthenurines suggests the feature may also be extensor of the forelimb) and m. infraspinatus related to forelimb browsing. muscles (rotator of the forelimb on the scapula). The condition of the supinator crest in G. In macropodids the supraspinous fossa is much bilamina resembles that of H. moschatus, P. reduced anteroposteriorly and the infraspinous oscillans (Ride et al. 1997) and species of fossa becomes dorsoventrally more extensive Dorcopsis in being tall and transversely narrow. (this development is most extreme in derived This would reduce area of attachment for the m. sthenurines, e.g. occidentalis; supinator longus (= m. brachioradialis which Murray 1995). The condition in G. bilamina rotates the forearm and flexes the carpal digits), a therefore suggests powerful extensors and common condition in quadrupedal cursors (Ride rotators were present as in T. vulpecula, H. etal.1997). moschatus and P. tridactylus, in which they act The teres tuberosity in G. bilamina is reduced to extend and rotate the forelimb during as in T. vulpecula, H. moschatus and P. osci//ans locomotion. The condition in macropodids (Ride et al. 1997) suggesting reduction of the m. indicates reduced emphasis on extension and latissimus dorsi and m. teres major respectively. development of a more extensive limb-rotating Both these muscles act as humeral retractors and musculaturepossibly for manipulation of food and are minimally developed (as in G. bilamina) in grooming. cursorial (Ride et al. 1997). The acromion process in G. bilamina is large The distal extremity of the humerus in G . and lobate similar to that of macropodids. The bilamina is similar to that of potoroines and acromion process anchors the spino~ macropodines ( except species of Dendrolagus ) acromiodeltoid portion of the m. deltoideus which in reduction of the medial entepicondyle. This serves to abduct the humerus. In forms limits the attachment area for the digital flexors consistently using quadrupedal gaits such as T. and indicates reduced strength in the digits of the vulpecula, H. moschatus and P. tridactylus the hand. In climbing such as T. vulpecula processis massive and steeply inclined indicating and species of Dendrolagus, the entepicondyle considerable humeral abduction. The reduced is strongly projecting and bears a prominent 132 AAP Memoir 25 (200 1) anterior groove (accommodating the flexor marsupials such as Sarcophilus harrisii and tendon). Absence of these features in G. bilamina Thylacinus cynocephalus in being transversely suggests insufficient gripping strength (in the compressed and spatulate in lateral view. This hand), to support the body weight, and thus increases the attachment area for the m. sartorius probably largely terrestrial habits. which acts to extend the upper portion of the The capitellum and trochlea in G. bilamina are hindlimb. unequal in size, a feature also occurring in T. The pectineal process is well developed and vulpecula, H. moschatus and P. tridactylus .This subcylindrical in G. bilamina, resembling that of may be an adaptation towards increasing macropodids ( except species of Dorcopsis and rotational flexibility of the elbow and/or Dendrolagus, tumbuna; Menzies & concentrating body weight through the more Ballard 1994, and H. puckridgi; Murray 1995). robust capitellar facet (on the ulna) during An extended pectineal process increases the area quadrupedal progression. for attachment of the m. psoas minor and m. pectineus (which act to flex the spine and hip and Ulna (Fig. 4D-F, Table I). TheulnainG. bilamina extend the pelvis), a common feature of resembles that of T. vulpecula, H. moschatus, macropodoid species which consistently use potoroines and sthenurines (Wells & Tedford saltating gaits (pers. obs.). 1995) in bearing a dorsoventrally deep olecranon The area of m. rectus femoris scarring in G . process and pronounced transverse dorsal crest. bilamina is anteroposteriorly extensive, similar This suggestsenlargement of the m. triceps brachii to that of potoroids and macropodids in which m. and m. anconeus muscles (which extend and rectus femoris abducts the femur retract the ulna during elbow flexion) and anteroposteriorly. Scarring is extensive in all strengthening of the forelimb for locomotion (e.g. macropodoids but is anteroposteriorly reduced in species of Dorcopsis and Dendrolagus, H. T. vulpecula, H. moschatus)and/or browsing (e.g. sthenurines; Wells & Tedford 1995). puckridgi (Murray 1995) and Protemnodon hopei (pers. obs., suggesting that reduction of the rectus The m. flexor digitorum profundus attachment muscle may be related to the reduced abductive in G. bilamina is reduced in its depth and width stressesofmore quadrupedalmodes in thesetaxa). similar to that of many macropodines.The m. flexor digitorum profundus serves to flex the wrist and Tibia and fibula (Fig. SD-E, Table I ). The metacarpals,therefore its reduction may be related possibility of limited movement between the tibia to stiffening of the wrist and limitation of the manus and fibula in G. bilamina is suggestedby: the large to a more locomotory role. posterior processof the proximal fibula epiphysis; The m. biceps brachii attachment is well prominent fibular sulcus on the lateral femoral developed in G. bilamina and, as in all condyle; and width of the tibial facet on the macropodoids (Dawson et al. 1989), would have proximal epiphysis of the fibula. These features formed a powerful flexor of the elbow. are also found in species of Dendrolagus (in which the tibia and fibula move freely relative to Pelvis (Fig. 5A-B, Table I). The ilium in G. each other, Flannery & Szalay 1982; Flannery et bilamina is similar that of other macropodoids in al. 1996; Bishop 1997) in which they facilitate being laterally flared relative to the long axis of lateral mobility in the lower hindlimb for the vertebral column, and having the sacro-iliac quadrupedal locomotion and climbing. contact situated far back on the iliac blade. This suggestsenlargement of the erector spinae which Pes (Fig. 6, Table I). The elongate, condyle-like support the upper half of the body during hopping navicular facet of G. bilamina resembles that of (Elftman 1929) and may also have acted to raise H. moschatus and P. tridactylus in which it the torso during browsing. The m. gluteus medius, provides flexibility to the navicular/cuneiform m. gluteus profundus and m. iliacus (which act to complex and hence the medial digits of the foot extend/evert the hip and abduct/evert the thigh) (digits I-III). This condition is unlike that in the attachments are large, representing the common majority of potoroines and macropodids in which condition in macropodoids. Enlargement of these the facet is shorter and more laterally oriented. muscle groups is indicative of saltating gaits This arrangement reduces lateral flexibility in the (Elftman 1929;Alexander & Vernon 1975). pes, stiffening it for more effective saltation. The The anterior ilial apex in G. bilamina resembles loss of a mesocuneiform/navicular contact that of T. vulpecula, H. moschatus, species of suggests the pes in G. bilamina was, as in all Dorcopsis and Dendrolagus and ambulating macropodoids, narrowed to some extent. AAP Memoir 25 (2001) 133

Taxon 1 I 2 I 3 I 4 9 10 I II I 12 I 13 I 14 I 15 I 16 I 17 I B. parVus ~I~I~ o 0 , ° I 0 u~~J_o 10 o I 0 I 7: vuloecula O I O I O 0 o I 0 I 0 0 o ) Qj --5!J~ ~I- o I IH. moschatus I o I 0 I 0 1 O . 0 o I o I o I o 0 (\ InI U

P. tridactylILY O o I o 01 0 Qj~_Q_I o o I G. bilamina 0 1 , o I 01 o J ~ 0 D. alrata 1 1 ~ rl o I 0 D.goodfellowi 1 H ~ :1 P. penicillata 1 o I ~ 1 I T. thetis 1 O ~ w: bicolor I H 1 o I ~ M parryi I 1 i ~ ~ S. stirlingi I o~~ ~1 H Table 2. Character state distribution for phylogenetic analysis of Ganguroo bilamina relationships. Plesiomorphic state = 0; apomorphic state = I. longer or subequal to ventral edge (I ). CHARACTER ANALYSIS parvus, T. vulpecula, H. moschatus, P. tridactylus Seventeenpostcranial characterswere selected and G. bilamina all exhibit a crescentic navicular forphylogenetic analysis. Character selection was in lateral view with the ventral edge longer than limited by specific element type preserved in the dorsal (0). A sub rectangular navicular is G. bilamina specimen. Because the specimen considered derived relative to a crescentic fonn representsa juvenile, comparison was made with and represents a synapomorphy for S. stirlingi juvenile specimens of parryi, and macropodines. Aepyprymnus rufescens and Potorous tridactylus 2. Fonn of navicularplantar eminence.Eminence to screen for ontogenetic variability within fonn in S. stirlingi and macropodines is reduced characters.Character polarity was established by (I) unlike that of B. parvus, T. vulpecula, H. comparison with outgroup taxa. Determination of moschatus, P. tridactylus and G. bilamina which the plesiomorphic statefollows Hennig ( 1966) and is condyle-like and prominent (0). A reduced Wiley et al. ( 1991) with referenceto the immediate eminence is considered derived relative to sister taxon to the ingroup (represented herein by prominent and condyle-like and is synapomorphic the phalangeriform Trichosurus vulpecula; Aplin for s. stirlingi and macropodines. & Archer 1987; Flannery 1987). The burramyid 3. Length and width of astragalar facet of Burramys parvus was included to aid in resolving navicular. Astragalar facet in G. bilamina, S. character polarities within the ingroup (see stirlingi and macropodines is transversely wide Stevens1980; Simmons 1993). and dorsoventrally shortened ( 1) relative to facet Character states were determined by fonn inB. parvus, T. vulpecula, H. moschatusand examination of representative specimens in the P. tridactylus which is transversely narrow and collections of the Australian Museum and dorsoventrally elongate (0). A wide short facet is National Museum ofVictoria. No single complete consideredderived relative to the narrow, elongate specimen of was available for study. fonn and represents a synapomorphy uniting G . Character state determination for this taxon was bilamina, S. stirlingi and macropodines. therefore made by reference to published 4. Mesocuneifonn/navicular contact. The descriptions and figures of Wells & Tedford mesocuneifonn does not contact the navicular (1 ) (1995). in macropodoids ( evidenced in G. bilamina by The following is a list of character definitions the absence of a mesocuneifonn facet on the and interpretations. Character distributions are navicular) unlikeB.parvus and T. vulpecula which listed in Table 2. exhibit a clear mesocuneifonn-navicular contact 1. Proportions of navicular. The navicular in S. (0). Absence of contact between the stirlingi and macropodines (represented herein mesocuneiform and navicular is considered by Dendrolagus goodfellowi, Dorcopsis atrata, derived relative to contact presence and is a Petrogale penicillata, Thylogale thetis, macropodoid synapomorphy. Wallabia bicolor and Macropus parryi) is 5. Form of acromion process of scapula. The subrectangular in lateral view having dorsal edge acromion process of the scapula in G. bilamina,

~ 134 AAP Memoir 25 (200 1)

S. stirlingi and macropodines is poorly developed macropodines; ridge is weakly developed and and rounded (I) unlike B. parvus, T. vulpecula, poorly evident (0) in B. parvus, T. vulpecula, H. H. moschatus and P. tridactylus in which the moschatus and G. bilamina. A well developed acromion is well developed and anterodorsally ridge is considered derived relative to a poorly produced (0). A poorly developed and rounded developed ridge and represents a synapomorphy acromion process is considered derived relative for S. stirlingi and macropodines. The condition to one which is well developed and anterodorsally in P. tridactylus may be autapomorphic or produced. Condition ( I) represents a represent a synapomorphy uniting P. tridactylus, synapomorphy for G. bilamina, S. stirlingi and s. stirlingi and macropodines which has been macropodines. secondarily lost in G. bilamina. 6. Condition of supraspinous fossa of scapula. 10. Condition of entepicondyle of humerus. The supraspinous fossa in S. stirlingi and The entepicondyle closely abuts the trochlea ( I ) macropodines is anteroposteriorly shortened inH. moschatus, P. tridactylus, G. bilamina and relative to the infraspinous fossa ( I ); D. atrata unlike B. parvus, T. vulpecula, s. supraspinous fossa is anteroposteriorly elongate stirlingi and macropodines in which the and subequal in length to the infraspinous fossa entepicondyle is widely separated from the (0) in B. parvus, T. vulpecula, H. moschatus, P. trochlea by a distinct groove (0). A closely tridactylus and G. bilamina. An anteroposteriorly abutting entepicondyle and trochlea is considered shortened supraspinous fossa is considered derived relative to their being widely separated derived relative to an elongate fossa and by a distinct groove. Condition (I) may represent represents a synapomorphy for S. stirlingi and a synapomorphy for macropodoids which has macropodines. been secondarily lost in S. stirlingi and 7. Condition of supinator crest on humerus. macropodines (except D. atrata). Loss of the The supinator crest in G. bilamina and D. atrata feature may thus represent a synapomorphy for is reducedand transversely narrow with a rounded this latter group. The absenceof the derived state apex (I). The crest is well developed and in D. atrata may be a further secondary reversal transversely broad with distinctly pointed apex or, more parsimoniously, a retention of the state (0) in B. parvus, T. vulpecula and all other present in plesiomorphic macropodoids. macropodoids. A reduced and narrow crest with Phylogenetic significance of this character is rounded apex is considered derived relative to a equivocal because placement of the shared well developed and broad crest with distinctly derived state can not be resolved. pointed apex. The latter condition appears to be II. Development of coronal depression on independently derived in G. bilamina and D. humerus. The coronal depression is deeply atrata. However, the condition may also represent excavated and strongly delineated (I) in G. a plesiomorphic state for macropodids which has bilamina, T. the/is, w. bicolorandM.panyiunlike been subsequently lost in all other taxa. The B. parvus, T. vulpecula, H. moschatus, P. character is uninformative because it represents tridactylus, s. s/irlingi, D. atrata, D. goodfellowi an autapomorphic state in this phylogeny. and P. penicillata, all of which exhibit a shallowly 8. Height of deltoid crest of humerus. The excavated and weakly delineated (0) coronal deltoid crest is of uniform height along its length depression. A deeply excavated and strongly with the distal extremity forming a shallow incline delineated depression is considered derived (I) inH. moschatus, G. bilamina andS. stirlingi. relative to a shallowly excavated and weakly The crest is of non-uniform height with its distal delineated depression. Condition ( I) is a extremity forming a steepincline which may bear synapomorphy for the clade containing T. thetis, a rugose boss at its apex (0) in B. parvus, T. w. bicolor and M. parryi and is autapomorphic in vulpecula, P. tridactylus and macropodines. A G. bilamina. uniform crest with shallowly inclined distal 12. Condition of capitellum and trochlea of extremity is considered derived relative to a non- humerus. The capitellum and trochlea are uniform crestwith steeplyinclined distal extremity. subequal in size (I) in S. stirlingi and Condition (I) is autapomorphic inH. moschatus, macropodines; but are of distinctly unequal size G. bilamina and S. stirlingi and is thus with the capitellum larger than the trochlea (0) in uninformative. B. parvus, T. vulpecula, H. moschatus, P. 9. Condition oflateral deltoid ridge ofhumerus. tridactylus and G. bilamina. Similarity in size of The lateral deltoid ridge is well developed and the capitellum and trochlea is considered derived prominent (I) in P. tridactylus, S. stirlingi and relative to dissimilarity in size, and is a synapomorphy for s. stirlingi and macropodines. AAPMemoir 25 (2001)

13. Condition of ulna shaft. The shaft of the prominent subcylindrical iliopectineal process is ulna inS. stirlingi and macropodines (except D. considered derived relative to a reduced process goodfellowi) has a distinctly sinuous ventral edge and is a synapomorphy for P. tridactylus, G. profile in lateral view (1) unlike B. parvus, T. bilamina, S. stirlingi and macropodines (except vulpecula, H. moschatus, P. tridactylus, G. D. goodfellowi). The condition in D. goodfellowi bilamina and D. goodfellowi in which the ulna represents an autapomorphic secondary reversal shaft has a straight ventral edge profile in lateral to the plesiomorphic state. view. A sinuous ventral edge profile is considered 17.Ventral profile of anterior lumbar centra.The derived relative to a straight ventral edge profile ventral profile bears a distinct median keel ( 1) in and is synapomorphic for S. stirlingi and G. bilamina and macropodines and is unlike B. macropodines (except D. goodfellowi). The parvus, T. vulpecula,H. moschatus,P. tridactylus condition in D. goodfellowi may represent an and s. stirlingi in which ventral profile is smooth autapomorphic secondary reversal or a retained and rounded (0). A distinct ventral keel is plesiomorphy. considered derived relative to a smooth and 14. Form of olecranon process of ulna. The rounded surface and is a synapomorphy for G . olecranon process in S. stirlingi and bilamina and macropodines. The condition in S. macropodines (except D. goodfellowi) is stirlingi is an autapomorphic secondary reversal dorsoventrally short (when element is horizontally to the plesiomorphic state. oriented), and rounded in lateral view (1). Those of B. parvus, T. vulpecula, H. moschatus, P. ~TS tridactylus, G. bilamina and D. goodfellowi are The analysis produced two equally dorsoventrally tall (when element is horizontally parsimonious trees (Fig. 7A, B. ), both of27 steps oriented) with a distinct, transverse dorsal crest with CI of 0.63 (expected CI = 0.667), RI of 0.818 (0). A dorsoventrally short and rounded olecranon and RCI of 0.515. Tree topology is identical for process is considered derived relative to a tall, heuristic and branch-and-bound search options. distinctly crested process. The derived condition Below expected consistency index values are represents a synapomorphy for S. stirlingi and produced by a polytomy including S. stirlingi and macropodineswith the condition inD. goodfellowi macropodines (except D. goodfellowi and D. being a retained plesiomorphy or a secondary atrata in tree 2; Fig. 7B.). Comparison with 'next- reversal to the plesiomorphic state. best' trees indicates poly to my resolution 15. Shapeof distal apex ofilium. The distal apex decreases with additional steps making of the ilium in P. tridactylus, S. stirlingi and interpretation of relationships within the S. macropodines is transversely broadened by stirlingi/macropodine group tentative. contribution of the raised distal end of the iliac crest (1); the apex is transversely narrowed by a Discussion poor contribution of the distal end of the iliac Macropodoid monophyly is unequivocally crest (which is reducedin height, making the distal supported in this phylogeny by an apomorphic ilial apex spatulatein lateral view; 0) inB. parvus, absence of contact between the mesocuneiform T. vulpecula, H. moschatus and G. bilamina. A and navicular. Potoroids (sensu Flannery 1989) transversely broad, distal ilial apex is considered do not form a discrete clade, with H. moschatus derived relative to one which is transversely placed as the sister taxon to all other narrow and is a synapomorphy for S. stirlingi macropodoids and P. tridactylus the sister taxon and macropodines.The condition in P. tridactylus to G. bilamina, S. stirlingi and macropodines may be autapomorphic or represent a (unequivocally united by the presence of a synapomorphy shared with S. stirlingi and prominent, subcylindrical iliopectineal process). macropodines which has been secondarily lost in Ganguroo bilamina is placed with S. stirlingi and G. bilamina. macropodines on the basis of: a transversely wide 16. Development of the ilio-pectineal process and dorsoventrally shortened astragalar facet on of ilium. The iliopectineal processinP. tridactylus, the navicular; a rounded and weakly developed G. bilamina, S. stirlingi and macropodines( except acromion process on the scapula; and presence D. goodfellowi) is prominent and subcylindrical of a distinct ventral keel on the anterior lumbar (1; more transversely compressed in S. stirlingi). vertebrae (not present in S. stirlingi). The process in B. parvus, T. vulpecula, H. Unequivocal apomorphies uniting S. stirlingi moschatus and D. goodfellowi is reduced and if and macropodines include: navicular rectangular present, restricted to a slight ridge (0). A in lateral view with dorsal edge longer than or 36 AAPMemoir25 (2001)

A B

B. parvus B. parvus

7: vulpecula T. vulpecula

H. moschatus H. moschatus

P. tridactylus P. tridactylus

G.bilamina G. bilamina D.goodfel/owi D.goodfel/owi D. atrata D. atrata ~ penicil/ata ~ penicillata S. stirlingi S.stirlingi 7:thetis 7:thetis W bicolor W bicolor M.parryi M.panyi

Fig. 7. Results of phylogenetic analyses. A, tree I topology includes D. atrata within the S. ster/ingi/macropodine polytomy. B, tree 2 topology includes D. atrata as sister taxon to the S. ster/ingi/macropodine polytomy. For both trees, length = 27, CI = 0.63, RI = 0.818, RCI = 0.515.

subequal to ventral edge; reduction of the supports the conclusions of Case (1984), navicular plantar eminence; anteroposteriorly Woodburne (1984) and Cooke (1997a, b) who shortened supraspinous fossa; and subequally regard bulungamayines as basal to Macropodidae. sized capitellum and trochlea on the humerus. This arrangement also supports Ride ( 1993), Dendrolagus goodfellowi forms the sister Cooke ( 1997a) and Wroe et al. ( 1998) in regarding taxon to a polytomy including s. stirlingi and the (sensuFlannery 1989) as polyphyletic. remaining macropodines, united by: a sinuous The possibilities of a sister group relationship of ventral edge to the ulnar shaft; and dorsoventrally potoroines to both bulungamayines and short and rounded olecranon process. Separation macropodids, or a polyphyletic Bulungamayinae, of D. atrata from the S. stirlingi/macropodine however, are plausible. Separationof H. moschatus polytomy in tree 2 (Fig. 7B.) is based on an from potoroids (Ride 1993; Szalay 1994; Cooke equivocal character state (a closely abutting 1997a;Wroeetal.1998;Burketal.1998) may also entepicondyle and trochlea on the humerus) and be suggested by the absence of any clear cannot be substantiated in this phylogeny. The synapomorphy. terminal clade formed by T. thetis, W. bicolor and M. parryi is united by an apomorphic deeply CONCLUSIONS excavated and strongly delineated coronal 1. Ganguroo bilamina was most likely partly depression on the humerus. This contradicts the quadrupedal, although bipedal saltation was immunological and DNA hybridisation data of probably used at higher speeds (as in species of Baverstocketal. (1989),Baverstocketal. (1990) Dendrolagus; Windsor & Dagg 1971 and and Kirsch et al. (1995) respectively, who include Potorous; Buchmann & Guiler 1974). species of Thylogale as the sister group to those 2. Bulungamayinae, represented herein by G. of Petrogale. bilamina, is the sister group to macropodids. This The inclusion of S. stirlingi within the supports the conclusions of Case (1984), macropodine complex may support the Woodburne(1984)andCooke(1997a,1997b). conclusions ofWoodburne ( 1967),Archer ( 1984), Szalay (1994) and Cooke (1997a) who suggest ACKNOWLEDGEMENIS origin of from within a paraphyletic We are indepted to B. Cooke and T. Myers for . The lack of resolution in this providing much neededcritical discussion. Many phylogeny, however, makes any conclusion thanks to T. Myers for reading earlier drafts of the extremely tentative. work. Thanks to S. Ingleby and the Australian Recognition of the bulungamayine G. bilamina Museum, H. Godthelp and the University of New as the sister taxon to s. stirlingi and macropodines South Wales for generousprovision of specimens.

~ AAP Memoir 25 (2001) 137

All photographs prepared by C. Bento of the BLACK, K., 1997. Diversity and biostratigraphy of Australian Museum. Support for this researchwas the Diprotodontoidea ofRiversleigh, northwestern provided by: the Australian Research Grant Queensland. Memoirs of the Queensland Museum Scheme (grants to M. Archer); National Estate 41, 187-192. Grants Scheme (grants to A. Bartholomai and M. BUCHMANN, O.L.K. & GUILER, E.R., 1974. Archer); Commonwealth Department of Locomotion in the . Journal ofMammalogy Environment, Sports and Territories; Queensland 55,203-206. National Parks and Wildlife Service; BURK, A., WESTERMAN, M. & SPRINGER, M., Commonwealth World Heritage Unit; University 1998. The phylogenetic position of the Musky Rat- of New South Wales; Queensland Museum; kangaroo and the evolution of bipedal hopping in Australian Museum; IBM Australia Pty Ltd; ICI (Macropodidae: ). Australia Pty Ltd; Australian Geographical Systematic Biology 47,457-474. Society; Wang Australia Pty Ltd; Century Zinc CASE, J.A., 1984. A new genus of Potoroinae Pty Ltd; Mount Isa Mines Pty Ltd; Surrey Beatty (Marsupialia: Macropodidae) from the Miocene & Sons Pty Ltd; Riversleigh Society Inc.; Royal Ngapakaldi Local Fauna, South Australia, and a Zoological Society of New South Wales; Linnean definition of the Potoroinae. Journal ofPaleontology Society of New South Wales; and private 58,1074-1086. supporters. CHRISTIAN, A. & PREUCHOFT, H., 1996. Deducing the body posture in extinct large vertebrates from REIiERENCFS the shape of the vertebral column. Palaeontology ALEXANDER, R.M. & VERNON, A., 1975. The 39,801-812. mechanics of hopping in kangaroos COOKE,B.N., 1997a.Researchesintofossil kangaroos (Macropodidae ). Journal of Zoology, London 177, and kangaroo evolution. Unpublished doctoral 265-303. thesis, University of New South Wales, Sydney. APL1N, K. P. & ARCHER, M., 1987. Recent advances COOKE, B.N., 1997b. New Miocene bulungamayine in systematics with a new syncretic kangaroos (Marsupialia: Potoroidae) from classification. xv-lxxii inArcher, M. (ed.), Possums Riversleigh, northwestern Queensland. Memoirs of and opossums:studies in evolution, 1, Surrey Beatty the Queensland Museum 41,281-294. & Sons, Sydney. COOKE, B.N., 1997c. Biostratigraphic implications ARCHER, M. HAND, S.I., GODTHELP, H. & of fossil kangaroos at Riversleigh, northwestern CREASER, P., 1997. Correlation of the Cainozoic Queensland. Memoirs of the Queensland Museum sediments of the Riversleigh World Heritage fossil 41,295-302. property, Queensland,Australia. 131-152 inAguilar, COOKE, B.N. & KEAR, B.K., 1999. Evolution and I.P., Legendre, S. & Michaux, I. (eds), Actes du diversity of macropodoids. Australian Mammalogy Congres BiochroM'97. Ecole Pratique des Hautes 21,27-29. Estudes Institut de Montpellier, Montpellier. DAWSON, T.J., FINCH, E., FREEDMAN, L., HUME, ARCHER, M. et al., 1994. List of the principal I.D., RENFREE, M.B. & TEMPLE-SMITH, P.D., Riversleigh local faunas and their interpreted relative 1989. Morphology and physiology of the ages. Unpublished Abstracts of the Riversleigh Metatheria. 451-504 in Walton, D. W. & Richardson, Symposium 1994, Supplement, 28-31. B.J. (eds) Fauna of Australia. Vol. 1B Mammalia. ARCHER, M., 1984. The Australasian marsupial Australian Government Publishing Service, radiation. 633-808 in Archer, M. & Clayton, G. Canberra. (eds), Vertebrate zoogeography and evolution in ELFTMAN, H.O., 1929. Functional adaptations of Australasia. Hesperian Press, Perth. the pelvis in marsupials. Bulletin of the American BAVERSTOCK, P.R., RICHARDSON, B.I., Museum of Natural History 58, 189-232. BIRRELL, I. & KRIEG, M., 1989. Albumin FINCH, M.E. & FREEDMAN, L., 1986. Functional immunologic relationships of the Macropodidae morphology of the vertebral column of Thylacoleo (Marsupialia). Systematic Zoology 37,38-50. carnifex Owen (Thylacoleonidae: Marsupialia). BAVERSTOCK, P.R., KRIEG, M. & BIRRELL, I., Australian Journal ofZoology 34,1-16. 1990. Evolutionary relationships of Australian FLANNERY, T.F., 1987. The relationships of the marsupials as assessedby albumin immunology. macropodoids (Marsupialia) and the polarity of Australian Journal ofZoology 37, 273-287. some morphological features within the BISHOP, N., 1997. Functional anatomy of the Phalangeriformes. 741-747 in Archer, M. (ed.), macropodoid pes. Proceedings of the Linnean Possums and opossums:studies in evolution. Surrey Society of New South Wales 117,17-50. Beatty & Sons, Sydney. 38 AAP Memoir 25 (2001'

FLA'NNERY, T.F., 1989. Phylogeny of the Woodburne, 1967 (Marsupialia: Macropodidiae). Macropodoidea; a study in convergence. 1-46 in Alcheringa 19, 119-170. Grigg, G., Jarman, P. & Hume, I. (eds), Kangaroos. RIDE, W.D.L., 1993. Jackmahoneya gen. nov. and the wallabies and rat-kangaroos. Surrey Beatty & Sons, genesis of the macropodifonn molar. Memoirs of Sydney. the Association of Australasian Palaeontologists 15, FLANNERY, T.F., ARCHER, M. & PLANE, M., 441-459. 1983. Middle Miocene kangaroos (Macropodoidea: RIDE, W.D.L., PRIDMORE, P.A., BARWICK, R.E., Marsupialia) from three localities in northern WELLS, R.T. & HEADY, R.D., 1997. Towards a Australia, with a description of two new subfamilies. biology of Propleopus oscillans (Marsupialia: BMRJournal of Australian Geology & Geophysics Propleopinae, ). Proceedings 7,287-302. of the LinneanSociety of New South Wales117,243- FLANNERY, T.F., MARTIN, R. & SZALAY, A., 1996. 328. Tree kangaroos: a curious natural history. Reed SIMMONS, N.B., 1993. The importance ofmethods: Books, Melbourne. 202p. archontan phylogeny and cladistic analysis of FLANNERY, T.F.&SZALAY;F., 1982.Bohrapaulae, morphological data. 1-61 in MacPhee,R.D.E. (ed.), a new giant fossil tree-kangaroo (Marsupialia: Primates and Their Relatives in Phylogenetic Macropodidae) from New South Wales, Australia. Perspective. Plenum Press, New York. Australian Mammalogy 5, 83-94. STEVENS, P.F., 1980. Evolutionary polarity of HENNIG, W., 1966. Phylogenetic Systematics. character states. Annual Review of Ecology and University of Illinois Press, Urbana. 263p. Systematics 11, 333-358. HOPWOOD, P.R., 1974. The intrinsic musculature of SZALAY, F.S., 1994. Evolutionary history of the the pectoral limb of the Eastern Grey Kangaroo. marsupials and an analysis of osteological Journal of Anatomy 118, 445-468. characters. Cambridge University Press,New York. HOPWOOD, P.R. & BUTTERFIELD, R.M., 1976. 455p. The musculature of the proximal pelvic limb of the WILEY, E.O., SIEGEL-CAUSEY, D., BROOKS, D.R. Eastern Grey Kangaroo. Journal of Anatomy 121, & FUNK, V.A., 1991. The compleat cladist: a primer 259-277. of phylogenetic procedures. University of Kansas HOPWOOD, P.R. & BUTTERFIELD, R.M., 1990. Natural History Museum Special Publication 19, 1- The locomotor apparatus of the crus and pes of the 158 Eastern Grey Kangaroo, Macropus giganteus. WINDSOR, D.E. & DAGG, A.I., 1971. Gaits in the Australian Journal ofZoology 38,397-413. Macropodinae (Marsupialia). Journal of Zoology, JOHNSON, P.M. & STRAHAN, R., 1982. A further London 163, 165-175. description of the Musky Rat Kangaroo, WELLS, R.T. & TEDFORD, R.H., 1995. Sthenurus Hypsiprymnodon moschatus Ram say, 1876 (Macropodidae: Marsupialia) from the (Marsupialia: Potoroidae), with notes on its biology. of Lake Callabonna, South Australia. Bulletin of the Australian Zoology 21,27-46. American Museum of Natural History 225, 1-111. KEAR, B.P., 1998. Postcranial morphology and WOODBURNE, M.O., 1967. The Alcoota Fauna, phylogenetics of Oligo-Miocene kangaroos central Australia: an integrated palaeontological and (Marsupialia: Macropodoidea) from Riversleigh. geological study. Bulletin of the Bureau of Mineral northwestern Queensland. Unpublished Honours Resources, Geology and Geophysics 87, 1-187. thesis, University of New South Wales. WOODBURNE, M.O., 1984. Wakiewakie lawsoni, a KIRSCH, J.A.W., LAPOINTE, F.J. & FOESTE, A., new genus and species ofPotoroinae (Marsupialia: 1995. Resolution of portions of the kangaroo Macropodidae) of medial Miocene age, South phylogeny (Marsupialia: Macropodidae) using Australia. Journal ofPaleontology 58, 1062-1073. DNA hybridization. Biological Journal of the WROE, S., BRAMMALL, J. & COOKE, B.N., 1998. Linnean Society 55,309-328. The skull of ima (Marsupialia: MENZIES, J.I. & BALLARD, C., 1994. Some new Hypsiprymnodontidae?): an analysis of some cranial records of Pleistocene megafauna from New features within Marsupialia and re-investigation of Guinea. Science in New Guinea 20, 113-139. propleopine phylogeny; with notes on the inference MURRAY, P.F., 1995. The postcranial skeleton of the of carnivory among mammals. Journal of Miocene kangaroo, Hadronomas puckridgi Paleontology 72,738-751.