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Kangaroo-related motor vehicle collisions are common but there is limited literature on this topic. Drivers confronted by kangaroos may choose to swerve or to directly collide with the kangaroo. The effect of these differing crash mechanisms, along with the effect of vehicle type or time of day, has not yet been reported.A retrospective cohort study was performed, examining patients admitted to our tertiary trauma centre for kangaroo-related motor vehicle collisions between 2000 and 2020. Data on patient demographics, crash characteristics, and hospital stay were collected and analysed.A total of 366 patients were included and were predominantly male (76%) with an median age of 40. Swerve crashes were more common (59%) than direct impact and swerving was a statistically significant predictor of reduced injury severity score on multivariable analysis (other significant factors were female sex and no rollover). Motor vehicle crashes and motorbike crashes had differing crash characteristics. Motor vehicle crashes were more likely the result of swerving, and swerving was less likely to cause ejection or require extrication but more likely to cause rollover. Motorbike crashes however, were more likely the result of head on collision and riders were more likely to be ejected from the vehicle, require extrication, or be involved in a rollover. In terms of time of day, there were more crashes at dawn and there was a trend towards higher injury severity score and length of stay for night-time crashes.For kangaroo-related motor vehicle crashes, predictors of increased injury severity score on multivariable analysis were male sex, direct impact, and rollover. Motorbikes and motor vehicles had differing crash mechanisms and characteristics, as did night-time crashes when compared to daytime or twilight crashes.IV, prognostic.

Bohra planei sp. nov. http://zoobank.org/ urn:lsid:zoobank.org:act: 47F6EBDE-2594-4710-B440-D1D940951248 cf. Dorcopsis, large form: Plane (1967): pp. 50–55, figures 11–13. Hoch & Holm (1986), p. 188. cf. Dendrolagus sp.: Woodburne (1967), p. 82. Dorcopsis large species: Flannery et al. (1983b), p. 77. cf. Dorcopsis sp.: Flannery et al. (1989), pp. 151–152. “ Dorcopsis ”: Menzies & Ballard (1994), p. 131. Silvaroo sp. indet.: Dawson (2004b), p. 284, table 1. Silvaroo sp. indet. 2: Dawson (2004b), pp. 286–287. ? Dorcopsis sp.: Dawson (2004b), p. 288. Dorcopsis sp. nov.: Black et al. (2012), p. 1050. Holotype. UCMP 70128, partial right adult hindlimb, including femur, distal half of tibia, distal end of fibula, talus and calcaneus. The fibula fragment could not be located in the UCMP collection in August 2007. Type locality. Watut 3 (UCMP V6356), located on the eastern bank of the Upper Watut River, downstream from Awe Fauna Type Locality (= Watut 1, V6234), Morobe Province, Papua New Guinea. Judging from the location map (figure 15 in Plane 1967), V6356 lies perhaps 500 m northeast of Awe Fauna Type Locality. UCMP 70128 was collected by Michael D. Plane in 1963 and is the only specimen recorded from Watut 3. The specimen was collected from a blue-grey claystone stratigraphically lower than Awe Fauna Type Locality. Referred specimens. Charlie Lawrence Locality (V 6172), Wiganda Creek, Upper Watut Valley, Morobe Province, Papua New Guinea. AM F49468, right adult maxilla (preserving M1–4). This specimen was collected in 1961 and presented to the Australian Museum by T. Brown. Before becoming submerged by a pond, the site was located “on the lower stretch of Wiganda Creek where the Gold and Power Company were working alluvial gold” (Plane 1967: 63). No stratigraphic details were recorded. Widubosh goldmine, Upper Watut Valley. NHMD 193283 (formerly GMK 55709), partial right calcaneus. This specimen was collected by Ella Hoch on 29 August 1983. Etymology. This species is named in honour of Michael D. Plane (1933–2017), who collected the holotype and first described the Awe LF and Otibanda Formation. Diagnosis. Bohra planei most closely resembles B. nullarbora and B. bila in upper molar size and morphology. Upper molars differ from those of B. nullarbora by having a cusp C region of the stylar crest on M1–3, incurved postpara- and premeta-cristae, and a broader, more open posterior metaloph face on M4. Upper molars differ from those of B. bila by having the bifurcation between the postparacrista and stylar cusp C region further from the paracone apex and a lingually thinner precingulum. Femur is transversely broad and craniocaudally compressed proximally with very broad greater trochanter, broad, shallow trochlea and rugose adductor scars running along the length of the caudomedial aspect of the shaft. Distal epiphysis and talar articular facet of the tibia are relatively narrow in comparison to other Bohra and Dendrolagus. Calcaneus is transversely broad and dorsoventrally flattened, with distinct plantar epiphyseal ridge and medial crest. Medial and lateral talar facets are mesially constricted and more confluent with each other than in all other species of Bohra. The distal extremity is narrow relative to the talar and fibular facets, and the calcaneocuboid facets relatively deep and narrow and with a more acutely stepped articulation than in other species of Bohra. Talus transversely broad, dorsoventrally flattened, with broad, shallow trochlea, medially extended malleolus, broad and shallow malleolar fossa, short talar neck and smoothly contiguous calcaneal facets. Description and comparisons. Maxilla and palatine. Very little is preserved of the maxilla of AM F49468, but there is sufficient to indicate that the base of the masseteric process was positioned above the anterior root of M 3 in lateral view (Figure 30A, C). The shape of the fracture through the palate that represents the mesial limit of this specimen is consistent with a line of breakage along the maxilla–palatine suture (Figure 30C). This is orientated anteromesially from adjacent to the M4 protoloph. What remains of the palatal portion of the maxilla is broadest adjacent to the anterior root of M3. The position of the masseteric process relative to M 3 in AM F49468 matches that observed in adult specimens of B. nullarbora, B. bila and B. bandharr. The anteromesial orientation of the inferred maxilla–palatine suture is a close match for that of B. nullarbora (Figure 27E), including the point at which it inflects anterolingually, as also is the relative width of the maxilla mesial to M3. By contrast, in B. bandharr, the maxilla–palatine suture runs anteroposteriorly for more of its length before inflecting anterolingually adjacent to M3 (Figure 14C), while it runs anteroposteriorly up to a position mesial to the M2 protoloph in B. bila (Figure 21B). Upper dentition. The single known maxillary specimen (AM F49468) preserves M1–4. The protoloph of M1 is marginally narrower than the metaloph, but from M2 to M4 the protoloph becomes increasingly wider than the metaloph (Figure 30C; Table 7). The molars are moderately worn, with dentine breached along the lengths of the lophs of M1–2, on the protoloph and metaconule of M3, and on the protocone of M4. Despite this, they were clearly low crowned, and it is evident from the less worn molars that the postparacrista, postprotocrista, premetacrista and postmetacrista are very fine and low (Figure 30). The postparacrista and premetacrista are distinctly incurved on all molars. On M1–3, the cusp C region of the stylar crest is evident, diverging from the postparacrista about halfway up the paracone face, such that it forms an inverted Y shape.The stylar crest remnant is projected buccally at 50° on M1 to the longitudinal axis of the crown, at 40° on M2 and at 30° on M3 (Figure 30C). The postmetaconulecrista forms a small but distinct shelf on the posterior face of M4. The upper molars of B. planei are smaller than in B. bandharr and B. illuminata, and closest in size and morphology to those of B. nullarbora and B. bila (Tables 1, 4–6). By comparison, B. nullarbora lacks any sign of a cusp C region of the stylar crest on any molars, and has anteroposteriorly orientated postpara- and premeta-cristae, a less lingually tapered precingulum, and a narrower, deeper, more ‘pocketed’ M4 posterior metaloph face. In B. bila, a cusp C region of the stylar crest is present on M1–2, but not M3, and differs by diverging from the postparacrista closer to the paracone apex. As in B. bila, the postpara- and premeta-cristae are incurved, but not as well developed as in B. wilkinsonorum. The configuration of these crests and the stylar crest on M1–3 is very like that of the species of Dorcopsulus, but in the much larger B. planei there is no obvious cusp D region of the stylar crest. It should be noted, however, that there is a small wear facet on the metacone anterior face buccal to the premetacrista on M1–2, which, in a minimally worn specimen, may reveal that this portion of the stylar crest was indeed retained in B. planei. The upper molars of B. planei share very weakly development of the anteroposterior crests with B. nullarbora and B. bandharr. Femur. Well preserved and largely complete, the holotype femur is missing only the lateral condyle of the distal epiphysis (Figure 31). The femoral head is large and roughly hemispheric. The proximal epiphyseal margin is more distally placed than the caudal margin of the head. The greater trochanter is very rugose at its apex, particular in cranial view, and strongly laterally flared at its base (Figure 31A). The lesser trochanter is robust and long, with a distally extended ridge and distinct sulcus on its craniodistal aspect (Figure 31C). The femoral shaft is slightly compressed in the craniocaudal plane, and is cranially convex. The caudal surface of the diaphysis is marked by a rugose adductor muscle scar, which runs obliquely from below the lesser trochanter to a depression marking the insertion for the m. quadratus femoris midway along the length of the bone (Figure 31B–C). A second scar marking the more distal portions of the mm. adductores femoris then continues along the posterior midline to the popliteal surface. The insertion of the m. quadratus femoris is rugose and concave (Figure 31B). The caudomedial fossa for the m. gastrocnemius is shallow and the caudolateral fossa for the m. flexor digitorum superficialis is very shallow. The distal femoral condyles are relatively low and the trochlea groove relatively shallow (Figure 31A–B, F). The medial condyle is transversely quite narrow. The femur of the B. planei holotype is approximately 33% larger in size than in D. lumholtzi and 20% larger than in B. illuminata. The greater trochanter is taller than in species of Dendrolagus, and its rugose apex is distinct. The base of the greater trochanter bears a distinct fossa on its cranial surface for the m. vastus lateralis, which is of similar depth in B. illuminata and B. nullarbora, but noticeably wider. The laterally flared distal extremity is most like Thylogale billardierii (Desmarest, 1822). The thickened lateral margin or crest upon which the m. gluteus medius inserts is proportionally shorter than in most macropodines, terminating proximally to the distal margin of the lesser trochanter, as in B. illuminata and species of Dendrolagus (Figure 7A–D). The lesser trochanter is more flared proximally than in species of Dendrolagus, Setonix and Thylogale, and may have been similar in B. illuminata and B. nullarbora, though the area is abraded in the only known specimens of each species. The shaft is similar in relative robustness to B. illuminata, and less robust than in B. nullarbora and species of Dendrolagus (Figure 7). The adductor muscle scars are more marked than in B. nullarbora, while in B. illuminata there are narrow but relatively deep longitudinal scars for the mm. adductores femoris along the medial aspect of the femoral shaft as in Dendrolagus (Figure 7C; Warburton et al. 2012). The concave insertion of the m. quadratus femoris is unusual, in contrast to the typically raised tubercle in most macropodines. In this attribute, B. planei resembles T. billardierii. The diaphysis flares toward its distal end of shaft more than in species of Dendrolagus and terrestrial species but similar to B. illuminata and B. nullarbora. The shallow fossae for m. gastrocnemius and m. flexor digitorum superficialis contrast markedly with the deep fossae in ground-dwelling species of Macropus Shaw, 1790, Osphranter Gould, 1842 and Thylogale, but are similar to the morphology in B. illuminata and species of Dendrolagus. The craniocaudally compressed distal epiphysis is reminiscent of species of Dendrolagus (Figure 7F). The transversely narrow medial condyle is similar in morphology to B. illuminata. Tibia. The B. planei holotype preserves only the distal two-thirds of the tibia, the diaphysis of which is relatively straight, being very slightly sinuous from cranial view, and similarly in the cranial face of the proximal portion from the lateral view (Figure 32B). Distally, the diaphysis is sub-rectangular in cross-section. The distal fibular facet is estimated to have extended for approximately the distal two-fifths of the tibial shaft. The interosseous border above the distal fibula facet is moderately rounded. The distal epiphysis is equal in width to the end of the diaphysis (Figure 32C). The medial malleolus is robust, moderately long and aligned in the parasagittal plane (Figure 32C). The talar articular facet is aligned square to the craniocaudal axis (Figure 32E). In overall shape, the tibial diaphysis and proportional length of the distal fibular facet of B. planei is very similar to that of B. illuminata. The rounded interosseous border is in contrast to the sharper interosseous crest observed in B. illuminata, B. nullarbora and the species of Dendrolagus (Figure 8) and Thylogale, but it is less rounded than in Setonix brachyurus. The distal epiphysis and talar articular facet are relatively narrower than in B. illuminata and species of Dendrolagus, and are not as laterally flared as in D. bennettianus. The medial malleolar process is more distally extended than in species of Dendrolagus, but narrower anteroposteriorly than in species of Thylogale and Setonix. In distal view, the process of the medial malleolus is more obliquely orientated than in other species of Bohra and Dendrolagus (Figure 8F), being more similar to Thylogale in both this and in cranial view, in which it is relatively flat and horizontal. Calcaneus. The calcaneus is stout, robust, broad and dorsoventrally compressed (Figure 33A–G). The tuber calcanei is expanded caudally and constricted cranially (Figure 33B). The rugose plantar surface is very broad, and is expanded along the length of the medial margin by a distinct flange (Figure 33A). The mid-lateral margin is particularly enlarged. The epiphyseal line on the rugose plantar surface is raised into a distinct, ventrally projected ridge. Paired tubercles (the lateral one much larger) on the craniolateral extremity of the plantar surface are positioned adjacent to the groove for the peroneal tendons, and are bounded on the opposite side by a projection cranial to the attachment site of the anterior calcaneofibular ligament. The transverse flexor sulcus is not well defined, being relatively shallow and with irregular margins. The sustentaculum tali is very broad, particularly caudally, extending well beyond the medial edge of the talar articular surface (Figure 33D). The ventral surface is smoothly curved and the flexor groove relatively deep, accentuated by the expanded medial margin of the plantar surface. The lateral and medial facets for the talus are very broad; the lateral facet is smaller, mesially constricted and smoothly contiguous with the oval medial facet. The fibular facet is very small, though the tubercles for the fibular ligaments, particularly the calcaneocuboid ligaments, are strongly developed. The anteromedial facet for articulation with the talar head is small and distinct. The calcaneocuboid articulation is transversely broad and dorsoventrally compressed (Figure 33C). The dorsomedial facet for the cuboid is rectangular, broader than deep, and convex in profile. The dorsolateral facet is narrower and subtriangular in shape, tapering from dorsal to ventral. The step between the two facets is distinct, only slightly smoothed and obliquely orientated in dorsal view (Figure 33A–G). The ventromedial cuboid facet is abraded on the holotype, but was apparently relatively small and smoothly continuous with the dorsolateral facet, but separate from the dorsomedial facet. The calcaneus of B. planei is similar overall in morphology to B. illuminata, B. nullarbora and the species of Dendrolagus (Figure 9). The distinct ridge formed at the plantar epiphyseal line distinguishes B. planei from all other species, but is most similar to D. bennettianus (Figure 9B), and may reflect augmentation of the digital flexor muscles in the pes. The rugose tuberosities at the cranial margin of the plantar surface are similar to those observed in B. wilkinsonorum (Figure 20A). The flaring of the medial plantar margin of the calcaneal tuber and flexor groove are developed to a greater extent than in other Bohra species. The morphology of the talar facets is most similar to that of B. paulae (Figure 12A), but even more confluent and more dorsoventrally flattened. In all other species of Bohra, the margins of the medial and lateral facets are more clearly delineated, even where they abut each other. The calcaneocuboid step is deeper and more angular than in other species of Bohra and Dendrolagus, being most similar to B. wilkinsonorum, and also approaching the depth in species of Setonix and Thylogale. When viewed dorsally, the cranial extremity (cuboid articulation) is relatively much narrower (does not extend to the width of the fibular facet) in comparison to other species of Bohra. The cuboid facets are relatively deep and narrow compared to other species of Bohra. The separation of the ventromedial facet appears most like B. wilkinsonorum. Talus. The talus is wider than long, dorsoventrally compressed (especially in cranial view) with low trochlear crests, which are subequal in height, and a shallow trochlea groove (Figure 33H–M). The malleolar fossa is moderately wide and shallow. The talar neck is very short. The talar head is medially displaced relative to the medial trochlear crest and transversely quite broad in cranial view. The articular facet for the navicular is ovoid around an axis that is approximately 40° from a dorsoventral plane (Figure 33M). The calcaneal facets are transversely broad, moderately deep and smoothly contiguous. The posterior plantar process is bulbous, transversely broad and dorsoventrally compressed. The trochlear groove is shallower than in B. illuminata, B. nullarbora, D. bennettianus (Figure 9H–I, K) and D. lumholtzi, but similar to that of B. sp. indet. 1. The medial trochlear crest is lower than in B. paulae, and also more similar to that of B. sp. indet. 1. The malleolar fossa is broad and shallower in B. planei than in other species of Bohra and species of Dendrolagus. The medial malleolus is broad and flattened, similar to the conditions seen in B. paulae and B. sp. indet. 1. It is relatively less projected than in B. nullarbora and species of Dendrolagus, Dorcopsulus, Setonix and Thylogale. The very short talar neck is similar to the morphology seen in species of Dendrolagus (Figure 9H), and is shorter than in B. illuminata and B. nullarbora, but not as short as in B. paulae (Figure 12K). The navicular facet of the talar head is set at a less oblique angle than in other species of Bohra, including B. paulae and especially B. illuminata and species of Dendrolagus, resulting in a narrower and more elongate outline. The lateral border of the navicular facet lies outside of, rather than in line with, the cranial extremity of the medial crest, as in B. paulae and D. bennettianus, D. lumholtzi. The talocalcaneal facets are smoothly conjoined and relatively shallow. The posterior plantar process is short and broad, as in B. nullarbora and B. illuminata, but proportionally smaller than in species of Dendrolagus. Remarks. Bohra planei is the only species i

Bohra paulae Flannery & Szalay, 1982 Bohra pauli: Dawson (2004a), p. 271, table 1; lapsus calami. Holotype. AM F62099 / AM F62100, right adult calcaneus and talus. The holotype was originally based solely on the calcaneus (AM F62099); the talus was listed and described separately as a referred specimen (AM F62100). However, given their precise anatomical fit and similar preservation, it is beyond reasonable doubt that these two elements belong to the same individual. Type locality. Wellington Caves (site not specified), New South Wales. No specific locality or collection details are associated with the holotype of B. paulae. However, the type locality is very likely a Pleistocene deposit (see ‘Locality information’). Referred specimens. Cathedral Cave, Wellington Caves. AM F62101, left tibia. AM F62102, left tibia (proximal half). Collected by Henry Barnes of the Australian Museum in 1881 (Ramsay 1882). The stratigraphic origins of the tibiae are unknown, but Ramsay’s Shaft No. 2, positioned in the alcove south of the ‘Altar’ formation in the main chamber, evidently produced much of the larger fossil material collected from Cathedral Cave in 1881 (Ramsay 1882). Wellington Caves (site not specified). AM F103802, adult right metatarsal V. AM F104075, left juvenile metatarsal IV. AM F104092, adult right metatarsal V. AM F104550, juvenile left metatarsal IV. These specimens are part of the ‘Old Collection’ of the Australian Museum (Dawson 1985), and probably originate from a Pleistocene deposit based on the same line of reasoning as for the holotype. Etymology. Named in honour of Flannery & Szalay’s draftsperson Paula Kendall. Revised diagnosis. A large species of Bohra, closest in size to B. wilkinsonorum in calcaneal dimensions. The species is distinguished by the following combination of calcaneal features: dorsal half of tuber calcanei barrel-shaped; plantar aspect less flared posteriorly, particularly laterally, than in the similarly large B. wilkinsonorum; medial and lateral talar facets of calcaneus continuous anteriorly but with facet contours distinct; calcaneocuboid articulation transversely broad, with step between dorsal facets smoothed, and confluent with ventromedial facet; talar and cuboid articular surfaces separated by short distance. The talus is distinguished from that of other species of Bohra by having a shallow trochlear groove that is slightly deeper medially, and a very short talar neck. The tibia is relatively more robust than in B. illuminata and B. nullarbora, particularly distally. The metatarsals IV and V are more robust than those of B. illuminata and B. nullarbora, and the metatarsal IV diaphysis is proportionally deeper anteriorly than in B. nullarbora. Description and comparisons. Tibia. The tibia of B. paulae (Figure 11) is significantly larger (Table 8) and more robust than that of B. illuminata and is more similar to species of Dendrolagus (Figure 8) in its stockiness. Other aspects of its morphology, including the relative size of the anterior tibial crest (Figure 8A), the curvature of the shaft, and the length of distal fibular facet (Figure 8B) are similar between B. paulae and B. illuminata, although the proximal fibular facet (Figure 8D) is more pronounced in B. paulae. The tibial crest is shorter and the lateral fossa for m. tibialis cranialis (Figure 8D) is deeper in both B. paulae (Figure 11) and B. illuminata than in species of Dendrolagus. Caudolaterally, the interosseous border of the tibia is more sinuous than that of B. illuminata and B. nullarbora, but less so than in species of Dendrolagus. The tibial diaphysis is more flared distally than in B. illuminata and B. nullarbora. The distal articular surface of AM F62101 has sustained significant post-mortem damage from rodent gnawing (Figure 11F), but the relatively short medial malleolus reflects the corresponding morphology of the talus. Calcaneus. The plantar surface of the holotype calcaneus was evidently expanded posteromedially, but the specimen is abraded in this area (Figure 12B). It is also abraded on the lateral side (Figure 12C), but the contour of the surrounding bone allows us to infer that it was flared to a lesser degree than in B. wilkinsonorum (Figure 20A–B). Indeed, the tuber calcanei of B. paulae is distinctly barrel-shaped in dorsal view (Figure 12A), and not as constricted anteriorly as in other tree-kangaroo species. Otherwise, the calcaneus is similar in general proportions and size to that of B. wilkinsonorum (Table 8). The distance between the talar and cuboid facets is even shorter in B. paulae than in other tree-kangaroos, e.g., D. bennettianus (Figure 9A). Although extended medially, the sustentaculum tali is relatively short, not extending as far posteriorly as in other tree-kangaroo species (Figure 9A–B). The plantar surface is flat, only slightly rugose, and very broad anteriorly (Figure 12B). The non-rugose flexor groove is shallow and wide, but not as wide as in D. lumholtzi (Figure 9B). The medial and lateral talar facets in B. paulae are broadly continuous (Figure 12B, E) and more smoothed than in D. bennettianus (Figure 9A), D. lumholtzi, B. illuminata and B. nullarbora. The lateral facet is only slightly tapered mesially, and the medial facet is large, ovoid and almost transverse in alignment (Figure 12A). The calcaneocuboid articulation is relatively shallow, and the characteristic macropodid ‘step’ between the medial and lateral facets is not much more than an undulation (Figure 12B, E, H; cf. Figure 9A). The ventromedial facet is broad and completely continuous with the medial and lateral facets (cf. Figure 9D). Overall, the cranial end of the calcaneus closely resembles that of D. lumholtzi. The sulcus for the m. flexor digitorum longus is short and narrow compared with all other species (cf. Figure 9B). Talus. The talus of B. paulae is short and wide, but the medial crest is moderately high and steep, and the lateral crest is low and broad, resulting in a moderately shallow but asymmetrical trochlear groove (Figure 12H, J). This marked asymmetry in crest contour is similar to that observed in D. lumholtzi, but less so in D. bennettianus (Figure 9H, K) and other species of Bohra, where the crests are more equally pronounced. The talar neck is very short (Figure 12K, N, L), shorter in fact than in all other tree-kangaroos for which the talus is known (e.g., Figure 9H). The navicular facet is slightly deeper relatively than that of D. lumholtzi, and most similar to that of D. bennettianus (Figure 9K). The malleolar fossa is broad and shallow in comparison to the conditions observed in B. nullarbora, D. bennettianus and the species of Dorcopsulus, Petrogale Gray, 1837, Setonix Lesson, 1842 and Thylogale Gray, 1837. The posteroventral process is similar in its posterior extension to the condition observed in B. illuminata, B. nullarbora, D. lumholtzi and D. bennettianus (Figure 9G), but more extended than in B. planei (Figure 33J). Metatarsals. Two juvenile metatarsal IV specimens are referred to B. paulae (Figure 13A–J) on the basis of their overall proportions and relative robustness, and the transversely wide, conjoined articular facet for metatarsal V, as in species of Dendrolagus (Figure 10B). AM F104075 is more robust and slightly shorter, and has a broader, more posteriorly restricted plantar ridge compared with AM F104550. Otherwise, they are very alike and morphologically distinct from the B. illuminata holotype metatarsal IV, which is also not fully grown and missing the distal epiphysis, and from the B. nullarbora holotype metatarsal IV, which is fully grown and has a fused distal epiphysis. The proximal articular facet for the cuboid is convex dorsally, while the peninsula of the cuboid facet that extends ventrally onto the posterior face of the plantar tuberosity is concave (Figure 13E, J). In comparison, the dorsolateral corner of the cuboid facet is more pointed in B. illuminata, while the ventral peninsula of the facet is much broader in B. nullarbora. In B. paulae, the plantar sesamoid facet on the anteroventral face of the plantar tuberosity abuts the conjoined metatarsal V facets (AM F104550) or is separated from them by a 1-mm gap (AM F104075), as in the B. illuminata holotype, while the gap is closer to 2 mm in the B. nullarbora holotype and D. bennettianus (Figure 10B). The metatarsal IV of B. paulae is more robust (broader relative to length) than that of the B. illuminata holotype. AM F104550 and the B. nullarbora holotype are very similar in size and general proportions, despite the ontogenetic difference between the specimens. However, the diaphysis of AM F104550 is comparatively deeper cranially (less concave ventrally). Two metatarsal V specimens are referred to B. paulae (Figure 13K–R) on the basis of their absolute size, proportions and distinctive robustness. Both are complete and fully grown with fused distal epiphyses. The proximal articular facet for the lateral facet of the cuboid is roughly triangular and concave, sweeping up onto the broad proximolateral process. The proximomedial surface has a single broad articular facet for metatarsal IV. There only noteworthy differences between them are that AM F103802 is 11% larger than AM F104092. AM F103802 and AM F104092 are 25% and 15% larger, respectively, than the B. nullarbora holotype metatarsal V (Table 8), and slightly broader posteriorly (evident primarily in the plantar rugosity). Other points of morphological distinction in metatarsal V morphology between B. paulae and B. nullarbora include a relatively larger surface area of the metatarsal IV facet, shorter proximolateral process and commensurately less deeply flexed cuboid facet in B. paulae. From the smaller, ontogenetically younger B. illuminata holotype, B. paulae differs by having a more rounded medial end of the metatarsal IV facet and a more rugose interosseous ligament attachment (Figure 13M, Q). Remarks. We are confident that the only known calcaneus and talus of Bohra paulae once articulated with each other and, as Flannery & Szalay (1982) originally suggested, that the two referred left tibiae belong to the same species as the tarsal bones. Both tibiae were collected from Cathedral Cave within the Wellington Caves complex (Figure 1), likely from Ramsay’s Shaft No. 2. Subsequent excavations within this area of the cave led by Lyndall Dawson & others in 1982–1986 (Dawson & Augee 1997) and by Diana Fusco and GJP since 2016 have revealed close spatial associations between limb elements of large macropodid individuals. It is conceivable that the holotype not only came from Shaft No. 2, but also belongs to the same individual as one of the referred tibiae. By contrast, ‘ Wellington Caves’ is the only provenance information associated with the four metatarsals. Their larger size and morphological distinction from the metatarsals of B. illuminata and B. nullarbora prompt us to refer them to B. paulae rather than B. bandharr. Basic metrics (Table 8) reinforce this: the holotype calcaneus of B. paulae is 23% longer than that of B. nullarbora, while metatarsal V length (mean of AM F103802 and AM F104092) is also 23% greater than that of B. nullarbora. Similarly, calcaneus length to metatarsal V length (B. paulae holotype against mean of AM F103802 and AM F104092) is 0.72, as it is for the B. nullarbora holotype.

Bohra Flannery & Szalay, 1982 Type species. Bohra paulae Flannery & Szalay, 1982. Revised diagnosis. Species of Bohra are distinguished from those of Dendrolagus based on the following craniodental features: larger cheek teeth relative to the size of the cranium; higher-crowned upper incisors; narrower P3, which lacks a large, distinct posterobuccal accessory cusp and deep cleft posterior to the anterior cusp of the main crest; relatively narrower molars, with a better developed postparacrista and no parametacristid. All known species of Bohra are also larger than those of Dendrolagus. Bohra most closely resembles Dorcopsis and Watutia in molar morphology, but the premetacristid, cristid obliqua and postprotocrista are less developed than in Dorcopsis, and the molars are narrower relative to their length than in Watutia. Species of Bohra may be distinguished from those of Dendrolagus on the basis of the following postcranial features: slightly less robust humerus, with a more pronounced pectoral crest, a slightly less developed deltoid tubercle, and a larger, more projecting medial epicondyle; relatively more robust femoral diaphysis; less sinuous tibial diaphysis with a relatively longer distal fibular facet; less obliquely orientated trochlear crests and groove on the talus; shorter and more robust metatarsals IV and V. Etymology. According to a legend of the Euahlayi people of inland eastern Australia, Bohra was a kangaroo that went about on four limbs and possessed canines before men removed them (Parker 1953). Flannery & Szalay (1982) considered this evocative of extant tree-kangaroos, which have relatively more equally proportioned fore- and hindlimbs and retain small canines. They did not designate a gender, but with the naming of B. illuminata, Prideaux & Warburton (2008) inferred that it was feminine. Temporal and geographic distribution. Late Pliocene of southeastern Queensland, central eastern New South Wales, northern South Australia, and Papua New Guinea. Early and/or middle Pleistocene of central eastern New South Wales and southern South Australia. Middle Pleistocene of eastern Queensland, southeastern Western Australia and northern South Australia. See Figure 1., Published as part of Prideaux, Gavin J. & Warburton, Natalie M., 2023, A review of the late Cenozoic genus Bohra (Diprotodontia: Macropodidae) and the evolution of tree-kangaroos, pp. 1-95 in Zootaxa 5299 (1) on page 21, DOI: 10.11646/zootaxa.5299.1.1, http://zenodo.org/record/8012666, {"references":["Flannery, T. F. & Szalay, F. S. (1982) Bohra paulae: a new giant fossil tree kangaroo (Marsupialia: Macropodidae) from New South Wales, Australia. Australian Mammalogy, 5 (2), 83 - 94.","Parker, K. L. (1953) Australian Legendary Tales. Halstead Press, Sydney, 237 pp.","Prideaux, G. J. & Warburton, N. M. (2008) A new Pleistocene tree-kangaroo (Diprotodontia: Macropodidae) from the Nullarbor Plain of south-central Australia. Journal of Vertebrate Paleontology, 28 (2), 463 - 478. https: // doi. org / 10.1671 / 0272 - 4634 (2008) 28 [463: ANPTDM] 2.0. CO; 2"]}

Prideaux, Gavin J., Warburton, Natalie M. (2023): A review of the late Cenozoic genus Bohra (Diprotodontia: Macropodidae) and the evolution of tree-kangaroos. Zootaxa 5299 (1): 1-95, DOI: https://doi.org/10.11646/zootaxa.5299.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5299.1.1

Bohra illuminata Prideaux & Warburton, 2008 Bohra sp. nov. 1: Prideaux (2006), p. 1524. Prideaux et al. (2007), p. 423, table 1. Bohra sp. indet.: Prideaux & Warburton (2008), pp. 463–464, figure 1 (partim). Prideaux & Warburton (2009), p. 166, figure 1 (partim). Holotype. WAM 03.5.10, partial adult skeleton including cranium, partial left and right dentaries. Minimal wear on P3 and M4 indicate that these teeth had only recently erupted. This reveals that the specimen is a young adult not yet fully grown as confirmed by the unfused epiphyses of vertebrae and limb elements. Postcranial elements: four cervical, four thoracic, five lumbar, and eight caudal vertebrae; numerous partial ribs; proximal fragments of right and left scapulae; left clavicle; right humerus (missing proximal epiphysis); right ulna; fragments of radius; triquetrum, hamatum; metacarpals; manual phalanges; innominate fragments, including partial left and right ilia with acetabular fossa; fragments of left femur and near-complete right femur; left and right tibiae; left calcaneus; right talus; right cuboid; partial left and right metatarsals IV; partial left metatarsal V; pedal phalanges, including left digits IV – V. Revised diagnosis. Bohra illuminata is distinguished from B. nullarbora and the species of Dendrolagus in the following cranial features: a distinct buccinator fossa; a less distinct supraorbital crest above the posterior end of the eye orbit; a broad groove between the supraorbital crest and superior lacrimal rugosity, which produces a constriction of the frontals; a relatively shallow zygomatic arch; a strongly domed dorsal neurocranial surface; a distally broader, more anteriorly hooked postglenoid process. The cheek teeth are large relative to the dimensions of the palate. P3 bears a distinct posterolingual cusp, but only the slightest hint of a thin cingulum anterior to it, which distinguishes B. illuminata from all other species for which P3 is known. The dentary is shallower relative to the height of the lower molars than in all other tree-kangaroo species. The i1 of B. illuminata has thicker dorsal and ventral enamel flanges than in B. nullarbora. Lower molars bear a strongly developed premetacristid. Compared with that of B. nullarbora, the clavicle of B. illuminata is more gracile and flattened, with less expanded articular facets and a more medially positioned anterior inflection. The humerus is more gracile with a weaker pectoral crest, deltoid ridge, bicipital groove and spinodeltoid insertion than in B. nullarbora, but with a higher m. teres major insertion. The distal articular surface of the humerus is relatively broad. The ilium is relatively narrower anteroposteriorly, with a straighter medial border compared with that of B. nullarbora. The distinct long adductor muscle grooves on the medial aspect of the femur distinguish B. illuminata from B. nullarbora and B. planei, and in this attribute resembles the species of Dendrolagus. The tibia of B. illuminata is smaller and more gracile than that of B. paulae, and the interosseous margin is less sinuous in cranial view. The talar articular surface on the calcaneus is wider and the medial malleolus is more obliquely orientated in distal view than in B. planei. The calcaneus has a longer anterior plantar sulcus than in B. nullarbora, and more confluent talar facets with a mesially tapered lateral talar facet. The cuboid facets on the calcaneus are smoothly confluent and the calcaneocuboid step is very shallow and smoothed. Metatarsal IV is less robust than in B. nullarbora, with a smaller cuboid facet, a more medially constricted facet for metatarsal V, and a more rounded, distally orientated sesamoid facet. Metatarsal V is less robust. Type Locality. Metasthenurus Chamber, Last Tree Cave, Nullarbor Plain, southeastern Western Australia. Most of the holotype was collected from a small grotto on the north side of the chamber by John A. Long, Lindsay M. Hatcher and GJP in July 2002 and April 2003. However, the right femur and several non-descript postcranial fragments were collected by Lindsay M. Hatcher and GJP in April 2014. Fossils from the floor of Last Tree Cave, although previously inferred to be of early middle Pleistocene age (Prideaux et al. 2007), are actually undated, but include specimens of several species known from well-dated middle and late Pleistocene sites elsewhere. At this point, the type locality age is best considered conservatively as Pleistocene. The precise location from which the holotype was collected is recorded with the Department of Earth and Planetary Sciences, Western Australian Museum. Paratype. Main Chamber, Last Tree Cave. WAM 02.7.16, partial juvenile skeleton. Craniodental elements: left P3, M1–2, right maxilla fragment (preserving M1); left i1, partial right dentary (preserving m 1–2 in occlusion, m 3 in crypt, isolated p3); Postcranial elements: fragments of numerous vertebrae including 14 caudal vertebrae; numerous rib fragments; proximal fragment of left scapula; proximal fragment of right humerus; numerous carpals and metacarpals; innominate fragments including left and right ilia with acetabula, right ischium and epipubic; proximal fragments of femora; distal epiphysis of right fibula; fragments of right and left proximal tibia epiphyses; distal fragment and epiphysis of right tibia; left and right calcanei (right with separate epiphysis); left talus; left cuboid; left navicular; left and right metatarsals IV; right metatarsal V (missing distal epiphysis); left proximal, left and right medial and distal phalanges of digit IV, plus several other distal phalanges. This specimen was collected by GJP on 7 July 2002 from beneath a boulder overhang on the east side of the southern end of the chamber, the deepest point in the cave at 40 m depth. Referred specimens. RF 95 fissure fill, Curramulka Quarry, Yorke Peninsula, South Australia. SAMA P55401, right M4; SAMA P55402, right M4; SAMA P55403, left M4; SAMA P55406, left M3; SAMA P55408, left m1, right m1 (these teeth are identical in morphology, wear and preservation, so are considered to belong to the same individual); P55409, right m2; SAMA P55410, right m1; SAMA P55416, left M1. These specimens were collected by James A. McNamara (South Australian Museum) in the mid-1990s. The deposit is considered to be of Pleistocene age. New Kalamurina, Warburton River, South Australia. SAMA P59526, left i1. Collected by Neville S. Pledge (South Australian Museum), 9 March 1972. Middle to late Pleistocene. Etymology. This refers to the insights provided by the holotype, the first-known specimen of a species of Bohra represented by associated hindlimb and cranial elements. This confirmed that Bohra paulae, known only from hindlimb elements, and B. wilkinsonorum, known only from one partial maxilla, were indeed congeneric, as argued by Dawson (2004a). Description and comparisons. Cranium. Previously described in detail (Prideaux & Warburton 2008). The incisor-bearing portion of the premaxilla (Figure 23A) is similar in proportion and relative depth to that observed in Dendrolagus bennettianus and D. inustus, but is shallower than in B. nullarbora and relatively longer than in most species of Dendrolagus.The narial aperture is slightly broader than deep(Figure23B),as in B.nullarbora,but the lateral margins are more concave than in species of Dendrolagus.The buccinator fossa is moderately deep(Figure 23A),much more distinct than in B. nullarbora and species of Dendrolagus, in which there is only the barest hint of a concavity immediately above the diastema. The masseteric process is small, extending ventrally to about the level of the basal enamel margin of M2, as in species of Dendrolagus. Above the posterior extremity of the eye orbit, there is a swelling of the frontal, triangular in lateral view, that is bounded dorsally by a slight supraorbital crest (Figure 23A).Above the centre of the orbit, a broad groove (Figure 23A, C) separates the supraorbital crest from the superior lacrimal rugosity. This groove manifests itself in dorsal view as a constriction of the frontals. This combination of supraorbital features distinguishes B. illuminata from all other tree-kangaroo species. Posteriorly, the supraorbital crest is more markedly projected laterally in B. nullarbora, and much smoother and lower over its entire length in species of Dendrolagus. The zygomatic process of the squamosal (Figure 23A–B, E) is shallower relative to the depth of the cranium than in all other tree-kangaroo species. The marked doming of the dorsal surface of the neurocranium, which is typically a feature of young juvenile macropodid individuals, is unique to B. illuminata among young adults or subadults of tree-kangaroo species. The relatively narrow ectoglenoid process (Figure 23D) and the manner in which the postglenoid process is broad at its distal extremity and curved anteriorly, providing the glenoid fossa with a semi-circular shape when viewed laterally (Figure 23A), are unique among tree-kangaroo species. Upper dentition. Bohra illuminata is most easily distinguished from all other tree-kangaroo species by having larger cheek teeth relative to the overall proportions of the palate and a weakly developed lingual cingulum on P3, but with a distinct cuspule on the anterolingual corner of the crown (Figures 23D, 24A). In addition, the central cuspules of the P3 main crest are very small and only just distinguishable (Figure 24A–B). The upper molars of B. illuminata (Figures 24A, C, 25A–J, Table 5) are most similar in size and morphology to those of B. bandharr, differing only by being marginally less flat-sided buccally. The upper molars are also similar to those of B. nullarbora, but differ by being slightly larger overall and longer relative to their width, by having a flatter lingual side of the crown base, and by having a low cingulum bordering the interloph valley lingually. They differ from those of B. bila by lacking any remnant cusp C region of the stylar crest, and by having a less incurved postparacrista. Dentary. The horizontal ramus of B. illuminata (Figure 23F–I) is shallower relative to the height of the lower molars than in B. nullarbora, B. bila and the species of Dendrolagus, although this may be because the holotype is a subadult. The mandibular condyle is not positioned as high above the level of the molar row as in B. nullarbora. The digastric sulcus is less distinct in B. illuminata than in B. nullarbora and B. bila, and the anterior insertion area for the internal superficial masseter muscle is thinner and restricted to the rim of the masseteric fossa. The general profile of the posterior end of the dentary is otherwise similar between these three species. Lower dentition. Although similar in size and overall morphology, the i1 of B. illuminata (Figure 24I–K) has thicker dorsal and ventral enamel flanges than that of B. nullarbora. The unerupted p3 of the paratype, WAM 02.7.16 (Figure 24D), although lacking its full complement of enamel, was clearly swollen posteriorly, but the posterobuccal eminence is not as large as in the holotype of B. bila. The lower molars (Figures 24E–H, 25K–O) closely resemble those of B. nullarbora, but they are larger and the premetacristid is more strongly developed, especially on m3–4, a morphology reminiscent of the condition seen in the species of Dorcopsis. The lophids of B. illuminata are slightly more swollen than in B. nullarbora, and the buccal and lingual sides of the interlophid valleys are less concave. Clavicle. The clavicle is more gracile and dorsoventrally flattened than in B. nullarbora, with reduced articular facets and a less laterally positioned cranial inflection. Humerus. The humerus is less robust than in B. nullarbora, with a weaker pectoral crest, deltoid ridge, bicipital groove and spino-deltoid insertion, and a higher teres major insertion. Ilium. The ilium is relatively narrower craniocaudally than in B. nullarbora, with a less sinuous medial border. Femur. The holotype right femur is abraded near its proximal epiphyseal margin, and the proximal and distal articular epiphyses are missing (Figure 26). The distal end of the lateral trochanteric crest terminates craniad of the distal margin of the lesser trochanter (Figure 26A). The trochanteric fossa is relatively open (Figure 26B), as in species of Dendrolagus (Figure 5A). The shaft is cranially convex (Figure 26C–D), less robust than in B. nullarbora, and tending to ovoid in cross-section rather than circular, reminiscent of the species of Dendrolagus (Figure 5A–D). Highly distinct muscle scars – one on the proximocaudal surface immediately distal to the trochanteric fossa and two elongate, depressed scars on the medial surface of middle third of the femur – appear to correspond to the three portions of the mm. adductores femoris (Figure 26C). These are much more strongly marked than in B. nullarbora, which has no sign of the proximal one, and B. planei in which the adductor scars are more rugose than depressed and more caudally than medially positioned, but are similarly strongly developed in species of Dendrolagus (Figure 7C; Warburton et al. 2012). The quadratus femoris tubercle is more raised than in B. nullarbora and B. planei. The distal fossa for the medial gastrocnemius is very shallow, even shallower than in species of Setonix and Thylogale, and most similar to those of B. nullarbora and species of Dendrolagus (Figure 7C). This is also the case with the very shallow popliteal fossa. The distal epiphysis resembles that of the species of Dendrolagus (Figure 7F), but the fibular groove of the lateral condyle is less reduced and the cranial epiphyseal width is only slightly less than the caudal width in B. illuminata. Tibia. The tibia of B. illuminata is smaller and more gracile than that of B. paulae (Figure 11). The two species are similar in the relative size of the tibial crest, curvature of the shaft and length of fibular contact, but the proximal fibular facet of B. illuminata is less pronounced than in B. paulae. In cranial view, the interosseous margin of the tibia is less sinuous than in B. nullarbora, and particularly in B. paulae. From the distal view, the talar articular surface is wider and the medial malleolus set at a more oblique angle in B. illuminata than in B. planei (Figure 32). Calcaneus. The calcaneocuboid step is much shallower and more smoothed than in all other species of Bohra, with the possible exception of B. paulae. Compared with that of B. nullarbora, the calcaneus of B. illuminata is smaller (which in part is very likely due to its younger ontogenetic age), lacks a partial separation of the talar facets, and has a longer anterior plantar sulcus, a poorer developed plantar tuberosity, a narrower sustentaculum tali, and a relatively wider, more mesially tapered lateral talar facet. Distinguished from that of B. paulae by being smaller, with a relatively shorter and less barrel-shaped tuber calcanei, shorter and wider talar facets, a longer sustentaculum tali, a larger anteroplantar tuberosity, a relatively smaller ventromedial cuboid facet and relatively wider dorsolateral cuboid facet. The calcaneus of B. illuminata is distinguished from that of B. wilkinsonorum in being much smaller in absolute size (Table 8), with much smoother and confluent talar and cuboid facets. The transversely wider, smoother, and more confluent cuboid facets similarly set the calcaneus of B. illuminata apart from that of B. planei (Figure 33). Talus. The holotypes of B. illuminata and B. nullarbora are very similar in talar morphology, but two clear differences are evident. The facets for articulation with the calcaneus on the ventral surface are smoothly conjoined in B. illuminata, whereas they are separated by a distinct groove in B. nullarbora. In addition, the malleolar fossa in B. illuminata is narrower than in B. nullarbora. Although the talus of B. illuminata is smaller and has a shallower trochlear groove, both of these differences may well be due to its younger ontogenetic age. All other differences evident between the two specimens appear to be superficial and the result of the incompleteness of several features due to post-depositional abrasion or breakage. By comparison, B. paulae (Figure 12K) and B. planei (Figure 33I) are characterised by a shorter talar neck and shallower malleolar fossa from the dorsal view, and a broader, less obliquely orientated navicular facet in cranial view. Metatarsals. The metatarsal IV of the B. illuminata holotype is less robust than in B. nullarbora and B. paulae, and the metatarsal V facet is more medially constricted. In B. illuminata, the dorsolateral corner of the cuboid facet is more pointed than in B. nullarbora and B. paulae, while the ventral peninsula of this facet is intermediate in relative breadth between the conditions manifested in the other two species. The proximity of the plantar sesamoid and metatarsal V facets in B. illuminata is intermediate between that observed in B. nullarbora and B. paulae. The metatarsal V of the B. illuminata holotype is less robust than that of B. nullarbora, but this may simply be because it is not fully grown. It also differs by having a relatively broader metatarsal IV facet. By comparison with B. paulae (Figure 13), B. illuminata differs by having a less rounded medial end of the metatarsal IV facet. It differs from both B. nullarbora and B. paulae by having a more rugose interosseous ligament attachment. Remarks. Most of what we know about B. illuminata is as it was in 2008, but the subsequent description of B. nullarbora, the belated 2014 discovery of the B. illuminata holotype right femur, and the allocation of pedal remains to other species have allowed the diagnosis and comparisons to be refined. Our understanding of the geographic distribution of B. illuminata has also been augmented through the referral to this species of loose molars from Curramulka Quarry, central Yorke Peninsula (Figure 1). Ancient DNA that aligned best with that of a tree-kangaroo has been recovered from a late Pleistocene archaeological deposit in Tunnel Cave, southwestern Australia (Figure 1; Murray et al. 2013), and may belong to B. illuminata or B. nullarbora, but confirmation of its tree-kangaroo identity must await further molecular work., Published as part of Prideaux, Gavin J. & Warburton, Natalie M., 2023, A review of the late Cenozoic genus Bohra (Diprotodontia: Macropodidae) and the evolution of tree-kangaroos, pp. 1-95 in Zootaxa 5299 (1) on pages 42-49, DOI: 10.11646/zootaxa.5299.1.1, http://zenodo.org/record/8012666, {"references":["Prideaux, G. J. & Warburton, N. M. (2008) A new Pleistocene tree-kangaroo (Diprotodontia: Macropodidae) from the Nullarbor Plain of south-central Australia. Journal of Vertebrate Paleontology, 28 (2), 463 - 478. https: // doi. org / 10.1671 / 0272 - 4634 (2008) 28 [463: ANPTDM] 2.0. CO; 2","Prideaux, G. J. (2006) Mid-Pleistocene vertebrate records: Australia. In: Elias, S. A. (Ed.), Encyclopedia of Quaternary Science. Elsevier, Oxford, pp. 1517 - 1537. https: // doi. org / 10.1016 / B 0 - 44 - 452747 - 8 / 00258 - 1","Prideaux, G. J., Long, J. A., Ayliffe, L. K., Hellstrom, J. C., Pillans, B., Boles, W. E., Hutchinson, M. N., Roberts, R. G., Cupper, M. J., Arnold, L. J., Devine P. D. & Warburton N. M. (2007) An arid-adapted middle Pleistocene vertebrate fauna from south-central Australia. Nature, 445 (7126), 422 - 425. https: // doi. org / 10.1038 / nature 05471","Prideaux, G. J. & Warburton, N. M. (2009) Bohra nullarbora sp. nov., a second tree-kangaroo (Marsupialia: Macropodidae) from the Pleistocene of the Nullarbor Plain, Western Australia. Records of the Western Australian Museum, 25 (2), 165 - 179. https: // doi. org / 10.18195 / issn. 0312 - 3162.25 (2). 2009.165 - 179","Dawson, L. (2004 a) A new Pliocene tree kangaroo species (Marsupialia, Macropodinae) from the Chinchilla Local Fauna, southeastern Queensland. Alcheringa, 28 (1), 267 - 273. https: // doi. org / 10.1080 / 03115510408619284","Warburton, N. M., Yakovleff, M. & Malric, A. (2012) Anatomical adaptations of the hind limb musculature of tree-kangaroos for arboreal locomotion (Marsupialia: Macropodinae). Australian Journal of Zoology, 60 (4), 246 - 258. https: // doi. org / 10.1071 / ZO 12059","Murray, D. C., Haile, J., Dortch, J., White, N. E., Haouchar, D., Bellgard, M. I., Allcock, R. J., Prideaux, G. J. and Bunce, M. (2013) Scrapheap challenge: a novel bulk-bone metabarcoding method to investigate ancient DNA in faunal assemblages. Scientific Reports, 3, 3371. https: // doi. org / 10.1038 / srep 03371"]}

Bohra wilkinsonorum Dawson, 2004a Holotype. QM F43277, right juvenile maxilla fragment (preserving dP2–3, M1–2; P3 removed from crypt). Type locality. Chinchilla Sand, Chinchilla Rifle Range (site not specified), north bank of the Condamine River, Chinchilla, western Darling Downs, Queensland. The holotype was collected by Doris and Cecil Wilkinson. Referred specimens. Probably Chinchilla. QM F675, right adult calcaneus. This specimen retains no associated locality or collection information, although it was likely collected in the late 19 th century. Chinchilla Sand is suggested by preservation. See earlier comments on specimens referred to B. sp. cf. B. bandharr putatively from Chinchilla. Childers Cove, Victoria. NMV P221950, left P3. Collected by David Pickering and others on 24 November 2011. NMV P222378, right M1. Collected by David Pickering and Wayne Gerdtz on 3 or 4 August 2004. Etymology. Named in honour of Doris and Cecil Wilkinson for their fossil collecting efforts at the Chinchilla Rifle Range locality. Revised diagnosis. The largest-known species of Bohra based on molar and calcaneal dimensions. The P3 is similar in size and morphology to that of B. bandharr, but is less markedly narrowed anteriorly. The postpara- and premeta-cristae on M2 are strongly developed and incurved, more so than in any other species of Bohra. The slight convexity of the buccal side and distinct interloph concavity of the lingual side of the M1–2 crown resembles that of the much smaller B. nullarbora. The species is also distinguished by the following combination of calcaneal features: massive, dorsoventrally compressed tuber calcanei; plantar aspect very broad and strongly flared posteriorly, particularly on the lateral aspect of the epiphysis; medial and lateral talar facets of the calcaneus discontinuous; pronounced and strongly demarked crest from the caudomesial border of the medial talar facet for the posterior talocalcaneal ligament; calcaneocuboid facets transversely broad, with oblique, moderately deep step; ventromedial facet separated from dorsomedial facet by deep fossa. Description and comparisons. Maxilla. Essentially all that is preserved of the holotype maxilla is that portion bearing the erupted dP2–3 and M1–2, the unerupted P3, and the masseteric process (Figure 19A–C). With its extremity positioned adjacent to the M2 protoloph in this juvenile specimen, the masseteric process is posteroventrally orientated and bears a low, slightly rugose crest along its anterolateral edge (Figure 19C). The juvenile maxilla is known for no other species of Bohra, and so no direct morphological comparisons are possible due to the changes that can occur from the ontogenetic stage where M2 has just come into occlusion versus a full adult where M4 is in occlusion. Nevertheless, it is worth noting that in D. bennettianus, the molars progress anteriorly relative to the position of the masseteric process between these two ontogenetic stages. When M2 first comes into occlusion, its protoloph is positioned adjacent to the masseteric process, but in adults the abutment of M2–3 is adjacent to the masseteric process. The same may well have been the case for B. wilkinsonorum. In shape, the masseteric process is more rounded than in B. illuminata and B. bila. Upper dentition. Previously described in detail (Dawson 2004a). The dP2–3 remain unknown for other species of Bohra, but comparisons with the much smaller D. bennettianus and D. inustus reveal very similar general crown outlines, although those of D. bennettianus are the closest match. Their dP2 crowns are relatively flat-sided oval shapes, and they share a low weak buccal cingulum. However, B. wilkinsonorum differs by lacking the large anterior cusp of the main crest bordered posteriorly by a deep cleft (Figure 19A–C), which is characteristic of both the dP2 and P3 of species of Dendrolagus. These species are also very similar in dP3 morphology, sharing: a strongly developed postparacrista, which abuts the end of the dP2 main crest; well-developed postpara- and premeta-cristae, which are anteroposteriorly orientated and confluent, rather than incurved as on M1–2; and a distinct preprotocrista that marks the lingual end of the transversely narrow precingulum. No preprotocrista is evident on the holotype M1, but there is a slight preprotocrista present on the unworn (unerupted) referred specimen, NMV P222378 (Figure 19L). There is a stylar eminence in the region of cusp C on M1, but not M2 (Figure 19C). Whether a urocrista (of which there is a slight manifestation in D. bennettianus and a very distinct expression in D. inustus) was also present in B. wilkinsonorum cannot be ascertained due to extensive wear of the metaloph. Bohra wilkinsonorum is most similar in P3 size and morphology to B. bandharr (Tables 1, 3). It differs by narrowing less markedly anteriorly, possessing three buccal cingular cuspules and bearing a slightly less pronounced posterobuccal eminence. Although the holotype P3 bears an anterior crest on the posterobuccal eminence (Figure 19D–E), this is lacking from NMV P221950 (Figure 19G–K). In M1–2 occlusal outline (Figure 19C, F, L), B. wilkinsonorum resembles the much smaller B. nullarbora. They share a slightly convex, but essentially quite flat, buccal side of the crown, a trait that also characterises B. bandharr. However, the lingual sides of M1–2 of B. bandharr are also flat lingually, rather than convex adjacent to the interloph valley. The strongly developed and incurved form of the postpara- and premeta-cristae on M2 distinguishes B. wilkinsonorum from all other species of Bohra. Absence from M2 of a stylar crest in the region of cusp C is a condition shared with B. bandharr, B. illuminata and B. nullarbora, but not B. bila and B. planei, in which this crest is distinct. Calcaneus. The referred calcaneus, QM F675, is well preserved and complete except for the caudolateral extremity of the tuber calcanei (Figure 20B, D). The calcaneus is large, stout, broad, stout and dorsoventrally flattened, with clearly defined articular surfaces and rugged ligamentous insertions. The tuber calcanei is strongly flared caudally (Figure 20B). From the plantar view, the medial margin diverges from the long axis more continuously along its length, in comparison to the lateral margin, which widens distinctly at the epiphysis. The rugose plantar surface is broad cranially, with a large anterolateral tubercle (Figure 20A). The transverse sulcus that separates the cranial margin of the rugose surface from the cuboid articulation is slightly oblique from medial to lateral sides. The sustentaculum tali is deep (from medial view), medially expanded and very slightly convex (Figure 20C). The medial and lateral talar facets are separated by a narrow but distinct, non-articular area (Figure 20B, E). The lateral facet is barrel-shaped (Figure 20B). The fibular facet is broad and rounded, and projects over the deep sulci for the posterior calcaneofibular ligament (Figure 20B, D). The tubercle and sulcus for the anterior calcaneofibular ligament are large, laterally expanded and rugose. The posterior margin of the oval medial talar facet projects dorsally and expands onto the dorsal surface toward the base of the calcaneal tubercle, with a marked projection for posterior talocalcaneal ligament (Figure 20B). Cranial to the talar facets, the mid-dorsal surface is has a deep scar for the ligamentum cervicis tali. The calcaneocuboid articulation is transversely very broad (laterally extending beyond the margin of the fibular facet; Figure 20B); and dorsoventrally compressed in cranial view (Figure 20E). The step between the rectangular dorsomedial and rounded dorsolateral facets is distinct, obliquely aligned and moderately smoothed (Figure 20A). The ventromedial facet is transversely broad, but dorsoventrally relatively shallow (Figure 20E). It is smoothly continuous with the dorsolateral facet but separated by a deep transverse groove from the dorsomedial facet. The lateral flaring of the tuber calcanei epiphysis is more pronounced than in all other species of Bohra, especially the similarly large B. paulae (Figure 12B). The transverse plantar sulcus is relatively deep and long in comparison to all other species of Bohra, in particular B. paulae and QM F51762 (Bohra sp. indet. 1), and is much deeper than in the species of Dendrolagus (Figure 9B). The expanded medial plantar margin of the sustentaculum tali accentuates the moderately deep flexor sulcus of the sustentaculum tali to a greater extent than in B. paulae, B. sp. indet. 1 and the species of Dendrolagus, though it is not as deeply flared as in B. planei. The gap between the medial and lateral talar facets resembles that observed in B. nullarbora, but differs from the condition seen in other species of Bohra and species Dendrolagus (Figure 9A), in which the facets are confluent. The calcaneocuboid articulation is transversely much broader than in any other species (Figure 20B, E cf. Figure 9B, D). The profile of the calcaneocuboid step and complete separation of the ventromedial and dorsomedial facets is most like B. planei, while the rounded and laterally expanded of the dorsolateral facet is more like other species of Bohra, in contrast to B. planei (Figure 33C). Remarks. The recognition of B. wilkinsonorum as a second member of the genus Bohra on the basis of one juvenile maxilla fragment (Dawson 2004a) was a highly perceptive deduction given that the type species was described and is still known only from hindlimb elements from a locality 665 km to the south (Flannery & Szalay 1982). Based on the dimensions of these specimens (Table 8), B. wilkinsonorum and B. paulae are the two largest species of Bohra, and one might be able to hypothesise on those grounds that the fossils belong to the same species. However, the recognition from the Darling Downs region (indeed, probably from the type locality) of a very large calcaneus clearly referable to a species of Bohra that is morphologically distinct from that of B. paulae supports its taxonomic separation. This calcaneus is much larger than, and differs in morphology from, the other two calcanei from the Chinchilla LF, which we refer to B. sp. indet. 1. Referral to B. wilkinsonorum of a loose upper premolar and molar from the putatively late Pliocene or early Pleistocene Childers Cove LF of southern Victoria (Rich et al. 2006) represents a major range extension of 1,500 km to the south (Figure 1)., Published as part of Prideaux, Gavin J. & Warburton, Natalie M., 2023, A review of the late Cenozoic genus Bohra (Diprotodontia: Macropodidae) and the evolution of tree-kangaroos, pp. 1-95 in Zootaxa 5299 (1) on pages 33-38, DOI: 10.11646/zootaxa.5299.1.1, http://zenodo.org/record/8012666, {"references":["Dawson, L. (2004 a) A new Pliocene tree kangaroo species (Marsupialia, Macropodinae) from the Chinchilla Local Fauna, southeastern Queensland. Alcheringa, 28 (1), 267 - 273. https: // doi. org / 10.1080 / 03115510408619284","Bartholomai, A. (1976 a) Notes on the fossiliferous Pleistocene fluviatile deposits of the eastern Darling Downs. Bureau of Mineral Resources, Geology and Geophysics, Bulletin, 166, 153 - 154.","Flannery, T. F. & Szalay, F. S. (1982) Bohra paulae: a new giant fossil tree kangaroo (Marsupialia: Macropodidae) from New South Wales, Australia. Australian Mammalogy, 5 (2), 83 - 94.","Rich, T. H., Piper, K. J., Pickering, D. & Wright, S. (2006) Further Ektopodontidae (Phalangeroidea, Mammalia) from southwestern Victoria. Alcheringa, 30 (1), 133 - 140. https: // doi. org / 10.1080 / 03115510608619349"]}

Dendrolagina subtribe nov. Included genera. Bohra Flannery & Szalay, 1982; Dendrolagus Müller, 1840 (tree-kangaroos). Diagnosis. Distinguished from other members of the Dendrolagini by the following combination of skeletal features: robust cranial rostrum; broader than deep occiput; absence of large palatal fenestrae; slightly broadened I1; broad lingual cingulum on P3; low-crowned molars; fine, low postprotocrista on upper molars; masseteric canal extends to posterior cheek teeth and separate from dental canal; posterobuccal eminence on main crest of p3; rectus femoris origin positioned immediately adjacent to acetabular rim; short, broad iliopubic process; medially displaced quadratus femoris scar; short, broad calcaneus with posteromedially expanded, dorsoventrally compressed tuber calcanei; large, medially expanded sustentaculum tali, extending far beyond the medial margin of the articular facets; relatively large, transversely broad calcaneotalar articulation, marked medial taper of lateral talar facet on calcaneus; laterally extended tubercle for calcaneofibular ligaments; relatively broad calcaneocuboid articulation; transversely broad, short, dorsoventrally compressed talus with shallow trochlea; relatively short, transversely broad, dorsoventrally compressed metatarsals IV and V., Published as part of Prideaux, Gavin J. & Warburton, Natalie M., 2023, A review of the late Cenozoic genus Bohra (Diprotodontia: Macropodidae) and the evolution of tree-kangaroos, pp. 1-95 in Zootaxa 5299 (1) on page 11, DOI: 10.11646/zootaxa.5299.1.1, http://zenodo.org/record/8012666, {"references":["Flannery, T. F. & Szalay, F. S. (1982) Bohra paulae: a new giant fossil tree kangaroo (Marsupialia: Macropodidae) from New South Wales, Australia. Australian Mammalogy, 5 (2), 83 - 94.","Muller, S. (1840) Bijdragen tot de Kennis van Nieuw-Guinea. In: Temminck, C. J. (Ed.), Verhandlungen over de Natuurlijke Geschiedenis der Nederlandische Overzeesche Bezittingen, Door de Leden der Natuurkundige Commissie in Indie en Andere Schrijvers. Zoologie. S. and J. Luchtmans and C. C. van der Hoek, Leiden, pp. 1 - 80."]}

Bohra sp. indet. 2 Referred specimen. UCMP 60850 a, right talus. SAM Hill (= Lake Kanunka Site 2, UCMP V5773), Lake Kanunka, eastern Lake Eyre Basin, South Australia (Figure 1). This specimen was collected by Richard H. Tedford et al. on 16 August 1961. Description and comparisons. In overall morphology, the talus is dorsoventrally compressed (especially in cranial view) with low trochlear crests, subequal in height and very shallow trochlea groove (Figure 35). The malleolar fossa is large, longer than wide, and moderately deep. The medial malleolus is broad and flattened. The neck of talus is intermediate in length, and extends cranially rather than craniomedially, such that lateral margin of navicular facet overlaps with medial trochlear crest from cranial view. The navicular facet of talar head is obliquely aligned relative to the craniocaudal plane, and relatively narrow and elongate. The talocalcaneal facets are smoothly conjoined and relatively shallow. The posterior plantar process is blunt (short) and broad. UCMP 60850a is intermediate between the holotype tali of B. illuminata and B. nullarbora in size and morphology, differing from the former by having a wider malleolar fossa and from the latter by bearing confluent calcaneal facets. The trochlear groove is more shallow than, but not as wide as, in B. illuminata, B. nullarbora, D. bennettianus and D. lumholtzi, but similar to B. planei. The medial trochlear crest is lower than in B. paulae, and is more similar to B. planei. The wide malleolar fossa is similar to D. bennettianus. The medial malleolus is relatively less projecting than in B. nullarbora, Dendrolagus, Dorcopsulus, Setonix, Thylogale, but similar to B. paulae and B. planei. The neck of talus is longer than in other species of Bohra and Dendrolagus, and the navicular facet of the talar head is narrower and more elongate, and less obliquely orientated than in B. paulae and B. planei, and particular B. illuminata and B. nullarbora. Although slightly water worn and abraded, the tibiotalar ligament fossa on the medial face of UCMP 60850a appears to have been very shallow and not extended above the dorsal border of the malleolus. The form of this fossa is most similar to the condition observed in B. planei in its medial restriction, but UCMP 60850a differs by lacking a distinct dorsal margin. Remarks. UCMP 60850a has suffered some post-depositional abrasion due to the fluviatile nature of the deposit, but it is complete enough to confidently say that it belongs to a species of Bohra. Although its morphology is not consistent with the talus of any of the four species for which this element is known (B. paulae, B. illuminata, B. nullarbora, B. planei), it is too incomplete to put a species name on. Nevertheless, UCMP60850a represents a definitive record for the genus in the late Pliocene of inland Australia. This is significant, because specimens from the Tirari Formation of the Lake Eyre Basin previously identified as belonging to large tree-kangaroos can now be attributed to other taxa. A maxilla fragment from Stirton Quarry (AMNH 144684 = SIAM 70) referred to cf. Dendrolagus (Tedford et al. 1992) belongs to a sthenurine (GJP, pers. obs.), while metatarsals from the Toolapinna LF tentatively referred to Dendrolagus (Tedford et al. 1992) belong to a species of Protemnodon (A. B. Camens, pers. comm., 2019; NMW, pers. obs.). Phylogenetic analysis Analysis of the 42 species and 85 characters (Appendices 1–2) produced 168 equally most parsimonious trees with a length of 251 steps, consistency index of 0.56, and retention index of 0.83. The strict consensus tree (Figure 36) collapses multiple nodes, the majority of which are within the Dendrolagina. Bohra is problematic due to the large proportion of missing data for most species, whereas Dendrolagus is challenging due to the markedly uniform osteology of its species. Macropodinae is supported by four unique synapomorphies (Figure 36): I3 bears a distinct anterobuccal crest (character 15); i1 rests on the palate, bordered by I2–3, during cheek-tooth occlusion (character 38); the ischium and ilium are aligned in the same plane (character 77); the ulnar olecranon process is moderately proportioned (character 85). Macropodines are also united by the possession of either a slightly or markedly broadened I1 (character 14), and by shared possession of bilophodont molars that lack a urocrista, a condition convergently characteristic of the sthenurine Hadronomas puckridgi Kear, 2002 (character 25). Macropodinae is supported by a bootstrap value of 72. Dorcopsoides fossilis Woodburne, 1967 is united with a Dorcopsis – Dorcopsulus – Watutia clade (Dorcopsini) as the basal macropodine branch in 64% of trees, where they are united by two unique synapomorphies: lateral constriction of P3 immediately anterior to the posterolingual cusp (character 21); a deep masseter muscle that extends to below the posterior cheek teeth, with the masseteric canal posteriorly confluent with the dental canal (character 35). In the strict consensus tree, D. fossilis forms a trichotomy with the dorcopsins and a clade giving rise to all other macropodines (Figure 36). Dorcopsis, Dorcopsulus and Watutia are united by two unique synapomorphies: the main crest of P3 is characterised by four cuspules with coarse ridgelets that extend to the crown base (character 17); the P3 crown is markedly deeper anteriorly than posteriorly (character 19). The clade giving rise to all other macropodines is supported by one unique synapomorphy (I1 markedly broadened; character 14) and four homoplastic features (Figure 36). Setonix brachyurus is reconstructed as the sister taxon to Dendrolagini + Macropodini in 82% of trees, but forms a trichotomy with those two tribes in the strict consensus tree. The monophyly of a clade composed of Wallabia Trouessart, 1904, Lagorchestes Gould, 1841, Prionotemnus Stirton, 1955, Kurrabi Flannery & Archer, 1984, Baringa Flannery & Hann, 1984, Onychogalea Gould, 1841, Osphranter and Macropus (Macropodini) is solidly supported (bootstrap value 74) by three unique podial synapomorphies: relatively short articular region for the talus on the calcaneus (character 53); rounded sustentaculum tali (character 60); medial malleolus on the talus characterised by a narrow, deep fossa and a small process (character 65) (Figure 36). Thylogale, Petrogale, Dendrolagus and Bohra (Dendrolagini) are united by two unique synapomorphies: triangular inflation of the squamosal between the occiput and zygomatic arch (character 11); relatively broad calcaneocuboid articulation (character 56). This clade is not supported in the bootstrap analysis. Dendrolagus and Bohra (Dendrolagina) are united by three unique synapomorphies (Figure 36), each a feature of the calcaneus. These are: posteromedial expansion of the tuber calcanei (character 51); relatively broad calcaneus–talus articulation (character 57); marked medial taper of the lateral talar facet (character 59). This alliance is reinforced by several homoplastic synapomorphies (Figure 36) and supported by a bootstrap value of 70. Bohra is recovered as paraphyletic in all 168 trees, and all species of Bohra collapse to form a comb with Dendrolagus in the bootstrap reconstruction due to having support values B. illuminata, B. nullarbora and B. paulae form a clade with Dendrolagus, united by three unique synapomorphies: smooth calcaneocuboid articulation (character 54); ventromedial facet of cuboid articulation on calcaneus reduced and confluent with dorsomedial facet (character 55); talus with medial malleolar fossa broad and moderately deep with large malleolar process (character 65). The species of Dendrolagus are united by four unique synapomorphies (Figure 36): a deep cleft on P3/p3 immediately posterior to the anterior cusp of the main crest (character 18); a relatively broad metatarsal V (character 69); a relatively long anterior crest on tibia (character 72); a short tibia relative to the femur (character 74). Two homoplastic traits may also be considered solid synapomorphies for the clade: a relatively shallow sustentaculum tali on the calcaneus otherwise only characterises the outgroup Hypsiprymnodon moschatus (character 62), while a short metatarsal IV relative to the length of the calcaneus is only seen elsewhere in Setonix brachyurus (character 66). Within Dendrolagus, the strict consensus tree produces a trichotomy between D. inustus, a D. bennettianus – D. lumholtzi clade and a clade composed of D. matschiei F̂rster & Rothschild, 1907, D. dorianus Ramsay, 1883, D. goodfellowi Thomas, 1908, D. scottae Flannery & Seri, 1990 and D. mbaiso. The New Guinea clade (excluding D. inustus) is united by two solid synapomorphies: relatively short distal fibular facet on the tibia (character 71); short pes (character 76). The latter is homoplastic, but only with the evolutionarily distant outgroup Hypsiprymnodon moschatus, and so is a very reliable synapomorphy. ......continued on the next page TABLE 9. (Continued) ......continued on the next page TABLE 9. (Continued) Body-size estimation Calcaneal measurements and body mass are highly positively correlated (p r 2 = 0.31) within tree-kangaroos and is not significant (p = 0.06), although it is significant within the larger macropodine sample (r 2 = 0.91). Ratio estimators range from 0.9649 to 1.4487 (Table 10). Corrected body-mass predictions vary depending on the dataset used to generate them, with those estimated from the modern tree-kangaroo dataset larger than those estimated from the macropodine dataset (Table 11). Body-mass estimates predicted from measurements where r 2 ≥ 0.70 fall within the following ranges: 25.7–43.5 kg for Bohra paulae; 25.6–47.4 kg for B. wilkinsonorum; 16.0– 26.9 kg for B. nullarbora; 15.9–26.7 kg for B. planei; and 18.7–29.9 kg for B. sp. indet. 1 (Table 11). No body-mass estimates were generated for B. illuminata, because the holotype is a young adult with unfused epiphyses, including that of the calcaneus., Published as part of Prideaux, Gavin J. & Warburton, Natalie M., 2023, A review of the late Cenozoic genus Bohra (Diprotodontia: Macropodidae) and the evolution of tree-kangaroos, pp. 1-95 in Zootaxa 5299 (1) on pages 67-74, DOI: 10.11646/zootaxa.5299.1.1, http://zenodo.org/record/8012666, {"references":["Tedford, R. H., Wells, R. T. & Barghoorn, S. F. (1992) Tirari Formation and contained faunas, Pliocene of the Lake Eyre Basin, South Australia. The Beagle, Records of the Northern Territory Museum of Arts and Sciences, 9 (1), 173 - 194. https: // doi. org / 10.5962 / p. 263124","Kear, B. P. (2002) Phylogenetic implications of macropodid (Marsupialia: Macropodoidea) postcranial remains from Miocene deposits of Riversleigh, northwestern Queensland. Alcheringa, 26 (2), 299 - 318. https: // doi. org / 10.1080 / 03115510208619259","Woodburne, M. O. (1967) The Alcoota fauna, central Australia. An integrated palaeontological and geological study. Bulletin of the Bureau of Mineral Resources, Geology and Geophysics, Australia, 87, 1 - 187.","Trouessart, E. L. (1904) Catalogus Mammalium tam Viventium quam Fossilium. Quinquennale Supplementum, Anno 1904. R. Friedlander and Son, Berlin, 288 pp. https: // doi. org / 10.5962 / bhl. title. 61820","Stirton, R. A. (1955) Late Tertiary marsupials from South Australia. Records of the South Australian Museum, 11, 247 - 268.","Flannery T. F. & Archer, M. (1984) The macropodoids (Marsupialia) of the early Pliocene Bow Local Fauna, central eastern New South Wales. The Australian Zoologist, 21 (4 - 5), 357 - 383.","Flannery, T. F. & Hann, L. (1984) A new macropodine genus and species (Marsupialia: Macropodidae) from the early Pleistocene of southwestern Victoria. Australian Mammalogy, 7 (4), 193 - 204.","F ˆ rster, F. & Rothschild, W. L. (1907) Description of a new tree kangaroo. Novitates Zoologicae, 14, 506. https: // doi. org / 10.5962 / bhl. part. 25383","Ramsay, E. P. (1883) Contributions to the zoology of New Guinea (part VII). Proceedings of the Linnean Society of New South Wales, 8 (1), 15 - 29. https: // doi. org / 10.5962 / bhl. part. 28637","Thomas, M. O. (1908) A new tree-kangaroo from British New Guinea. Annals and Magazine of Natural History, Series 8, 2 (11), 452 - 453. https: // doi. org / 10.1080 / 00222930808692512","Flannery, T. F. & Seri, L. (1990) Dendrolagus scottae n. sp. (Marsupialia: Macropodidae): a new tree-kangaroo from Papua New Guinea. Records of the Australian Museum, 42 (3), 237 - 245. https: // doi. org / 10.3853 / j. 0067 - 1975.42.1990.117"]}

Bohra bila (Dawson 2004b) comb. nov. Wallabia indra (De Vis, 1895): Bartholomai (1976b), pp. 375, 378 (partim), plate 53: figures 1–3. Not W. indra (De Vis, 1895). Silvaroo bila Dawson, 2004b: Dawson (2006), p. 114. Piper & Herrmann (2006), p. 130. Louys & Price (2015), pp. 565, 567–568, figure 8M. Piper (2016), p. 238. Eldridge et al. (2019), p. 822. Holotype. QM F43280, partial left adult dentary (preserving p3, m1–4). Type locality. Chinchilla Rifle Range (site not specified), north bank of the Condamine River, Chinchilla, western Darling Downs, southeastern Queensland. Chinchilla Sand. Pliocene. Collected by Doris and Cecil Wilkinson. Paratypes. QM F43276, partial left maxilla (preserving M1–4). Chinchilla Rifle Range (site not specified). Collected by Doris and Cecil Wilkinson. A second paratype (QM F43292, left i1) from Chinchilla is deemed here to be referable to a species of Congruus. Referred specimens. Lion Hill, Chinchilla Rifle Range, Queensland. QM F58668, left m1; QM F58669, left juvenile dentary fragment (preserving m3). Collected by Doris and Cecil Wilkinson. Chinchilla Sand, Chinchilla Rifle Range (site not specified). QM F58667, left M2. Collected by Doris and Cecil Wilkinson. Chinchilla (presumably Chinchilla Sand). QM F4740, partial left adult maxilla (preserving M2–4). Collected by Kendall Broadbent in 1887. Probably Darling Downs, Queensland. QM F4749, partial left juvenile dentary (preserving dp3, m1 trigonid). No locality details are associated with this specimen. Etymology. In the Wiradjuri language of central New South Wales, bila means ‘river’, a reference to the surmised riparian habitat of this species. Revised diagnosis. Bohra bila is distinguished from all other species of Bohra by having the palatine extend anteriorly to a position mesial to the M2 protoloph. The upper molars are similar in size to those of B. bandharr and B. illuminata, but its M4 is slightly smaller. They are most similar in morphology to those of the slightly smaller B. planei. They differ by having the cusp C region of the stylar crest diverge from the postparacrista closer to the paracone apex, and by lacking any sign of the stylar crest on M3. The ventral margin of the dentary is very smoothly curved posteriorly in lateral profile, lacking the digastric eminence of B. illuminata and B. nullarbora. It differs from that of B. bandharr by being deeper and having a shorter postalveolar shelf, which bears a small mesial process. The lower molars are similar in size and morphology to those of B. illuminata, but they lack a distinct premetacristid and the lophids are thinner in buccal view. The p3 is large and robust. Description and comparisons. Maxilla and palatine. QM F4740 is a chunk of maxilla that retains M2–4 and the complete masseteric process, and has attached to it a thin strip of palatine (Figure 21D–E). QM F43276 retains M1–4, but the end of its masseteric process is abraded off. It also has a thin strip of palatine fused to it, which extends anteriorly to a position mesial to the M2 anterior root (Figure 21B). The masseteric process of QM F4740 is triangular in lateral view. Its tip aligns with the M3 interloph valley (Figure 21D), and its leading edge is slightly twisted anterolaterally. Although the tip of the masseteric process is missing in QM F43276, we can extrapolate that it would have been positioned adjacent to the M3 protoloph. In B. bila, the maxilla–palatine suture extends anteriorly to a position adjacent to the M2 anterior root, but in B. nullarbora, B. illuminata, B. planei and B. bandharr it stops and inflects lingually at or posterior to a position mesial to the anterior edge of M3. The position of the masseteric process in B. bila is consistent with that observed in adult specimens of B. nullarbora, B. planei and B. bandharr. The form of the masseteric process is a good match for that of B. illuminata, but in the holotype of that species it is instead positioned adjacent to the M2 metaloph. This may be an artefact of its younger ontogenetic age (M4 is in the process of rotating into occlusion). Upper dentition. Previously described in detail for QM F43276 (Dawson 2004b). The referred unworn M2, QM F58667 (Figure 21F–J), is indistinguishable from the M2 of QM F 43276 in size and morphology, except for possessing a very slightly weaker cusp C region of the stylar crest. Bohra bila is a close match for B. bandharr and B. illuminata in upper molar size, but it has a slightly smaller M4 (Tables 1, 4–5). In morphology, including general shape of the crown outline, B. bila is most similar to B. planei, and to a slightly lesser degree, B. nullarbora, both of which have smaller molars. In contrast to that of B. planei, the cusp C region of the stylar crest diverge from the postparacrista closer to the paracone apex in B. bila rather than further up the paracone. In addition, there is no manifestation of this stylar crest region on M 3 in B. bila. The upper molars of B. bila differ from those of B. nullarbora by having a cusp C region of the stylar crest on M1–2, incurved postpara- and premeta-cristae, and a broader, more open posterior metaloph face on M4. Dentary. Previously described in detail for the holotype, QM F43280 (Dawson 2004b). The two referred juvenile dentary fragments are too incomplete to add to the knowledge of dentary morphology, but QM F58669 does confirm the absence of a digastric eminence. The holotype dentary of B. bila (Figure 22A–C) is most similar in overall form to that of B. nullarbora, but its ventral margin has a much more evenly curved profile due to the barely discernible digastric sulcus and absence of a digastric eminence (Figure 22A–B). It also has a larger and more rugose anterior insertion area for the internal superficial masseter muscle, and a more distinct buccinator sulcus (Figure 22A).The latter feature is also characteristic of the smaller B. bandharr, which is distinguished by its longer, rounded postalveolar shelf. A postalveolar shelf that bears a mesial process is shared with B. nullarbora, but the process on the latter is more pronounced. Lower dentition. The lower cheek teeth of the holotype have been described (Dawson 2004b), but the molars are heavily worn. Here we describe the teeth of three referred specimens, which have sustained little to no wear, one of which (QM F4749) preserves the previously unknown dp3. The dp3 is narrow, especially anteriorly due to the compressed form of the trigonid (Figure 22D). The protolophid is very narrow and inflects at the protoconid into a paracristid that rises to a point anteriorly that matches the protoconid in height. From there a crest descends lingually to the anterolingual corner of the crown. If this is a continuation of the paracristid, then its terminus is likely the position of the paraconid; however, it is also conceivable that the high point on the paracristid marks the paraconid (Figure 22H, L). The buccal face of the crown is sheer; there is no distinct cingulum, although the crown bulges slightly near to the base. There is no parastylar crest. The metaconid is higher than the protoconid and gives rise to short, fine pre- and post-metacristids (Figure 22D). Although broader than protolophid, the hypolophid is quite narrow relative to the m1 protolophid, and sits substantially lower than it in buccal view. Like the protolophid, the hypolophid slopes slightly buccally, resulting in the hypoconid being noticeably lower than the metaconid. The cristid obliqua descends into the interlophid valley and smoothly ascends to the protoconid apex, such that there is no distinct postprotocristid (Figure 22D). The posterior face of the hypolophid is smooth and featureless. QM F58668 is a complete, isolated, unworn m1 crown (Figure 22F–G, J, M–N), whereas QM F4749 is missing the m1 talonid. The only slight difference is that the most posterior portion of the paracristid is slightly inflected in QM F58668 and straight in QM F4749. The m1 protolophid is narrower than the hypolophid, and its crest is straight for much of its length, as opposed to posteriorly convex like the hypolophid. The paracristid trends anterolingually from the protoconid apex before turning lingually, forming a distinct anterior shelf with a rounded lingual border (Figure 22F, M). The small precingulid is at a distinctly lower level than the anterior portion of the paracristid (Figure 22M). A short, distinct parametacristid is directed anterobuccally from the metaconid apex, but terminates high on the protolophid anterior face. The cristid obliqua terminates low on the protolophid posterior face, buccal to the tooth midline. There is no distinct premetacristid, but the anterior face of the hypolophid is broadly convex on the lingual side. The posterior face of the hypolophid is smooth and featureless, except for a slight bulge low on the crown on the buccal side (Figure 22F, N). The m3 is generally similar in form to the much smaller m1, but differs in the following ways: the lophid sides are more convex, the protolophid is not as narrow relative to the hypolophid, the precingulid is broader, and the parametacristid is very low and barely discernible (Figure 22E–F). The dp3 is not known for any other species of Bohra. It is closest in overall morphology to that of the much smaller Setonix brachyurus (Quoy & Gaimard, 1830), with these species sharing a very similar trigonid topology. The main difference is that S. brachyurus lacks the crest that descends lingually to the anterolingual corner of the crown from the anterior high point on the paracristid. In addition, the base of the crown is not as narrow anteriorly relative to posteriorly in S. brachyurus, and the hypolophid curves more smoothly into the cristid obliqua and premetacristid than in B. bila. In species of Dendrolagus and dorcopsins, there is no distinct protolophid and paracristid, but rather a single crescentic crest (paralophid), which is met halfway along its length by the cristid obliqua. Species of Dendrolagus (e.g., D. bennettianus) may also have a distinct parastylar crest, but this is absent from B. bila. The p3 of B. bila is large and robust relative to the molars compared with its proportions in B. illuminata and B. sp. cf. B. bandharr, which are the only other species of Bohra for which p3 is known. The lower molars of B. bila are similar in size and morphology to those of B. illuminata (Tables 4–5), but the lophids are thinner in buccal view and the m3 lacks a distinct premetacristid. Remarks. Bohra bila is known definitively only from the Pliocene Chinchilla LF of southeastern Queensland (Figure 1). The referred specimen QM F4749 is from the Darling Downs, and is probably from Chinchilla, judging from the preservation of the bone, teeth and adhering sediment. This species is one of three in the Chinchilla LF, which is the most of any known assemblage. By recognising that Silvaroo bila Dawson, 2004b belongs in Bohra Flannery & Szalay, 1982, we render Silvaroo Dawson, 2004b a junior synonym of Bohra, because S. bila is its type species (Dawson 2004b). Furthermore, we recognise that the species originally named Protemnodon bandharr Dawson, Muirhead & Wroe, 1999, before it was shifted to Silvaroo by Dawson (2004b), also belongs in Bohra. Silvaroo sp. indet. 1 and Silvaroo sp. indet. 2 of Dawson (2004b) include specimens referable to B. bandharr and B. planei, respectively. The one remaining species that was allocated to Silvaroo is Protemnodon buloloensis Plane, 1967 from the Pliocene Awe LF of Papua New Guinea. Because it lacks the generic attributes of Protemnodon Owen, 1874 (Dawson 2004b; Kerr & Prideaux 2022), and because Silvaroo is a junior synonym of Bohra, the generic allocation of this species is uncertain. Although resolving this problem is outside of the scope of this work, it is worth noting that the form of the p3 is dorcopsin-like: it is markedly elongate relative to the molars and broader anteriorly before constricting just forward of the posterior end of the tooth. It is within the size range of Watutia novaeguineae, which is likewise from the Awe LF (Flannery et al. 1989), but it is clearly distinct from it, minimally at the species level., Published as part of Prideaux, Gavin J. & Warburton, Natalie M., 2023, A review of the late Cenozoic genus Bohra (Diprotodontia: Macropodidae) and the evolution of tree-kangaroos, pp. 1-95 in Zootaxa 5299 (1) on pages 38-42, DOI: 10.11646/zootaxa.5299.1.1, http://zenodo.org/record/8012666, {"references":["Dawson L. (2004 b) A new fossil genus of forest wallaby (Marsupialia, Macropodinae) and a review of Protemnodon from eastern Australia and New Guinea. Alcheringa, 28 (1), 275 - 290. https: // doi. org / 10.1080 / 03115510408619285","De Vis, C. W. (1895) A review of the fossil jaws of the Macropodidae in the Queensland Museum. Proceedings of the Linnean Society of New South Wales, Second Series, 10, 75 - 133.","Bartholomai, A. (1976 b) The genus Wallabia Trouessart (Marsupialia: Macropodidae) in the Upper Cainozoic deposits of Queensland. Memoirs of the Queensland Museum, 17 (3), 373 - 377.","Dawson, L. (2006) An ecophysiological approach to the extinction of large marsupial herbivores in middle and late Pleistocene Australia. Alcheringa, 30 (Supplement 1), 89 - 114. https: // doi. org / 10.1080 / 03115510609506857","Piper, K. J. & Herrmann, N. (2006) Additions to knowledge of the early Pleistocene wallaby, Baringa nelsonensis Flannery and Hann 1984 (Marsupialia: Macropodinae). Proceedings of the Linnean Society of New South Wales, 127, 125 - 131.","Louys, J. & Price, G. J. (2015) The Chinchilla Local Fauna: an exceptionally rich and well-preserved Pliocene vertebrate assemblage from fluviatile deposits of south-eastern Queensland, Australia. Acta Palaeontologica Polonica, 60 (3), 551 - 572. https: // doi. org / 10.4202 / app. 00042.2013","Piper, K. J. (2016) The Macropodidae (Marsupialia) of the early Pleistocene Nelson Bay Local Fauna, Victoria, Australia. Memoirs of Museum Victoria, 74, 233 - 253. https: // doi. org / 10.24199 / j. mmv. 2016.74.18","Eldridge, M. D., Beck, R. M. D., Croft, D. A., Travouillon, K. J. & Fox, B. J. (2019) An emerging consensus in the evolution, phylogeny, and systematics of marsupials and their fossil relatives (Metatheria). Journal of Mammalogy, 100 (3), 802 - 837. https: // doi. org / 10.1093 / jmammal / gyz 018","Quoy, J. R. C. & Gaimard, P. (1830) Voyage de Decouvertes de l'Astrolabe Execute par Ordre du Roi, Pendant les Annees 1826 - 1827 - 1828 - 1829, sous la Commandement de M. J. Dumond d'Urville. Zoologie. Vol. 1. [1 - 4]. J. Tastu, Paris, 268 pp.","Flannery, T. F. & Szalay, F. S. (1982) Bohra paulae: a new giant fossil tree kangaroo (Marsupialia: Macropodidae) from New South Wales, Australia. Australian Mammalogy, 5 (2), 83 - 94.","Dawson, L., Muirhead, J. & Wroe, S. (1999) The Big Sink Local Fauna: a lower Pliocene mammalian fauna from the Wellington Caves complex, Wellington, New South Wales. Records of the Western Australian Museum, Supplement No. 57, 265 - 290.","Plane, M. D. (1967) Stratigraphy and vertebrate fauna of the Otibanda Formation, New Guinea. Bulletin. Bureau of Mineral Resources, Geology and Geophysics, Australia, 86, 1 - 64.","Owen, R. (1874) On the fossil mammals of Australia, Part VIII. Family Macropodidae: genera Macropus, Osphranter, Phascolagus, Sthenurus and Protemnodon. Philosophical Transactions of the Royal Society, 164, 245 - 288. https: // doi. org / 10.1098 / rstl. 1874.0008","Prideaux, G. J., Kerr, I. A. R., van Zoelen, J. D., Grun, R., van der Kaars, S., Oertle, A., Douka, K., Grono, E., Barron, A., Mountain, M. J. & Westaway, M. C. (2022) Re-evaluating the evidence for late-surviving megafauna at Nombe rockshelter in the New Guinea highlands. Archaeology in Oceania, 57 (3), 223 - 248. https: // doi. org / 10.1002 / arco. 5274","Flannery, T. F., Hoch, E. & Aplin, K. P. (1989) Macropodines from the Pliocene Otibanda Formation, Papua New Guinea. Alcheringa, 13 (2), 145 - 152. https: // doi. org / 10.1080 / 03115518908619048"]}

Bohra sp. cf. B. bandharr cf. Dendrolagus sp. 1: Flannery & Archer (1984), pp. 358–359, 361–362, figure 1F–G, table 1. Rich (1991), p. 1049. cf. Dendrolagus sp. 2: Flannery & Archer (1984), pp. 358–359, 362–363, figure 1F–G, table 1. Rich (1991), p. 1049. Tedford et al. (1992), p. 187. cf. Dendrolagus: Tedford et al. (1992), p. 190, table 3; partim. Protemnodon ?cf. bandharr: Dawson et al. (1999), pp. 281, 287, table 10. Dendrolagus sp. 2: Dawson et al. (1999), p. 281. Dendrolagus sp.: Dawson (2004a), pp. 267, 270–272, table 1; partim. Silvaroo sp. indet. 1: Dawson (2004b), pp. 283, 286, figure 3. Silvaroo sp. 1: Dawson (2004b), p. 288, table 2. Dendrolagus sp. indet.: Prideaux & Warburton (2008), pp. 463–464, figure 1; partim. Silvaroo sp.: Louys & Price (2015), pp. 565, 567, 569, figure 8N. Referred specimens. Chinchilla Rifle Range (site not specified), Queensland. QM F43281, partial right adult dentary (preserving m3–4). QM F58665, partial right adult dentary (preserving m3–4); QM F61040, right i1. Collected by Doris and Cecil Wilkinson. Probably Chinchilla. QM F4750, partial right adult dentary (preserving incomplete p3, m1–4). This specimen retains no associated locality information. Chinchilla Sand is suggested by preservation (chocolate-brown bone, blue-grey enamel and yellow- to orange- to chocolate-brown adhering matrix), which has been used to retrospectively, albeit tentatively, differentiate Chinchilla Sand specimens from other Darling Downs material (e.g., Bartholomai 1963, 1975; Bartholomai & Woods 1976; Louys & Price 2015). Collection details are unknown, but it was very likely collected in the late 19 th century. The specimen was ascribed the number 8808 in De Vis’s catalogue. Bow, New South Wales. AM F59576, partial right adult maxilla (preserving M2–3); AM F64004, posterior half of left P3; AM F64007, right M4; AM F64075, partial left maxilla (preserving very worn M2–4). AM F64245, left M4. Fisherman’s Cliff, Warrananga Station, New South Wales. NMV P38267, left M2; MNV P38351, left M3 metaloph; NMV P38325, right p3. Collected by Edmund Gill and others in the late 1960s or James Warren and others in the early 1970s. FUPS Quarry, Bone Gulch, Moorna Station, New South Wales. NMV P257273, partial left P3; NMV P257274, left M3; NMV P257275, right M1 metaloph. These specimens were picked from concentrate sieved from sediment collected by a team led by GJP. NMV P257273 was collected on 14 February 2016, NMV P257274 on 16 August 2015, and NMV P257275 on 15 February 2016. Remarks. An assortment of specimens have been grouped under this limbo nomen (Figure 16–18) because they share dentary or cheek-tooth traits with specimens of B. bandharr (e.g., comparatively rectangular upper molars, dentary with a long, rounded postalveolar shelf), but differ by having narrower, much smaller molars (Table 2). NMV P38325, by contrast, is a p3 from the early Pleistocene Fisherman’s Cliff LF (Figure 1) referred to B. sp. cf. B. bandharr by association with several loose molars, and the general similarity of its form (Figure 18J–L) to that of the only other unworn p3 known for a species of Bohra, that of the paratype of B. illuminata (Figure 24D). The barrier to defining a species from among this collective is the lack, in any one specimen, of sufficient diagnostic attributes that would permit its selection as a reliable holotype. Obtaining further tree-kangaroo fossils from the Chinchilla and Bow localities (Figure 1) may offer the greatest chance of solving this impasse. Nevertheless, although it lacks a name, it would be reasonable to assume that within the sample referred to B. sp. cf. B. bandharr there is yet another late Cenozoic species of Bohra., Published as part of Prideaux, Gavin J. & Warburton, Natalie M., 2023, A review of the late Cenozoic genus Bohra (Diprotodontia: Macropodidae) and the evolution of tree-kangaroos, pp. 1-95 in Zootaxa 5299 (1) on page 31, DOI: 10.11646/zootaxa.5299.1.1, http://zenodo.org/record/8012666, {"references":["Flannery T. F. & Archer, M. (1984) The macropodoids (Marsupialia) of the early Pliocene Bow Local Fauna, central eastern New South Wales. The Australian Zoologist, 21 (4 - 5), 357 - 383.","Rich, T. H., Archer, M., Hand, S. J., Godthelp, H., Muirhead, J., Pledge, N. S., Flannery, T. F., Woodburne, M. O., Case, J. A., Tedford, R. H. & Turnbull, W. D., Lundelius, E. L., Rich, L. S. V., Whitelaw, M. J., Kemp, A. & Rich, P. V. (1991) Australian Mesozoic and Tertiary terrestrial mammal localities. In: Vickers-Rich, P., Monaghan, J. M., Baird, R. F. & Rich, T. H. (Eds.), Vertebrate Palaeontology of Australasia. Pioneer Design Studio and Monash University Publications Committee, Melbourne, pp. 1005 - 1070.","Tedford, R. H., Wells, R. T. & Barghoorn, S. F. (1992) Tirari Formation and contained faunas, Pliocene of the Lake Eyre Basin, South Australia. The Beagle, Records of the Northern Territory Museum of Arts and Sciences, 9 (1), 173 - 194. https: // doi. org / 10.5962 / p. 263124","Dawson, L., Muirhead, J. & Wroe, S. (1999) The Big Sink Local Fauna: a lower Pliocene mammalian fauna from the Wellington Caves complex, Wellington, New South Wales. Records of the Western Australian Museum, Supplement No. 57, 265 - 290.","Dawson, L. (2004 a) A new Pliocene tree kangaroo species (Marsupialia, Macropodinae) from the Chinchilla Local Fauna, southeastern Queensland. Alcheringa, 28 (1), 267 - 273. https: // doi. org / 10.1080 / 03115510408619284","Dawson L. (2004 b) A new fossil genus of forest wallaby (Marsupialia, Macropodinae) and a review of Protemnodon from eastern Australia and New Guinea. Alcheringa, 28 (1), 275 - 290. https: // doi. org / 10.1080 / 03115510408619285","Prideaux, G. J. & Warburton, N. M. (2008) A new Pleistocene tree-kangaroo (Diprotodontia: Macropodidae) from the Nullarbor Plain of south-central Australia. Journal of Vertebrate Paleontology, 28 (2), 463 - 478. https: // doi. org / 10.1671 / 0272 - 4634 (2008) 28 [463: ANPTDM] 2.0. CO; 2","Louys, J. & Price, G. J. (2015) The Chinchilla Local Fauna: an exceptionally rich and well-preserved Pliocene vertebrate assemblage from fluviatile deposits of south-eastern Queensland, Australia. Acta Palaeontologica Polonica, 60 (3), 551 - 572. https: // doi. org / 10.4202 / app. 00042.2013","Bartholomai A. (1963) Revision of the extinct macropodid genus Sthenurus Owen in Queensland. Memoirs of the Queensland Museum, 14 (3), 51 - 76.","Bartholomai, A. (1975) The genus Macropus Shaw (Marsupialia: Macropodidae) in the Upper Cainozoic deposits of Queensland. Memoirs of the Queensland Museum, 17 (2), 195 - 236."]}

Bohra nullarbora Prideaux & Warburton, 2009 Bohra sp. nov. 2: Prideaux (2006), p. 1524. Prideaux et al. (2007), p. 423, table 1. Bohra sp. nov.: Warburton & Prideaux (2010), pp. 138–139, 143, 147, 149, figures 13.1, 13.5–13.6. Holotype. WAM 05.4.70, partial adult skeleton. Craniodental elements: fragmentary cranium (preserving left P3, left and right M1–4); partial left dentary (preserving base of i1, m1–3); right dentary preserving posterior half of m2, m3–4). Postcranial elements: vertebrae including atlas, axis, cervical 3–5, thoracic 1,?5–6,?10,?12, dorsal lumbar fragment, two mid-caudal and two distal-caudal elements; one mid-caudal chevron; left and right clavicles; six ribs; proximal portion of left humerus and distal fragments; partial diaphysis of right humerus; left ulna; left radius diaphysis fragment; right hamatum; left scaphoid; left metacarpal III; right metacarpal III; left metacarpals IV – V; manual phalanges, including those of digits I– IV proximal, digits III– V medial, digits III– IV distal; partial left innominates; fragments of right innominate; left epipubic; proximal portion of left femur; abraded distal fragments of right femur; fragments of left distal tibia diaphysis; right tibia diaphysis; distal portion of right fibula; left calcaneus; left talus; left navicular; right entocuneiform; right metatarsals IV – V, pedal phalanges including those of digits II, IV – V proximal, digits II – V medial, digits IV – V distal. TABLE 6 (continued). Type Locality. Leaena’s Breath Cave, Nullarbor Plain, southeastern Western Australia. The holotype was collected by Paul D. Devine and GJP in July 2002 from a tight space beneath an overhanging boulder atop the main boulder pile, adjacent to the ‘gypsum powder floor’. Fossils from the Leaena’s Breath Cave boulder floor, although previously inferred to be of early middle Pleistocene age (Prideaux et al. 2007), are actually undated, but are associated with remains of other species known from dated middle and late Pleistocene sites elsewhere. At this point the type locality age is best considered conservatively as Pleistocene. The precise location from which the holotype was collected is recorded with the Department of Earth and Planetary Sciences, Western Australian Museum. Referred specimens. Leaena’s Breath Cave (stratigraphic unit 3), Nullarbor Plain, southeastern Western Australia. WAM 2020.3.201, right cuboid. RF 95 fissure fill, Curramulka Quarry, Yorke Peninsula, South Australia. SAMA P55204, left M2; SAMA P55205, left M2; SAMA P55206, right M2; SAM P55389, right M1; SAMA P55390, right M1; SAMA P55391, right M1; SAMA P55392, right M1; SAMA P55393, left M1; SAMA P55394, left M1; SAMA P55395, left M1; SAMA P55396, left M1; SAMA P55397, right M1; SAMA P55398, left M2, right M2 (these teeth are identical in morphology, wear and preservation, so are considered to belong to the same individual); SAMA P55400, right M3; SAMA P55407, right M1; SAMA P55411, left m2; SAMA P55412, left m2; SAMA P55413, left dP3; SAMA P55417, left m1; SAMA P55419, left m2 trigonid. These specimens were picked from concentrate sieved from sediment collected in 1997 by Jim McNamara (South Australian Museum). Revised diagnosis. Bohra nullarbora is distinguished from B. illuminata in the following cranial features: a shallow buccinator fossa; a supraorbital crest that is more markedly projected laterally above the posterior end of the eye orbit; a less-domed dorsal neurocranial surface; a wider ectoglenoid process; a more distally narrowed postglenoid process. The cheek teeth are smaller relative to the dimensions of the palate than in B. illuminata. P3 is wider anteriorly than posteriorly, but narrow overall relative to M1, and it bears a small, low posterolingual cusp and narrow lingual cingulum. The dentary is characterised by a well-developed digastric eminence and sulcus, and a distinctly hooked mesial process on the postalveolar shelf. The dorsal and ventral enamel flanges on i1 are thinner than in B. illuminata. The lower molars are distinguished from those of other species of Bohra by having a short anterior portion of the trigonid, and a very low paracristid and cristid obliqua. The m4 has a very low parametacristid, which extends to centre of trigonid basin. Bohra nullarbora is distinguished from B. illuminata in the following postcranial features. The clavicle is more robust and less flattened, with expanded articular facets and a more laterally positioned cranial inflection. The humerus is more robust with a stronger pectoral crest, deltoid ridge, bicipital groove and spinodeltoid insertion, but with a less pronounced teres major tubercle. The distal articular surface (especially the capitulum) of the humerus is relatively narrow. The ilium is relatively broader anteroposteriorly, and with a more sinuous medial border.A marked oval fossa is contiguous with a rugose muscle scar of the m. rectus femoris dorsal to the acetabulum. The epipubic is very large. The calcaneus is larger with partial separation of posterior talar facets, a very short anteroplantar sulcus, a more strongly developed plantar tuberosity, a wider sustentaculum tali, and a relatively narrow, less mesially tapered lateral talar facet. The talus is relatively longer with a higher lateral trochlear ridge. Metatarsal IV is more robust with a larger cuboid facet, a less medially constricted facet for metatarsal V, and a less rounded, less distally orientated sesamoid facet. Metatarsal V is more robust. Bohra nullarbora is distinguished from B. paulae in the following pedal features: smaller absolute size (Table 8); separation of the talar facets, less complete assimilation of the ventromedial cuboid facet with the dorsal cuboid facets, and very short anteroplantar sulcus on the calcaneus; higher trochlear crests (deeper trochlear groove) and a longer neck on the talus; more gracile metatarsals IV and V. Bohra nullarbora is distinguished from B. planei by having a less laterally flared base of the greater trochanter, distinct posterior talar facets and more smoother cuboid step on the calcaneus, and higher trochlear crests on the talus, and from B. wilkinsonorum in its much smaller overall size and more complete assimilation of the ventromedial cuboid facet on the calcaneus. Etymology. The species name refers to the Nullarbor Plain, the region whence the holotype was collected. Description and comparisons. Cranium. Among the species of Bohra, only B. nullarbora (Figure 27) and B. illuminata (Figure 23) are known from parts of the cranium other than the tooth-bearing portion of the maxilla. The incisor-bearing portion of the premaxilla is slightly deeper in B. nullarbora than in B. illuminata. The buccinator fossa is shallower and less distinct than in B. illuminata. The supraorbital crest is markedly projected laterally above the posterior end of the eye orbit and underlain by a deep sulcus, whereas the supraorbital crest is more of a triangular swelling in B. illuminata. The zygomatic arch is relatively deeper in B. nullarbora than in B. illuminata, and the dorsal surface of the neurocranium is less domed. The ectoglenoid process is wider and the postglenoid process is more narrowed distally (ventrally). Upper dentition. The upper cheek teeth (Figures 27E, 28A–O) are similar in size to those of B. planei and B. sp. cf. B. bandharr, but smaller than in B. illuminata, B. bandharr and B. bila (Tables 1–7). Greater width of the P3 anteriorly compared with posteriorly (Figure 27E) is a condition unique to B. nullarbora. P3 is narrower relative to M1 than in any other species of Bohra. As in B. illuminata, there is no manifestation of a posterobuccal eminence on P3, whereas it is slight in B. bandharr and distinct in B. wilkinsonorum. The upper molars are most similar in morphology to those of B. illuminata, but differ by being slightly smaller overall and shorter relative to their width, and by having a more concave lingual margin of the crown adjacent to the interloph valley and a more protuberant lingual metaloph base. Absence of a stylar crest in the region of cusp C is a condition shared with B. bandharr, B. illuminata and B. wilkinsonorum, but not B. bila and B. planei, in which this crest is distinct. Dentary. The well-developed digastric eminence and sulcus (Figure 29A–C), and distinctly hooked mesial process on the postalveolar shelf (Figure 29G) distinguish B. nullarbora from other species of Bohra. Although it is broken off, it is evident that the mandibular condyle was positioned at a higher level relative to the cheek teeth than in B. illuminata. The anterior insertion area for the internal superficial masseter muscle (Figure 29A) is larger than in B. illuminata, similar in proportions to B. sp. cf. B. bandharr (QM F4750), but smaller than in B. bandharr and B. bila. Lower dentition. The i1 of B. nullarbora is similar in size to that of B. illuminata, but it has thinner dorsal and ventral enamel flanges. In overall size and morphology, the lower molars are most similar to those of B. sp. cf. B. bandharr (e.g., QM F4750, QM F58665), but they have a shorter anterior portion of the trigonid (precingulid + anterior portion of paracristid) than in any other species of Bohra, and the paracristid and cristid obliqua are also comparatively very low. The species is also distinguished by the presence, on m4, of a very low parametacristid, which extends to the centre of the trigonid basin, terminating against the paracristid (Figure 29F). Clavicle. The clavicle is more robust and less flattened than in B. illuminata, with expanded articular facets and less-pronounced cranial curvature. Humerus. The humerus is more robust than in B. illuminata, with a stronger pectoral crest, deltoid ridge, bicipital groove and spino-deltoid insertion. The teres major insertion is less protruding. The distal articular surface, in particular the capitulum, is relatively narrower than in B. illuminata. Innominate. The base of the ilium is relatively broader in the craniocaudal axis, and the medial border is sinuous by comparison with the straight medial border in B. illuminata. An oval fossa contiguous with the rugose muscle scar of m. rectus femoris dorsal to the acetabulum is distinct and unique to B. nullarbora. Femur. The femur of B. nullarbora is distinguished from that of B. illuminata in its more caudally positioned scars for the mm. adductores femoris, and from B. planei by having a less laterally projected base of the greater trochanter. Calcaneus. Morphologically, the calcaneus of B. nullarbora is distinguished from those of B. illuminata, B. paulae and B. planei by having distinctly separate medial and lateral talar facets, and from B. wilkinsonorum and B. sp. indet. 1 by having a straight lateral edge to the medial facet, which extends both cranially and caudally from the facet margins. It shares with B. illuminata and B. paulae a less acute, more smoothed cuboid step than in B. planei, B. wilkinsonorum and B. sp. indet. 1. It shares with B. illuminata a ventromedial cuboid facet that is confluent with the dorsal facets, though the entire surface is not as smoothed as in B. paulae. The relatively very short anteroplantar sulcus between the rugose plantar surface and anterior plantar tubercle is unique to B. nullarbora. Talus. The talus of B. nullarbora is distinguished from all other species of Bohra and Dendrolagus (e.g., Figure 9F–K) in its relatively higher trochlear crests, and thus deeper trochlear groove, and in the distinct separation between the calcaneal facets. It also has a relatively deeper malleolar fossa, more projected malleolar tubercle, and longer neck than in other species of Bohra. Metatarsals. Metatarsal IV and, in particular, metatarsal V of B. nullarbora are more robust than in B. illuminata, though this might be accounted for by the ontogenetically younger age of the holotype of B. illuminata versus that of B. nullarbora. The facet for metatarsal V is relatively smaller and the gap between this and the sesamoid facet is relatively wider in B. nullarbora compared with the conditions of B. illuminata and B. paulae, though this may be within the normal range of intraspecific variation given their differing ontogenetic ages. Metatarsal V of B. nullarbora is more gracile than in B. paulae, with a relatively narrower facet for metatarsal IV, a more elongate proximolateral process and a more deeply flexed cuboid facet. Remarks. Our understanding of B. nullarbora is fundamentally as it was when it was named and first described in 2009, but the broader comparisons made here mean that the species is now better circumscribed. In addition, several loose molars referable to B. nullarbora have been identified in a Pleistocene assemblage from Yorke Peninsula, 1,000 km southeast of the Nullarbor type locality (Figure 1). Whether the ancient DNA retrieved from Tunnel Cave in southwestern Australia (Murray et al. 2013) belongs to B. nullarbora, B. illuminata or neither, remains to be verified with further molecular work., Published as part of Prideaux, Gavin J. & Warburton, Natalie M., 2023, A review of the late Cenozoic genus Bohra (Diprotodontia: Macropodidae) and the evolution of tree-kangaroos, pp. 1-95 in Zootaxa 5299 (1) on pages 49-59, DOI: 10.11646/zootaxa.5299.1.1, http://zenodo.org/record/8012666, {"references":["Prideaux, G. J. & Warburton, N. M. (2009) Bohra nullarbora sp. nov., a second tree-kangaroo (Marsupialia: Macropodidae) from the Pleistocene of the Nullarbor Plain, Western Australia. Records of the Western Australian Museum, 25 (2), 165 - 179. https: // doi. org / 10.18195 / issn. 0312 - 3162.25 (2). 2009.165 - 179","Prideaux, G. J. (2006) Mid-Pleistocene vertebrate records: Australia. In: Elias, S. A. (Ed.), Encyclopedia of Quaternary Science. Elsevier, Oxford, pp. 1517 - 1537. https: // doi. org / 10.1016 / B 0 - 44 - 452747 - 8 / 00258 - 1","Prideaux, G. J., Long, J. A., Ayliffe, L. K., Hellstrom, J. C., Pillans, B., Boles, W. E., Hutchinson, M. N., Roberts, R. G., Cupper, M. J., Arnold, L. J., Devine P. D. & Warburton N. M. (2007) An arid-adapted middle Pleistocene vertebrate fauna from south-central Australia. Nature, 445 (7126), 422 - 425. https: // doi. org / 10.1038 / nature 05471","Warburton, N. M. & Prideaux, G. J. (2010) Functional pedal morphology of the extinct tree-kangaroo Bohra (Diprotodontia: Macropodidae). In: Coulson, G. C. & Eldridge, M. D. B. (Eds.), Macropods: The Biology of Kangaroos, Wallabies and Rat-kangaroos. Surrey Beatty and Sons, Sydney, pp. 137 - 151.","Murray, D. C., Haile, J., Dortch, J., White, N. E., Haouchar, D., Bellgard, M. I., Allcock, R. J., Prideaux, G. J. and Bunce, M. (2013) Scrapheap challenge: a novel bulk-bone metabarcoding method to investigate ancient DNA in faunal assemblages. Scientific Reports, 3, 3371. https: // doi. org / 10.1038 / srep 03371"]}

Funder: McKenzie Fellowship Program, Funder: University of Melbourne, Lead-based ammunition (gunshot and bullets) frequently leaves small lead fragments embedded in the meat of wild-shot game animals. Australia produces several commercial game meat products from wild animals harvested with lead-based ammunition and has a growing population of recreational hunters. However, no studies have previously investigated the frequency of lead fragments or lead concentrations in Australian game meat. We examined 133 Australian minced game meat items of four types for evidence of lead contamination. Samples were meat from kangaroos (Macropus and Osphranter spp.; n=36) and Bennett's wallabies (Notamacropus rufogriseus; n=28) sold for human consumption, and deer ('venison'; multiple spp.; n=32) and stubble quail (Coturnix pectoralis; n=37) harvested for private consumption by recreational hunters. All packages were studied by digital radiography to detect the presence of radio-dense fragments, assumed to be lead fragments from ammunition. Visible fragments were absent in commercially available kangaroo products, but were present in 4%, 28% and 35% of wallaby, venison and quail, respectively. Mean meat lead concentrations (mg/kg wet weight) were 0.01 ± 0.01 for kangaroo, 0.02 ± 0.01 for wallaby, 0.12 ± 0.07 for venison, and 1.76 ± 3.76 for quail. The Australian food standards threshold for livestock meat (0.1 mg/kg w.w.) was not exceeded by any kangaroo or wallaby products but was exceeded by 53% and 86% of venison and quail, respectively. Radiography only detected 35% of samples that were above the food safety threshold. While average lead concentrations in commercially available macropod (kangaroo and wallaby) meat were low, those in recreationally harvested game meat may pose health risks for hunters and associated consumers.

An albino infant wallaby was born to a mother with the wild-type body color. PCR and sequencing analyses of TYR (encoding tyrosinase, which is essential for melanin biosynthesis) of this albino wallaby revealed a 7.1-kb-long DNA fragment inserted in the first exon. Because the fragment carried long terminal repeats, we assumed it to be a copy of an endogenous retrovirus, which we named walb. We cloned other walb copies residing in the genomes of this species and another wallaby species. The copies exhibited length variation, and the longest copy (>8.0 kb) contained open reading frames whose deduced amino acid sequences were well aligned with those of gag, pol, and env of retroviruses. It is not known through which of the following likely processes the walb copy was inserted into TYR: endogenization (infection of a germline cell by an exogenous virus), reinfection (infection by a virus produced from a previously endogenized provirus), or retrotransposition (intracellular relocation of a provirus). In any case, the insertion into TYR is considered to have been a recent event on an evolutionary timescale because albino mutant alleles generally do not persist for long because of their deleterious effects in wild circumstances., 白いワラビーの遺伝的原因を初めて解明 --世界各地の動物園で相次いで誕生--. 京都大学プレスリリース. 2022-01-18.