from the Early Miocene of Australia

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A new cracticid (Passeriformes : Cracticidae) from the Early Miocene of Australia Jacqueline M. T. Nguyen A,D, Trevor H. Worthy B, Walter E. Boles C, Suzanne J. Hand A and Michael Archer A A

School of Biological, Earth and Environmental Sciences, University of New South Wales, Sydney, NSW 2052, Australia. B School of Biological Sciences, Flinders University, Adelaide, SA 5001, Australia. C Research and Collections Branch, Ornithology Section, Australian Museum, 6 College Street, Sydney, NSW 2010, Australia. D Corresponding author. Email: [email protected]

Abstract. The Cracticidae (Passeriformes) is an endemic Australo-Papuan family that, for the purposes of this paper, comprises the butcherbirds and Australian Magpie (Cracticus), currawongs (Strepera) and peltops (Peltops). Here we describe a new genus and species of cracticid from an Early Miocene deposit in the Riversleigh World Heritage Area, northwestern Queensland, Australia. Kurrartapu johnnguyeni, gen. nov., sp. nov. is described from a proximal tarsometatarsus that is similar in size to that of the extant Black Butcherbird (C. quoyi). This new species shares morphological features with the Strepera–Cracticus clade to the exclusion of Peltops, which suggests that it is a representative of the crown-group Cracticidae. Kurrartapu johnnguyeni represents the first Tertiary record of the Cracticidae in Australia, and is in concordance with molecular estimates for the timing of the cracticid radiation. We also describe morphological differences of the tarsometatarsus between cracticids and woodswallows (Artamus), which have at times been considered confamilial. We add this new cracticid to the expanding Tertiary fossil record of passerines in Australia, which plays a significant role in our understanding of early passerine evolution. Additional keywords: Artamus, Cracticus, fossil bird, passerine, Riversleigh, songbird, Strepera, tarsometatarsus. Received 11 March 2013, accepted 5 June 2013, published online 18 September 2013

Introduction The Cracticidae (sensu Dickinson 2003) is a family of predatory passerines that are characteristic of the Australo-Papuan avifauna. As here construed, the family comprises the currawongs (Strepera, three species), peltops (also known as shieldbills) (Peltops, two species) and butcherbirds (Cracticus, eight species), including the Australian Magpie (C. tibicen), which was formerly segregated in the genus Gymnorhina (see Christidis and Boles 2008). Strepera is endemic to Australia and Peltops to New Guinea, whereas Cracticus is distributed throughout these two land masses. These birds range in size from medium (Peltops) to large (Australian Magpie and Strepera), and occupy a wide range of terrestrial habitats, from alpine forests to coastal shrublands. Most species of Cracticus and Strepera are commonly found in open sclerophyll woodlands and forests, although some species occur in denser habitats, such as the Black Butcherbird (C. quoyi), which inhabits rainforests and mangroves. The widespread Australian Magpie and Pied Currawong (Strepera graculina) are familiar birds of farmland and suburban areas. Peltops, however, is restricted to rainforests and swamp forests (Coates 1990; Higgins et al. 2006). Journal compilation Ó BirdLife Australia 2013

All cracticids are strong fliers and predominantly feed on insects and other invertebrates, but also eat fruit, small vertebrates and carrion. These birds have adapted to a range of foraging niches that enable them to occupy all vertical strata, from the ground to the upper canopy. The flycatcher-like peltops are aerial feeders, whereas the robust, long-legged Australian Magpie is a terrestrial forager. Butcherbirds are perch-and-pounce hunters that frequent the ground to mid-storey, whereas currawongs forage through all vertical strata (Debus 1996; Schodde and Mason 1999). Australasia is one of the centres of diversity for the core Corvoidea (sensu Barker et al. 2004), a large assemblage of oscine passerines that includes the crows (Corvidae), true shrikes (Laniidae), birds-of-paradise (Paradisaeidae), fantails (Rhipiduridae), whistlers and allies (Pachycephalidae), orioles (Oriolidae) and the Cracticidae, among others. Within the core Corvoidea, cracticids are currently understood to be part of the shrike-like passerine assemblage Malaconotoidea (sensu Cracraft et al. 2004; e.g. Jønsson et al. 2011; Hugall and Stuart-Fox 2012). This assemblage comprises the Australasian woodswallows (Artamus), African bush-shrikes (Malaconotiwww.publish.csiro.au/journals/emu

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dae), helmet-shrikes (Prionopidae), batises (Platysteiridae), vangas (Vangidae) and Asian ioras (Aegithinidae). Also included in this assemblage are the Australo-Papuan boatbills (Machaerirhynchus), Mottled Whistler (Rhagologus leucostigma) and Bornean Bristlehead (Pityriasis gymnocephala), the last of which was previously included with the cracticids (e.g. Ahlquist et al. 1984; see Moyle et al. 2006). A close affinity between cracticids and Artamus has been supported by morphological studies (Pycraft 1907; Beecher 1953; McEvey 1976; Manegold 2008a), protein electrophoresis (Christidis and Schodde 1991) and DNA–DNA hybridisation data (Sibley and Ahlquist 1990). Several analyses of DNA sequences recovered mixed support for a sister relationship between Artamus and cracticids but were also based on limited sampling of Artamus species, cracticids and other malaconotoid taxa (e.g. Moyle et al. 2006; Norman et al. 2009; Fuchs et al. 2012). Analyses that have incorporated more representatives of these groups and additional genes yielded weak support for a nested position of Artamus within the Cracticidae (Hugall and Stuart-Fox 2012; Jetz et al. 2012). A recent mitochondrial phylogeny of all cracticid species by Kearns et al. (2013, fig. 1) also did not find support for an sister relationship between Artamus and Cracticidae, but recovered moderate support for one between cracticids and the African bush-shrikes and allies. The uncertainty in the molecular estimates of these relationships is also reflected in the current taxonomic treatment of Artamus and cracticids. The assignment of family-group ranks to these taxa and their circumscription are still unresolved. Following Sibley and Ahlquist’s (1990) classification, some authors have combined these birds into a single family, Artamidae (e.g. Schodde and Mason 1999; Higgins et al. 2006; Christidis and Boles 2008), but others maintain the traditional separation of Artamus into its own family (e.g. Dickinson 2003; Johnstone and Storr 2004; Russell and Rowley 2009). The Australian fossil record of the Cracticidae was, until now, restricted to the Quaternary. The Pleistocene fossil material includes Cracticus tibicen (previously listed as Gymnorhina) from Naracoorte Caves World Heritage Area, South Australia (Moriarty et al. 2000; Reed and Bourne 2000, 2009) and Byaduk Caves, Victoria (Baird 1991). Fossils of C. tibicen, as well as Strepera graculina and S. versicolor, were also identified from Seton Rock Shelter, South Australia (Hope et al. 1977). Remains of S. versicolor were recovered from Devil’s Lair, Western Australia (Baird 1986), and McEachern’s Cave, Victoria (Rich and van Tets 1982), which also produced fossils of C. tibicen. Fossils referable to Cracticidae were reported from the Pleistocene Coolup Bore, Western Australia (Rich and van Tets 1982) and Buchan Caves, Victoria (Baird 1991), as well as the Holocene Amphitheatre Cave, Victoria (Baird 1992). The earliest representative of the Cracticidae in New Zealand is an indeterminate scapula from the Miocene St Bathans Fauna (Worthy et al. 2007). Here we describe the first pre-Quaternary cracticid species from Australia, based on a proximal tarsometatarsus. This specimen is from the Riversleigh World Heritage Area, north-western Queensland, which is the richest Tertiary site for fossil passerines in Australia. Passerines reported so far from the Riversleigh deposits include a lyrebird Menura tyawanoides, logrunner Orthonyx kaldowinyeri, oriolid Longmornis robustirostrata, a corvid-like passerine Corvitalusoides grandiculus and honeyea-

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ters (Meliphagidae) (Boles 1993, 1995, 1999, 2005, 2006). We add this new species to the diversity of Riversleigh passerines and discuss its significance regarding calibration of nodes in molecular estimates of the passerine phylogeny. We also discuss tarsometatarsal characters that either differ or are shared between cracticids and Artamus. The fragmentary nature of the fossil precludes an informative phylogenetic analysis of its relationships. A comprehensive phylogenetic analysis of the Australasian passerines, based on morphological data, is in preparation and outside the scope of the present contribution. Materials and methods The fossil specimen described here is registered in the palaeontological collections of the Queensland Museum, Brisbane, Australia (QM F). Comparisons were made with specimens of extant taxa (Appendix 1) in the collections of the Australian Museum, Sydney (AM) and Museum Victoria, Melbourne (NMV). Measurements were made with digital vernier callipers accurate to 0.01 mm and rounded to the nearest 0.1 mm. Terminology of anatomical structures follows Baumel and Witmer (1993) and Baumel and Raikow (1993). Some of the abbreviations used in the text are: lig., ligamentum; M., musculus; proc., processus; tub., tuberculum. Taxonomic nomenclature Nomenclature and systematic grouping of taxa follows Dickinson (2003), with the exception of the Australian Magpie, which is regarded here as Cracticus tibicen. This taxonomic treatment of Australian Magpie follows Storr (1952), Johnstone and Storr (2004) and Christidis and Boles (2008), who suggest that its retention in a separate genus, Gymnorhina, on the basis of terrestrial adaptation (e.g. Amadon 1951; Schodde and Mason 1999) is unwarranted. These authors consequently include Gymnorhina in Cracticus, which is supported by recent genetic studies (Kearns et al. 2013). Systematic palaeontology Order PASSERIFORMES Linnaeus, 1758 Family CRACTICIDAE Chenu & des Murs, 1853 (1836) Kurrartapu Nguyen, gen. nov. Type-species Kurrartapu johnnguyeni Nguyen, sp. nov. Included species Type-species only. Differential diagnosis The fossil is referred to Passeriformes because of the combined presence of the following features: prominent crista plantaris lateralis; hypotarsal canal for M. flexor hallucis longus (fhl) tendon is separate from and lateral to canal for M. flexor

A new cracticid from Early Miocene Australia

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digitorum longus (fdl) tendon; in proximal view, tub. M. fibularis brevis is lateral to fhl canal. Kurrartapu is assigned to Cracticidae based on the following combination of character states (Figs 1, 2):

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clade to the exclusion of Peltops. The exact relationships between cracticids and Artamus, which have at times been collectively treated as a single family Artamidae (e.g. Christidis and Boles 2008; see above), are unresolved in the molecular data. Because of this uncertainty, we have also included here comparisons of the fossil with Artamus. Character states 1 and 2, which are considered apomorphies for the Strepera–Cracticus clade, are shared with Kurrartapu but not with Peltops or Artamus. Character state 2, in particular, is not observed in any passerines except Strepera, Cracticus and Kurrartapu. Peltops and Artamus differ from these birds by having both the fp3–4 canal (character state 1) and retinaculum extensorium tarsometatarsi (character state 2) completely ossified. The retinaculum in Peltops forms an osseous arch (arcus extensorius) that is situated at the midline of the shaft, but is, ossified or otherwise, located more medially in Strepera, Cracticus, Kurrartapu, Artamus and most passerines. This feature is an autapomorphy for Peltops. Also, the lateral margin of the retinaculum extensorium tarsometatarsi in Peltops is level with the dorsal rim of the cotyla lateralis, not more distally as in other cracticids, Artamus and most passerines. Kurrartapu is also distinguished from Peltops and Artamus by its considerably larger size. The total length of the tarsometatarsus of Peltops or Artamus is less than the preserved length of the fossil proximal tarsometatarsus. These similarities in morphology that Kurrartapu shares with other cracticids, but not with Peltops and Artamus, suggest that it is more closely related to the Strepera– Cracticus clade than to a more inclusive clade including Peltops and Artamus. Kurrartapu is further distinguished from Artamus by the following features. The medial shaft surface of the tarsometatarsus in Kurrartapu, Strepera and some species of Cracticus is deep, whereas in Artamus it is very shallow and essentially a sharp crest. It is shallow in C. quoyi, C. nigrogularis (Pied Butcherbird),

(1) The hypotarsal canal for M. flexor perforatus digit III and IV (fp3–4) tendons is incompletely ossified and plantarly open. Although the distal section of the lateral bony ridge that bounds the fp3–4 canal is broken, the proximal section is completely preserved. It does not show any sign of breakage that would indicate that the canal was once plantarly closed. A plantarly open fp3–4 canal was observed in all cracticid specimens and does not appear to ossify completely with age. (2) The retinaculum extensorium tarsometatarsi that guides the tendon of the M. extensor digitorum longus is incompletely ossified, but marked by two bony ridges that define its location, length and depth. As with the fp3–4 canal, incomplete ossification was observed in all available cracticid specimens and does not appear to be an ontogenetic effect. (3) The groove that holds the tendon of the M. extensor digitorum longus is directed proximo-laterally to the midpoint of eminentia intercotylaris. (4) The crista plantaris lateralis is relatively low compared to many other passerines. Its plantar extent is less than that of the hypotarsus. (5) The tub. for insertio M. gastrocnemius pars medialis protrudes further medially than the medial rim of cotyla medialis. (6) The fossa on the dorso-lateral side of the eminentia intercotylaris is deeply excavated. (7) The cotyla medialis is situated proximal to the cotyla lateralis. (8) Both foramina vascularia proximalia are situated immediately proximal to the tub. M. tibialis cranialis. Recent molecular studies (e.g. Fuchs et al. 2012; Kearns et al. 2013) have shown that Strepera and Cracticus form a

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Fig. 1. Proximal right tarsometatarsus of Kurrartapu johngnuyeni, gen. nov., sp. nov. (QM F56251) in dorsal (a), plantar (b), lateral (c), medial (d) and proximal (e) views. fdl, canal for M. flexor digitorum longus; fhl, canal for M. flexor hallucis longus; fp3–4, canal for M. flexor perforatus digit III and IV; gft, grooves for flexor tendons; gpm, impressio M. gastrocnemius pars medialis; ilcm, impressio lig. collateralis medialis; lret, lateral ridge of retinaculum extensorium tarsometatarsi; mret, medial ridge of retinaculum extensorium tarsometatarsi; tfb, tub. fibularis brevis; ttc, tuberositas M. tibialis cranialis. Scale bar: 2 mm.

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Fig. 2. Proximal right tarsometatarsus of Kurrartapu johnnguyeni, gen. nov., sp. nov. (b) in comparison with tarsometatarsi of (a) Strepera graculina (AM O.66132) and (c) Cracticus tibicen (AM O.71169), viewed dorsally. See caption for Fig. 1 for abbreviations. Scale bar: 2 mm.

C. mentalis (Black-backed Butcherbird) and Peltops, but not to the extent seen in Artamus. In Kurrartapu and all other cracticids, the retinaculum extensorium tarsometatarsi is aligned at a less obtuse angle to the shaft long axis than in Artamus. The crista plantaris medialis immediately distal to the hypotarsus is poorly developed in Kurrartapu and all other cracticids. In Artamus, however, this crista is sharp and prominent, and merges with the crista medialis hypotarsi. Artamus also possesses the autapomorphic presence of a distinct fossa dorsal to the impressio for M. gastrocnemius pars medialis. This fossa is shallow in some individuals, but in others it is very deep and extends distally to form a groove that trends dorsally of the impressio lig. collateralis medialis. Kurrartapu differs from all other cracticids in possessing the following suite of features. The groove for the M. extensor digitorum longus is deeply incised into the dorsal rim of cotyla medialis, whereas it is shallow in Strepera and indistinct in Cracticus. There is a deep fossa plantar to the eminentia intercotylaris, which is very shallow in C. mentalis and C. nigrogularis, and absent in all other cracticids. The bony ridges of the retinaculum extensorium tarsometatarsi are low and proximo-distally long, whereas in Strepera and Cracticus these ridges are short. Plantarly, the longitudinal grooves immediately distal to the hypotarsus for the flexor tendons of the digits are moderately deep, but shallow in Strepera, C. mentalis and C. tibicen, and indistinct in other cracticids. The fossa infracotylaris is very deep, but shallow in Strepera and Cracticus, and very shallow in Peltops. Kurrartapu is also peculiar in that the medial edge of the cotyla medialis is pointed proximally, which is absent in all other cracticids. Peltops is further distinguished from Strepera and Cracticus by the following features of the distal tarsometatarsus: the trochlea metatarsi II and III have equal distal extent, rather than trochlea metatarsi III extending more distally; the medial and lateral rims of trochlea metatarsi III are

equal in length, instead of the medial rim being proximo-distally longer; and the presence of a shallow groove on the distal surface of the trochlea metatarsi II, which is absent in all other extant cracticids and Artamus. Other groups of large passerines currently found in Australia include the lyrebirds (Menuridae), crows and ravens (Corvidae), White-winged Chough (Corcoracidae : Corcorax melanorhamphos), bowerbirds (Ptilonorhynchidae), birds-ofparadise (Paradisaeidae) and a number of large species of honeyeater (Meliphagidae). The fossil is distinguished from these groups because the fp3–4 hypotarsal canal is incompletely ossified (character state 1), despite osteological maturity of the specimen. Although a plantarly open canal for the fp3–4 tendons was observed in one specimen of Corvus orru (Torresian Crow (AM O.65439)), it was completely ossified in other specimens of C. orru, as well as in C. mellori (Little Raven) and C. coronoides (Australian Raven) (Appendix 1). The plantar wall of this canal is ligamentous in younger individuals and progressively ossifies with age. Kurrartapu differs from the Riversleigh raven-like passerine Corvitalusoides grandiculus (see Boles 2006), which was described from a distal tibiotarsus, by its distinctly smaller size. Kurrartapu is excluded from families characterised by small to very small species, such as the fairy-wrens and grasswrens (Maluridae), because of its considerably larger size and plantarly open fp3–4 canal. Etymology From the northern Queensland Aboriginal word kurrartapu, for magpie (Breen 1981); gender is masculine. Kurrartapu johnnguyeni Nguyen, sp. nov. Holotype QM F56251, a proximal right tarsometatarsus with most of the plantar section of the hypotarsus broken off. Type-locality and age Price is Right Site, Riversleigh World Heritage Area, northwestern Queensland, Australia. Based on stage-of-evolution biocorrelation, multivariate studies and radiometric dating currently underway, sites in Riversleigh have been allocated a relative age and categorised into Faunal Zones A to D (Archer et al. 1989, 1997, 2006; Creaser 1997; Travouillon et al. 2006, 2009). Price is Right Site has been interpreted to represent Faunal Zone B, hence Early Miocene in age (Archer et al. 1997; Travouillon et al. 2006). Measurements of holotype Total length as preserved 19.9 mm; proximal width 7.2 mm; proximal depth, from cotyla lateralis to plantar edge of fp3–4 canal 5.8 mm; depth of cotyla lateralis 3.8 mm; depth of cotyla medialis 4.2 mm. Diagnosis As for genus, by monotypy.

A new cracticid from Early Miocene Australia

Description and comparison The following description concerns features that are not already described in the genus differential diagnosis (above). In our comparisons, we note only those features that differ between Kurrartapu and the extant cracticids. The fossil tarsometatarsus is comparable in size to that of Cracticus quoyi. Its proximal width is approximately twice the shaft width immediately distal to the tuberositas M. tibialis cranialis. The plantar wall of the hypotarsal canal for the fdl tendon is broken off, but most of the plantar wall of the fhl canal is preserved. The fdl canal appears to have a slightly smaller diameter than the fhl canal. In Strepera graculina, S. versicolor and C. tibicen this canal is considerably smaller in diameter than the fhl canal. Kurrartapu has a low and rounded eminentia intercotylaris. Its proximal margin is slightly angled proximo-laterally. The bony ridges of the retinaculum extensorium tarsometatarsi are slightly abraded but show no evidence of having been completely ossified. The medial and lateral bony ridges of the retinaculum extensorium tarsometatarsi are at a shallow angle to the shaft long axis. The groove for the M. extensorium digitorum longus tendon trends proximo-laterally to the midpoint of the eminentia intercotylaris. The tuberositas M. tibialis cranialis is elongate, directed dorso-laterally, and is situated slightly medial to the shaft midline. The proximal part of the tuberosity is damaged but the distal part is markedly elevated dorsally. The shaft becomes slightly narrower distally. Kurrartapu has a very shallow sulcus extensorius with a rounded lateral margin, but in Cracticus this margin is sharply convex. The medial margin of the sulcus extensorius is more sharply defined in the fossil than in all other cracticids examined. The medial shaft surface is plantarly deep and sharply convex. It is also deep in Strepera graculina, S. versicolor, C. tibicen and C. torquatus (Grey Butcherbird), but more rounded or planar. In C. quoyi, C. mentalis, C. nigrogularis and Peltops montanus (Mountain Peltops), the medial shaft surface is relatively shallower compared to the proximal width of the tarsometatarsus. The impressio lig. collateralis medialis is aligned with the medial edge of the sulcus extensorius. In Kurrartapu it is narrow and pronounced, but in C. quoyi, C. torquatus, C. mentalis and P. montanus the impressio lig. collateralis medialis is low. In Kurrartapu, the sulcus flexorius is very shallow and gently sloped medially. In Strepera versicolor and all Cracticus species examined, the sulcus is also shallow but is level medio-laterally. Within this sulcus, the two grooves immediately distal to the hypotarsus for the flexor tendons are separated by a low ridge and become shallower distally. The fossa parahypotarsalis medialis is absent in Kurrartapu and all other cracticids except for C. quoyi, C. nigrogularis and Peltops, in which the fossa is shallow to moderately deep. In the fossil, the proximo-plantar corner of the crista plantaris lateralis is rounded. In C. torquatus, C. nigrogularis, C. tibicen and Peltops, the proximo-plantar corner of this crista is a sharp edge. The plantar scar for attachment of M. fibularis longus is flat and elongate, whereas in all Strepera species examined this muscle scar is protuberant. The plantar passage of the groove

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for M. fibularis longus is indistinct in the fossil but shallow in Strepera versicolor, C. tibicen, C. quoyi, C. torquatus and C. nigrogularis. The lateral surface of the shaft is shallowly concave, unlike in S. graculina, C. torquatus, C. mentalis, C. nigrogularis and Peltops. On the lateral surface of the shaft the impressio lig. collateralis lateralis is protuberant in S. versicolor and in some specimens of S. graculina and C. tibicen, but is indistinct or low in Kurrartapu and other cracticids. Etymology The species name is dedicated in loving memory of the father of the first author, John Van Giao Nguyen. Discussion Kurrartapu johnnguyeni is the geologically oldest and first Tertiary fossil cracticid to be described from Australia. Although its description is based on a single proximal tarsometatarsus, it is distinct enough from other cracticids to warrant its assignation to a new genus and species. Kurrartapu johnnguyeni is more similar to species of Strepera and Cracticus than to Peltops because it shares a plantarly open hypotarsal canal for the fp3–4 tendons (character state 1) and an incompletely ossified retinaculum extensorium tarsometatarsi (character state 2), which are apomorphies for the Strepera–Cracticus clade. It is also similar to this clade in lacking a centrally located retinaculum that is level with the dorsal rim of the cotyla lateralis, which is the autapomorphic condition observed in Peltops. Kurrartapu is distinguished from Artamus, which is sometimes treated as confamilial with cracticids (e.g. Christidis and Boles 2008), in having character states 1 and 2. It also lacks a fossa situated dorsally of the impression for M. gastrocnemius pars medialis, which is an autapomorphy of Artamus. Species of Kurrartapu, Strepera and Cracticus are also considerably larger in size than Peltops and Artamus. These features suggest that Kurrartapu johnnguyeni is more closely related to the Strepera–Cracticus clade than to the Peltops or Artamus lineages and, when Strepera, Cracticus and Peltops comprise Cracticidae (sensu Dickinson 2003), it can be placed within the crown Cracticidae rather than on the stem lineage. Complete ossification of all hypotarsal tendinal canals except the fp3–4 canal is rare in passerines. Among the Australasian groups, this feature was found only in the Noisy Scrub-bird (Atrichornis clamosus) and White-throated Treecreeper (Cormobates leucophaea) of Australia and the Kokako (Callaeas cinerea) of New Zealand (J. M. T. Nguyen, unpubl. data; see Appendix 1). A plantarly open fp3–4 canal has been reported in the Old World broadbills (Eurylaimidae) (Manegold et al. 2004), neotropical ovenbirds (Furnariidae sensu Manegold 2008b) and woodcreepers (Dendrocolaptidae), and the ‘climbing Certhioidea’ (Manegold 2008b). Incomplete ossification of the retinaculum extensorium tarsometatarsi, which may be a reversal to an ancestral state, is not known in any other passerine group except the Strepera–Cracticus clade. It is possible that these two diagnostic features were lost in the Strepera–Cracticus clade after divergence from the Peltops lineage. Comparisons with the African bush-shrikes and allies (which were not available for the current study) will

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contribute to understanding how these features evolved in this part of the passerine radiation. As a probable representative of the crown Cracticidae, Kurrartapu johnnguyeni is significant because it provides a minimum age for the divergence between the Peltops and the Strepera–Cracticus clades. Accordingly, it can be used as a fossil calibration in molecular estimates of divergence times within this part of the passerine radiation. After the Riversleigh oriolid Longmornis robustirostrata (see Boles 1999) and the as-yet-unnamed cracticid from St Bathans in New Zealand (Worthy et al. 2007), this is the third Tertiary representative of the core Corvoidea (sensu Barker et al. 2004) to be described so far. Owing to similarities with members of Corvidae and Cracticidae (Boles 2006), Riversleigh’s Corvitalusoides grandiculus is also possibly a member of the core Corvoidea. However, it cannot be regarded as a reliable fossil calibration for dating the passerine evolutionary time-scale because of its uncertain taxonomic position beyond subordinal level. The Early Miocene palaeoenvironment for Kurrartapu johnnguyeni has been interpreted to be closed forest (Archer et al.1989; Travouillon et al. 2009; Black et al. 2012). Faunal Zone B assemblages are characterised by very high species diversity, presence of rainforest-indicative taxa such as rat-kangaroos (Hypsiprymnodon), lyrebirds (Menura) (Boles 1995), striped possums (Dactylopsila), cuscuses (Onirocuscus) and an abundance of arboreal marsupials (e.g. Archer et al. 1997; Roberts et al. 2007) and hipposiderid bats (Microchiroptera) (Hand 1997; Hand and Archer 2005) typical of closed forest communities. Other birds that have been identified from the Price Is Right Site include the casuariid Emuarius gidju (see Worthy et al., in press) and the dromornithid Barawertornis tedfordi (see Nguyen et al. 2010). Extant cracticids occupy a diverse range of foraging niches and are found in many terrestrial habitats, including rainforest, so it is not surprising to find a representative of this family in this palaeoenvironment. Kurrartapu johnnguyeni may be older or of similar age to the fossil cracticid from the late Early Miocene St Bathans Fauna (19–16 million years ago) (Worthy et al. 2007). The St Bathans cracticid is similar in size to Cracticus tibicen and Strepera, and may have been slightly larger than Kurrartapu johnnguyeni. The age of Kurrartapu johnnguyeni is consistent with molecular estimates for the evolutionary time-scale of the Cracticidae. Several studies have estimated the divergence of the Cracticidae from the Artamus lineage to be between 20 and 36.3 million years ago (e.g. Fuchs et al. 2006; Jønsson et al. 2011; Kearns et al. 2013). A multilocus species tree for the cracticids yielded an estimated date of 16.9– 28.3 million years ago for the split between the Peltops lineage and Strepera–Cracticus clade, and 9.8–17.3 million years ago for the divergence between Strepera and Cracticus (Kearns et al. 2013). Greater resolution of the phylogenetic relationships among Cracticidae, Artamus and the African bush-shrikes and allies will be improved with additional morphological data from other skeletal elements, supplemented by expanded molecular analyses with wider sampling of these groups. Together, these would help in assessing whether Artamus and cracticids should be merged into a single family, and in resolving the evolutionary history of the Malaconotoidea.

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Acknowledgements We thank Jaynia Sladek (AM) for access to comparative material and providing J. M. T. Nguyen a workspace. We also thank Wayne Longmore (NMV) for facilitating access to additional reference material. Research at Riversleigh is supported by the Australian Research Council (LP100200486, DP1094569, DP130100197 DE130100467 grants to M. Archer, S. J. Hand and K.H. Black at the University of New South Wales (UNSW)), XSTRATA Community Partnership Program (North Queensland), the University of New South Wales, Queensland Parks and Wildlife Service, Environment Australia, the Queensland Museum, the Riversleigh Society Inc., Phil Creaser and the CREATE Fund, Outback at Isa, Mount Isa City Council, private supporters, including Ken and Margaret Pettit, the Carpentarian Land Council and the Waanyi people of north-western Queensland. We are grateful for the field assistance of many volunteers at Riversleigh as well as staff and postgraduate students of UNSW. J. M. T. Nguyen is supported by grants from the Linnean Society of New South Wales (Betty Mayne Scientific Research Fund for Earth Sciences), BirdLife Australia (Stuart Leslie Bird Research Award) and the CREATE Fund. We thank Simon Ho for providing helpful comments on an earlier draft of this manuscript, and Gerald Mayr and an anonymous reviewer for their constructive feedback.

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Appendix 1. Skeletons of a wide range of passerines were examined in the collections of the Australian Museum, Sydney (AM) and Museum Victoria, Melbourne (NMV). Detailed comparisons were made with the following taxa which were most relevant owing to their close affinity, similar size and geographical source in Australia. Menuridae: Menura novaehollandiae (Superb Lyrebird) AM O.60220; Atrichornithidae: Atrichornis clamosus (Noisy Scrub-bird) NMV R11354; Climacteridae: Cormobates leucophaea (White-throated Treecreeper) AM O. 66622; Ptilonorhynchidae: Ailuroedus crassirostris (Green Catbird) AM O.64754, Sericulus chrysocephalus (Regent Bowerbird) AM O.60064, Ptilonorhynchus violaceus (Satin Bowerbird) AM O.64555; Meliphagidae: Anthochaera carunculata (Red Wattlebird) AM O.60082, Philemon citreogularis (Little Friarbird) AM O.65440; Callaeidae: Callaeas cinerea (Kokako) AM A1983; Artamidae: Artamus leucorynchus (White-breasted Woodswallow) AM O.60048, AM O.71161, Artamus personatus (Masked Woodswallow) AM O.72300, AM O.66305, Artamus superciliosus (White-browed Woodswallow) AM O.71701, AM O.64740, AM O.60054, Artamus cinereus (Black-faced Woodswallow) AM O.71159, AM O.71580, Artamus cyanopterus (Dusky Woodswallow) AM O.58036, AM O.65088, AM O.64840, Artamus minor (Little Woodswallow) AM O.65129, AM O.67050; Cracticidae: Cracticus quoyi (Black Butcherbird) AM O.64878, Cracticus torquatus (Grey Butcherbird) AM O.60037, Cracticus mentalis (Black-backed Butcherbird) AM O.59645, Cracticus nigrogularis (Pied Butcherbird) AM O.66326, AM O.58063, Cracticus tibicen (Australian Magpie) AM O.71169, AM S10, AM O.70776, O.65177, Strepera graculina (Pied Currawong) AM O.66110, AM O.66331, AM O.66139, AM O.57129, AM O.71396, Strepera versicolor (Grey Currawong) AM O.71398, AM O.71363, Peltops montanus (Mountain Peltops) AM O.46418; Corvidae: Corvus coronoides (Australian Raven) AM O.60383, Corvus mellori (Little Raven) AM O.70738, Corvus orru (Torresian Crow) AM O.65439, AM O.71330; Corcoracidae: Corcorax melanorhamphos (White-winged Chough) AM O.59864; Paradisaeidae: Ptiloris victoriae (Victoria’s Riflebird) AM O.68473, Manucodia comrii (Curl-crested Manucode) AM S815; Paradisaea rudolphi (Blue Bird-ofparadise) AM S1477.

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