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New specimens of the logrunner Orthonyx kaldowinyeri (Passeriformes: Orthonychidae) from the Oligo-Miocene of Australia Jacqueline M.T. Nguyen, Walter E. Boles, Trevor H. Worthy, Suzanne J. Hand & Michael Archer Published online: 10 Dec 2013.
To cite this article: Jacqueline M.T. Nguyen, Walter E. Boles, Trevor H. Worthy, Suzanne J. Hand & Michael Archer , Alcheringa: An Australasian Journal of Palaeontology (2013): New specimens of the logrunner Orthonyx kaldowinyeri (Passeriformes: Orthonychidae) from the Oligo-Miocene of Australia, Alcheringa: An Australasian Journal of Palaeontology, DOI: 10.1080/03115518.2014.861732 To link to this article: http://dx.doi.org/10.1080/03115518.2014.861732
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New specimens of the logrunner Orthonyx kaldowinyeri (Passeriformes: Orthonychidae) from the Oligo-Miocene of Australia JACQUELINE M.T. NGUYEN, WALTER E. BOLES, TREVOR H. WORTHY, SUZANNE J. HAND and MICHAEL ARCHER
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NGUYEN, J.M.T., BOLES, W.E., WORTHY, T.H., HAND, S.J. & ARCHER, M., 2014. New specimens of the logrunner Orthonyx kaldowinyeri (Passeriformes: Orthonychidae) from the Oligo-Miocene of Australia. Alcheringa 38, 000–000. ISSN 0311–5518. Logrunners (Orthonychidae) are a family of ground-dwelling passerines that are endemic to the Australo-Papuan region. These peculiar birds are part of an ancient Australo-Papuan radiation that diverged basally in the oscine tree. Here we describe eight fossil tarsometatarsi of the logrunner Orthonyx kaldowinyeri, and a distal tibiotarsus tentatively assigned to this species from sites in the Riversleigh World Heritage Area, Australia. The new fossil material ranges in age from late Oligocene to early late Miocene, and extends the temporal range of the Orthonychidae into the late Oligocene; this is the geologically oldest record of the family. These specimens also include the oldest Cenozoic passerine fossils from Australia that can be confidently referred to an extant family. The distinctive features of the tarsometatarsus and tibiotarsus of extant logrunners, which are probably related to their unusual method of foraging, are also present in O. kaldowinyeri. Assuming that O. kaldowinyeri had vegetation requirements similar to those of extant logrunners, its presence in various Riversleigh sites provides clues about the palaeoenvironment of these sites. Jacqueline M.T. Nguyen [
[email protected]] (author for correspondence), Suzanne J. Hand [
[email protected]], Michael Archer [
[email protected]], School of Biological, Earth and Environmental Sciences, University of New South Wales, Sydney, NSW 2052, Australia; Walter E. Boles [
[email protected]], Ornithology Section, Australian Museum, 6 College Street, Sydney, NSW 2010, Australia; Trevor H. Worthy [trevor.worthy@flinders.edu.au], School of Biological Sciences, Flinders University, Adelaide, SA 5001, Australia. Received 19.9.2013; revised 11.10.2013 accepted 25.10.2013. Key words: logrunner; Orthonyx; passerine; Oligo-Miocene; tarsometatarsus; Riversleigh; Australia.
LOGRUNNERS (Orthonychidae) are a unique Australo-Papuan family of ground-dwelling passerines. This small family contains a single genus with three extant species, two of which are represented in Australia: the Chowchilla Orthonyx spaldingii Ramsay, 1868, restricted to northeastern Queensland, and the Australian Logrunner, O. temminckii Ranzani, 1822, confined to southeastern Queensland and eastern New South Wales (Higgins & Peter 2002). The New Guinean Logrunner, O. novaeguineae Meyer, 1874, has at times been treated as conspecific with O. temminckii, but morphological and molecular studies show a deep divergence between the two (Joseph et al. 2001, Norman et al. 2002). Orthonyx spaldingii, the largest of the three species, inhabits tropical rainforests. The smaller O. temminckii is found in subtropical rainforests and adjacent wet sclerophyll forest, and O. novaeguineae in highland montane forests (Coates 1990, Higgins & Peter 2002). All three species are sexually dimorphic, with males larger with a white throat and upper breast, whereas females are smaller and rufous (Boles 2007).
© 2013 Association of Australasian Palaeontologists http://dx.doi.org/10.1080/03115518.2014.861732
Logrunners display a remarkable method of foraging, which involves a sweeping motion of their powerful feet, either alternately or with several successive kicks of one foot, in wide lateral arcs to push aside leaf litter (summarized by Boles 2007). During sweeping, the opposite leg is considerably flexed at the ankle to support the body and, for additional stability, the spiny tail is sometimes angled downwards into the litter. Upon removal of the large debris, logrunners use their tails as a brace and scratch the exposed soil to uncover insects and other invertebrates (Hindwood 1934, Boles & Shields 1980). This foraging manner is associated with a distinctive structure of the pelvis and hind limb (reviewed by Schodde & Mason 1999). For example, the pelvis is short and broad, and the fossae iliacae dorsales are large and deeply excavated to accommodate an expanded M. iliotrochantericus caudalis. A similar pelvic structure is also found in other passerines, such as the Yellowhead Mohoua ochrocephala (Gmelin 1789) and fernbirds Bowdleria spp. Rothschild, 1896, which also use their feet to move vegetation and other debris while foraging (Olson 1990a, b). However, the sideways sweeping action of the legs is unique to logrunners. Unlike Mohoua and Bowdleria, the femur
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in Orthonyx is extremely stout with wide proximal and distal ends relative to the shaft, and has strongly developed projections on the trochanter and distal end for muscle attachments (Olson 1990a, b, JMTN, pers. obs.). Other structural adaptations to this unusual foraging manner include thickly scaled ankles, syndactylous toes and stiffened rectrices that protrude beyond the vanes as spines (Zusi 1978). Previously grouped with quail-thrushes Cinclosoma Vigors & Horsfield, 1827, jewel-babblers Ptilorrhoa Peters, 1940 and allies (e.g., Deignan 1964), logrunners were shown to form a distinct monotypic family based on DNA-DNA hybridization data (Sibley & Ahlquist 1990). Recent molecular analyses have demonstrated that the Orthonychidae is an ancient endemic lineage that is part of the early Australo-Papuan passerine radiation. Along with lyrebirds (Menuridae), scrub-birds (Atrichornithidae), Australasian treecreepers (Climacteridae), bowerbirds (Ptilonorhynchidae), Meliphagoidea (sensu Gardner et al. 2010) and Australasian babblers (Pomatostomatidae), logrunners are one of the sequential sister groups to all other oscine passerines (e.g., Barker et al. 2004, Hugall & Stuart-Fox 2012). Owing to their basal position among the oscines, the fossil record of logrunners has a potentially significant role in interpreting the evolution of crown passerines. The fossil record of this family indicates that they had a more extensive geographical range than is currently demonstrated. The Pleistocene record is represented by Orthonyx hypsilophus Baird, 1985 from Green Waterhole Cave, southeastern South Australia, which was larger than the extant O. spaldingii. A second Pleistocene orthonychid, O. wakefieldi Baird, 1993, slightly smaller than O. temminckii, was described from Pyramids Cave, eastern Victoria. The earliest known representative is O. kaldowinyeri Boles, 1993, described from a complete femur from the middle Miocene of the Riversleigh World Heritage Area, northwestern Queensland. Here we describe new fossil material from Riversleigh that is referred to Orthonyx kaldowinyeri. One complete tarsometatarsus, seven partial tarsometatarsi and one partial tibiotarsus have been recovered from sites at Riversleigh that span the late Oligocene to early late Miocene. These fossils extend the temporal range of the Orthonychidae back into the Paleogene. We also discuss the palaeoecological implications of these fossils and their significance for calibrating molecular estimates of the early passerine radiation.
Materials and methods The fossil specimens described here are registered in the palaeontology collections of the Queensland Museum, Brisbane (QM). Direct comparisons were made with the following specimens of extant taxa in the ornithology collections of the QM, Australian Museum, Sydney (AM), Museum Victoria, Melbourne
ALCHERINGA (NMV) and Australian National Wildlife Collection, CSIRO, Canberra (ANWC). The sex of specimens (M male, F female) is given if known. Orthonychidae: Chowchilla Orthonyx spaldingii AM O.3515 M, AM O.67652 F, QM O21124, QM O27876 F; Australian Logrunner O. temminckii AM O.59360, AM O.59361, AM O.62494, AM O.64813 F, ANWC 20000 F, ANWC 46911 M, QM O22530 F, QM O25471 M; Dasyornithidae: Rufous Bristlebird Dasyornis broadbenti (McCoy, 1867) NMV B.20931; Pomatostomidae: Grey-crowned Babbler Pomatostomus temporalis (Vigors & Horsfield 1827) AM O.68150. Measurements of fossil material were made with a Wild MMS 235 digital length-measuring unit on a Leica Wild M3B microscope at the University of New South Wales (UNSW). Measurements of specimens of extant species and the O. kaldowinyeri holotype femur (QM F16867) were made with digital vernier callipers and rounded to the nearest 0.1 mm. Terminology of anatomical structures follows Baumel & Witmer (1993) and Baumel & Raikow (1993). Some abbreviations used in the text: lig., ligamentum; M., musculus; proc., processus; tub., tuberculum. Taxonomic nomenclature follows Christidis & Boles (2008). Log-ratio diagrams Log-ratio diagrams were constructed following Simpson (1941) to compare the relative proportions of the fossil tarsometatarsi with those of O. spaldingii and O. temminckii. Because the O. kaldowinyeri holotype femur can not be directly compared with the fossil tarsometatarsi, we incorporated in the diagrams several femoral measurements from O. kaldowinyeri and extant Orthonyx species. The log-ratio diagram makes it possible to compare different elements and postulate whether they belong to the same taxon based on similarity in ratios of dimensions. The fragmentary nature of the fossil tibiotarsus precluded meaningful comparisons of measurements with those of extant logrunners in the log-ratio diagram. Various measurements were made on the tarsometatarsus and femur for all specimens (Table 1). These measurements were averaged for each taxon and converted to their common logarithm; measurements of incomplete landmarks were omitted. One taxon was arbitrarily chosen as the standard for comparison to represent zero difference in logarithmic values. Here we chose an unrelated passerine, Dasyornis broadbenti, as the standard taxon to emphasise the morphological pattern of logrunners and any variation between the fossil and extant taxa. The differences between the logarithmic values of the standard and those of the other specimens were calculated and plotted on a graph. Points representing different measurements of the same specimen were connected with a line. Values that are smaller than the standard fall below the standard baseline, whereas values that are larger than the standard
ALCHERINGA
FOSSIL LOGRUNNERS FROM AUSTRALIA Orthonyx kaldowinyeri
Orthonyx spaldingii n = 4
Orthonyx temminckii n = 8
ca 26.1 >3.9 (>3.7– >4.0) >3.0 (>2.8– >3.2) 1.7 3.3 (2.3–3.8) 1.7 (1.3–1.9) 1.7 (1.2–1.9) 1.3 (>0.9– >1.5)
45.3 (42.9–49.5) 7.4 (7.1–8.0) 7.9 (7.3–8.5) 2.7 (2.1–3.2) 5.5 (5.4–6.0) 3.4 (3.2–3.8) 2.8 (2.5–3.2) 2.5 (2.2–2.9)
31.9 (30.1–32.9) 4.9 (4.5–5.2) 5.3 (4.7–5.8) 1.9 (1.8–2.1) 3.7 (3.6–4.0) 2.3 (2.0–2.6) 2.0 (1.8–2.8) 1.6 (1.3–1.8)
ca 4.4
6.9 (6.5–7.4)
4.7 (4.3–5.2)
19.7 6.1 3.5 2.8 2.6 2.1 5.7 3.8 2.8 2.9
31.6 (30.1–33.4) 9.2 (8.8–9.8) 5.5 (5.3–5.7) 4.4 (4.1–4.9) 4.0 (3.8–4.4) 3.1 (3.0–3.2) 9.0 (8.6–9.7) 6.0 (5.5–6.9) 4.4 (3.9–5.0) 5.0 (4.0–6.0)
23.5 (>22.0–24.5) 7.0 (6.7–7.4) 4.3 (4.0–4.9) 3.4 (3.1–3.6) 2.9 (2.7–3.3) 2.4 (2.2–2.6) 6.4 (>5.2–6.7) 4.3 (>2.4–4.6) 3.1 (2.8–3.4) 3.6 (>2.1–4.2)
Tarsometatarsus Total length Proximal width Proximal depth Least shaft width Distal width Trochlea metatarsi II depth Trochlea metatarsi III depth Trochlea metatarsi IV depth Tibiotarsus Distal width
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Femur Total length Proximal width Proximal depth Depth, caput femoris Mid-shaft width Mid-shaft depth Distal width Depth, condylus lateralis Depth, condylus fibularis Depth, condylus medialis
Table 1. Measurements (mm) of the tarsometatarsi and femora of Orthonyx kaldowinyeri compared with those of extant Orthonyx species. The mean and range of measurements for each species are given, where possible. The means were used to construct a log-ratio diagram of these species (see Fig. 4).
are above it. Specimens with lines that are parallel to the standard share similar proportions regardless of size. Where possible, the log maximum and log minimum values of landmark measurements for each taxon were also plotted.
Systematic palaeontology Order PASSERIFORMES Linnaeus, 1758 Family ORTHONYCHIDAE Gray, 1840 Orthonyx Temminck, 1820 Type species. Orthonyx temminckii Ranzani, 1822, by subsequent designation. Included species. Orthonyx spaldingii Ramsay, 1868; O. novaeguineae Meyer, 1874; O. hypsilophus Baird, 1985; O. wakefieldi Baird, 1993; O. kaldowinyeri Boles, 1993. Identification. The fossils are referred to Passeriformes because of the following suite of features: (tarsometatarsus) crista plantaris lateralis is prominent; hypotarsal canal for M. flexor hallucis longus (fhl) tendon is separate from and lateral to canal for M. flexor digitorum longus (fdl) tendon; in proximal view, tub. m. fibularis brevis is lateral to fhl canal; trochleae metatarsorum are aligned in the same dorso-plantar plane; (tibiotarsus) pons supratendineus is completely ossified; medial and lateral bony ridges of retinaculum m. fibularis are present; incisura intercondylaris is deep.
The fossil tarsometatarsi possess the following autapomorphies for Orthonyx. The impressio lig. collateralis lateralis is low but very large (Fig. 1E, F, 2C). The plantar surface immediately proximal to the incisura intertrochlearis lateralis is deeply excavated, whereas the plantar surface immediately proximal to the incisura intertrochlearis medialis is very shallowly excavated (Fig. 1C, D, 2E, G, I, K, M, O). The assignment of these specimens to Orthonyx is also based on the following combination of character states. The eminentia intercotylaris is low and has a very deep fossa on its dorso-lateral surface. The arcus extensorius is short relative to its width, and the groove for the M. extensor digitorum longus tendon beneath it is very deep. The proximal margin of the tuberositas m. tibialis cranialis is adjacent to the distal margin of the arcus extensorius. The lateral foramen vasculare proximale is distinctly larger than its medial companion. The crista plantaris medialis immediately distal to the hypotarsus is a sharp ridge. The trochlea metatarsi II is broad, has no furrow dorsally and distally, and is similar in width to trochlea metatarsi III. The incisura intertrochlearis medialis is very wide along its entire length. The fossa metatarsi I is shallow and situated far proximally of the trochlea metatarsi II. The trochlea metatarsi IV is distinctly narrower than the other trochleae metatarsorum, and its lateral margin is not contiguous with that of the shaft but is situated more medially. The fossil tibiotarsus shares the following character states with Orthonyx. The distal end and shaft are broad and robust. The sulcus extensorius immediately
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Fig. 1. Tarsometatarsus of Orthonyx kaldowinyeri (QM F56329, Neville’s Garden Site) (A, C, E, G) compared with that of the extant species Orthonyx temminckii (AM O.62494) (B, D, F, H). A, B, dorsal view; C, D, plantar view; E, F, lateral view; G, H, medial view. Abbreviations: ae, arcus extensorius; cpl, crista plantaris medialis; cpm, crista plantaris medialis; fd, foramen vasculare distale; fm, fossa metatarsi I; fo, fossa on eminentia intercotylaris; fpm, fossa parahypotarsalis medialis; hyp, hypotarsus; ilcl, impressio lig. collateralis lateralis; ilcm, impressio lig. collateralis medialis; in, very shallow plantar surface proximal to incisura intertrochlearis medialis; lfp, lateral foramen vasculare proximale; mfp, medial foramen vasculare proximale; pf, peroneal foramen; ri, ridge proximal to trochlea metatarsi II; ttc, tub. M. tibialis cranialis. Scale bar = 2 mm.
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ALCHERINGA
FOSSIL LOGRUNNERS FROM AUSTRALIA
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Fig. 2. Tarsometatarsi specimens of the fossil logrunner Orthonyx kaldowinyeri in dorsal (A, D, F, H, J, L, N), plantar (B, E, G, I, K, M, O) and lateral (C) views. A–C, QM F56330; D, E, QM F56328; F, G, QMF 30,244; H, I, QM F57237; J, K, QM F57238; L, M, QM F24461; N, O, QM F56331. Abbreviations: fu, fusion line of the metatarsal synostoses; mr, medial bony ridge of arcus extensorius. See Fig. 1 for other abbreviations. Scale bar = 2 mm.
Fig. 3. Distal tibiotarsus of Orthonyx sp. cf. O. kaldowinyeri QM F57239 (A, C, E) compared with that of O. temminckii (AM O.62494) (B, D, F). A, B, cranial view; C, D, caudal view; E, F, lateral view. Abbreviations: el, epicondylus lateralis; em, epicondylus medialis; lre, lateral tuberositas retinaculi extensoris; lrf, lateral tub. retinaculi fibularis; mct, medial crista trochlearis; mre, medial tuberositas retinaculi extensoris; mrf, medial tub. retinaculi fibularis; tcr, ridge on trochlea cartilaginis tibialis. Scale bar = 2 mm.
proximal to the pons supratendineus is very shallow. The distal part of the medial tuberositas retinaculi extensoris is level with the bony ridges of the retinaculum m. fibularis (Fig. 3A, B). In cranial view, the
condylus medialis is narrower than the condylus lateralis. The proximo-distal length of the pons supratendineus is about equal to or less than its width. The cranial rim of the condylus lateralis is thickened and the
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JACQUELINE M.T. NGUYEN et al.
epicondylaris lateralis is a prominent tuberosity that protrudes far laterally (Fig. 3E, F). Four of the specimens represent immature birds because they exhibit incomplete ossification of the articular facets and a pitted appearance of the surface of the bone (Campbell 1979). Specifically, QM F24461 (Fig. 2L, M) and QM F57238 (Fig. 2J, K) have incompletely ossified trochleae metatarsorum, QM F56331 (Fig. 2N, O) has visible fusion lines at the metatarsal synostoses, and the eminentia intercotylaris and cotylae of QM F56330 (Fig. 2A–C) are incompletely ossified. These specimens, however, can still be referred to Orthonyx because they possess the diagnostic features described above.
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Orthonyx kaldowinyeri Boles, 1993 (Figs 1, 2) Referred material. A right tarsometatarsus with hypotarsus and trochlea metatarsi IV broken off—QM F56329 (Neville’s Garden Site); proximal left tarsometatarsus with hypotarsus broken off—QM F56330 (Camel Sputum Site); distal left tarsometatarsi—QM F24461 (Encore Site), QM F56328 (Dirk’s Towers Site), QM F56331 (Upper Site), QM F57238 (Judith’s Horizontalis Site); distal right tarsometatarsi—QM F30244 (Hiatus Site), QM F57237 (Judith’s Horizontalis Site). See Table 1 for measurements. Occurrence. Riversleigh World Heritage Area, northwestern Queensland, Australia. Based on stage-ofevolution biocorrelation, stratigraphy and multivariate studies of more than 200 local faunas at Riversleigh, together with radiometric dating currently under way, sites in Riversleigh have been attributed to Faunal Zones A to D. These Faunal Zone assemblages are interpreted to be late Oligocene (A), early Miocene (B), middle Miocene (C) and late Miocene (D) in age (Archer et al. 1989, 1997, 2006, Creaser 1997, Travouillon et al. 2006, 2009). The fossil tarsometatarsi were recovered from deposits that represent Faunal Zones A (Hiatus Site), B (Neville’s Garden Site, Camel Sputum Site, Upper Site, Dirk’s Towers Site, Judith’s Horizontalis Site) and D (Encore Site) (ibid). Specific details of the locations of Riversleigh fossil sites noted in this paper have been lodged with and are available from the Queensland Museum. Identification. The fossil tarsometatarsi are conservatively referred to O. kaldowinyeri because they exhibit the expected size and proportions of an individual represented by the holotype femur QM F16867. They indicate that Orthonyx kaldowinyeri is distinguished from all other orthonychids in the following combination of features: overall small size; fossa parahypotarsalis medialis is deeper; fossa infracotylaris dorsalis is deeper; trochlea metatarsi III is surpassed distally by trochlea metatarsi II; ridge proximal to trochlea metatarsi II is more prominent; incisura intertrochlearis lateralis is narrower. Orthonyx kaldowinyeri differs from
ALCHERINGA O. temminckii by having a shallower fossa metatarsi I and trochleae metatarsorum III and IV of about equal distal extent. It differs from O. spaldingii in having an incisura intertrochlearis medialis that extends further proximally, and the medial depth of the shaft is relatively shallower. Description and comparison. All observations in the following descriptions were made on the basis of direct comparison with specimens of O. spaldingii and O. temminckii (see Materials and methods). In proximal view, the cotyla medialis protrudes much further dorsally than the cotyla lateralis. Also in proximal view, the cotyla lateralis is dorsally protuberant of the eminentia intercotylaris, forming a notch between the cotyla and the eminentia. In O. kaldowinyeri, the sulcus ligamentosus is shallow, but in O. spaldingii and O. temminckii it is deep. The muscle attachment scar on the medio-plantar corner of the cotyla medialis protrudes only slightly beyond the cotyla. The plantar section of the hypotarsus is missing. In dorsal view, the cotylae have equal proximal extent. The eminentia intercotylaris is very low; its proximal extent is less than the length of the tuberositas m. tibialis cranialis. There is a deeply incised fossa at the base of the eminentia intercotylaris. The fossa infracotylaris is deeper than in O. temminckii (Fig. 1A, B). Within the fossa, the lateral foramen vasculare proximale is situated laterally of the tuberositas m. tibialis cranialis. It is twice as long as the medial foramen vasculare proximale, which is adjacent to the proximal margin of the tuberositas m. tibialis cranialis. In QM F56329, there is some breakage within the fossa infracotylaris dorsalis proximal to the lateral foramen vasculare proximale. The tuberositas m. tibialis cranialis is low and not as elongate as in other passerines. It begins immediately distal to the arcus extensorius. The arcus extensorius is proximo-distally shorter than wide, but not as proportionally short as in O. temminckii and O. spaldingii. Its alignment to the shaft long axis is at a slightly more acute angle than in the extant logrunners, in which the arcus extensorius is at a perpendicular angle (Fig. 1A, B). The groove for the M. extensorium digitorum longus tendon is very deep. The sulcus extensorius is deeper than in O. temminckii (Fig. 1A, B). The medial depth of the shaft is shallow at about level with the tuberositas m. tibialis cranialis, then becomes broader distally (Fig. 1G). The impressio lig. collateralis medialis is very low and elongate. The fossa parahypotarsalis medialis is moderately shallow. The lateral shaft surface is more concave than that in O. spaldingii. The crista plantaris lateralis is damaged in QM F56329 and QM F56330. In O. temminckii (Fig. 1F) and O. spaldingii, an ossified tendinal bridge connects the crista plantaris lateralis to the hypotarsus and encloses a large foramen (‘peroneal foramen’ in Orenstein [1977]). This ossified tendinal
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ALCHERINGA bridge is present in only a few passerine groups, e.g., Pomatostomidae, Climacteridae (see Manegold 2008, Worthy et al. 2010). The tub. m. fibularis brevis, which in Orthonyx is enlarged and proximally protuberant, is also broken off in the fossils. Unlike in other passerine groups, the impressio lig. collateralis lateralis is low, very large and elongate in Orthonyx species, being flat in O. spaldingii and O. temminckii (Fig. 1F) but slightly elevated in O. kaldowinyeri (Fig. 1E, 2C). In plantar view, the crista plantaris medialis is proximally a sharply developed ridge that merges with the crista medialis hypotarsi, but further distally becomes less prominent. Immediately distal to the hypotarsus, the sulcus flexorius is moderately deep and becomes shallower distally. The muscle scar on the medio-plantar corner of the cotyla medialis is wider than long in O. kaldowinyeri (Fig. 1C) and O. spaldingii, but is more elongate in O. temminckii (Fig. 1D). The distal end is broad and bends slightly plantarly. In dorsal aspect, there is a shallow groove between trochleae metatarsorum II and III that extends proximally on the shaft surface. This groove merges with a shallow fossa that has a few small foramina situated medially of the foramen vasculare distale (Fig. 1A, 2D, F, H, J). The plantar shaft surface of the distal end is not planar but rounded. The fossa metatarsi I is very shallow and situated far proximally of trochlea metatarsi II by more than the length of trochlea metatarsi II. Distal to the fossa is a low ridge (Fig. 1C, 2) that is more prominent in the fossils than in O. temminckii and O. spaldingii. The trochleae metatarsorum are widely separated and parallel in distal view. The trochlea metatarsi II is broad and about two-thirds the length of trochlea metatarsi III. The medial rims of trochleae metatarsorum II and III protrude further dorsally than the lateral rim. In dorsal view, the medial rim of trochlea metatarsi III is distinctly larger and angled disto-medially. The trochlea metatarsi IV is narrow and widely separated from trochlea metatarsi III. It extends slightly further distally relative to trochlea metatarsi III than in O. spaldingii and O. temminckii. Orthonyx sp. cf. O. kaldowinyeri Boles, 1993 (Fig. 3) Referred material. A distal right tibiotarsus with the caudal and distal portions of the condyles broken off —QM F57239. Measurements (mm). Total length >10.3; distal width ca 4.4; depth of condylus medialis >3.7; depth of condylus lateralis >3.4. Occurrence. Golden Steph Site, Riversleigh World Heritage Area, northwestern Queensland, Australia. Golden Steph Site is tentatively assigned to lower Faunal Zone C (early middle Miocene) until more taxa from this site are recovered and studied to allow confident biocorrelation (Archer et al. 1997, Worthy & Boles 2011).
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Identification. We refer the fossil tibiotarsus to Orthonyx sp. cf. O. kaldowinyeri because it falls within the expected size range of O. kaldowinyeri, as represented by the holotype femur QM F16867 and the described fossil tarsometatarsi. Because of its preservation, however, we acknowledge that this specimen can not be assigned to O. kaldowinyeri with confidence until more is discovered about this species. The fossil differs from O. spaldingii because it is much smaller in size, has an epicondylus lateralis that is larger and more protuberant, and an incisura intercondylaris that is slightly deeper. It differs from O. temminckii in that the bony ridges of the retinaculum m. fibularis are less developed, the ridge on the trochlea cartilaginis tibialis is more prominent (Fig. 3C, D) and the epicondylus medialis is more pronounced. The distal width of the fossil tibiotarsus is less than the average distal width for O. temminckii specimens measured (Table 1), but similar in size to the smallest representative of O. temminckii examined. The fossil is distinguished from all other Orthonyx species in that the pons supratendineus is slightly longer relative to its width, the bony ridges of the retinaculum m. fibularis are shorter, the difference in the widths of the condyles is more pronounced, and the lateral tuberositas retinaculi extensoris is flat (Fig. 3A). Description and comparison. In cranial view, the medial tuberositas retinaculi extensoris is low and elongate. Its distal portion is level with the proximal portion of the lateral tub. retinaculi m. fibularis. The lateral tuberositas retinaculi extensoris is also elongate but much less prominent than its medial counterpart. This tuberositas is located on the lateral portion of the pons supratendineus and does not extend beyond the proximal margin of the pons (Fig. 3A). The tuberositates retinaculorum extensorium are close together, being only separated by less than the length of the medial tuberositas. The sulcus m. fibularis is very shallow. The bony ridges of the retinaculum m. fibularis are very low, short and set widely apart. The proximal margins of the condyles are about level with the distal margin of the pons supratendineus, whereas in some specimens of O. temminckii (AM O.64813, QM O25471) they are slightly distal to the pons. The width of the condylus lateralis is about 1.5 times that of the medial condyle. In distal view, the condylus medialis projects further cranially than the condylus lateralis. There is breakage to the distal and caudal portions of the condylus lateralis, and to the caudal portion of the condylus medialis. The incisura intercondylaris is wide and moderately deep, although the distal part of the incisura is damaged. In cranial view, the distal profile of the incisura intercondylaris in O. temminckii (Fig. 3B) and O. spaldingii is very asymmetrical, with the lateral portion of the incisura being more proximally excavated than the medial
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Measurements of tarsometatarsus and femur
Fig. 4. Log-ratio diagram of measurements (tarsometatarsus and femur) of Orthonyx kaldowinyeri and extant Orthonyx species (O. spaldingii, O. temminckii). The standard taxon is Dasyornis broadbenti. The shaded areas represent the log maximum and log minimum values of landmark measurements for each taxon. Measurements on the horizontal axis are as follows. Tarsometatarsus: 1, total length; 2, proximal width; 3, proximal depth; 4, least shaft width; 5, distal width; 6, trochlea metatarsi II depth; 7, trochlea metatarsi III depth; 8, trochlea metatarsi IV depth. Femur: 9, total length; 10, proximal width; 11, proximal depth; 12, depth of caput femoris; 13, mid-shaft width; 14, mid-shaft depth; 15, distal width; 16, depth of condylus lateralis; 17, depth of condylus fibularis; 18, depth of condylus medialis.
portion. The epicondylus lateralis is a large, elongate and prominent protuberance (Fig. 3E). The epicondylus medialis is also large and pronounced, but is rounded and not as protuberant as the epicondylus lateralis. Viewed caudally, only the proximal sections of the trochlea cartilaginis tibialis and cristae trochlearum are preserved. The trochlea cartilaginis tibialis has a low ridge that is situated laterally of its medio-lateral midpoint (Fig. 3C). Comparison of Orthonyx kaldowinyeri with extant Orthonyx species The proportions of the fossil tarsometatarsi, as captured by the log ratio diagram (Fig. 4), are congruent with those observed for O. spaldingii and O. temminckii, supporting their referral to Orthonyx. The fossils overlap with the minimum size limit observed for O. temminckii. The fossil tarsometatarsi and tarsometatarsus of Orthonyx temminckii differ from those of O. spaldingii in having a proportionately narrower distal width and shallower trochlea metatarsi IV. Orthonyx temminckii is shown to have a proportionately larger least shaft width than O. spaldingii, although the least shaft widths differ markedly between the one male (AM O.3515) and two female (AM O.67652, QM O27876) specimens of O. spaldingii measured. The fossil tarsometatarsi differ from those of extant logrunners in having a proportionately shallower trochlea metatarsi II. When we removed the juvenile fossils from the log-ratio diagram, the size range was substantially reduced but the patterns in shapes changed only slightly. The distal width of the fossil tarsometatarsi was slightly larger proportionately, as in O. spaldingii, and the depth of trochlea metatarsi
III was slightly deeper relative to that of trochlea metatarsi IV, as in O. temminckii. The log-ratio diagram also shows that the relative dimensions of the O. kaldowinyeri holotype femur are very similar to those observed in the extant species. Minor differences between the species include a slightly narrower mid-shaft width in O. temminckii compared with those of O. spaldingii and O. kaldowinyeri. The femur of O. kaldowinyeri is relatively smaller and has a shallower condylus medialis than those of O. spaldingii and O. temminckii. The patterns for the fossil tarsometatarsi show a similar size depression to those of the O. kaldowinyeri holotype femur from extant taxa, and similar relative proportions to tarsometatarsi of the extant taxa. Together this supports the conservative referral of the fossil tarsometatarsi to O. kaldowinyeri, which can be confirmed with discovery and study of additional fossil material. When we used an alternative standard taxon, Pomatostomus temporalis, to construct the log-ratio graph, we recovered a similar pattern for the fossils (not shown).
Discussion Here we have described fossil tarsometatarsi that we refer to Orthonyx kaldowinyeri from various sites at Riversleigh that sequentially span the late Oligocene to early late Miocene. These fossils provide new osteological data on this taxon, and expand its temporal range, which was previously known only from the middle Miocene (Boles 1993). The O. kaldowinyeri tarsometatarsus from Hiatus Site (QM F30244), which is interpreted to be late
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ALCHERINGA Oligocene in age, represents the geologically oldest record of the Orthonychidae. We have also described a distal tibiotarsus referred to Orthonyx sp. cf. O. kaldowinyeri from the middle Miocene of Riversleigh, which was contemporaneous with the O. kaldowinyeri holotype femur. Orthonyx kaldowinyeri was smaller than the living O. spaldingii and O. temminckii and the Pleistocene O. hypsilophus (see Baird 1985) and O. wakefieldi (T. Park, pers. comm.). To date, the fossil specimens of Orthonyx kaldowineri include the oldest Cenozoic crown group passerine material described from Australia. This is the third pre-Quaternary representative of the Australo-Papuan basal oscine radiation. Other fossils of this ancient radiation include a carpometacarpus of the lyrebird Menura tyawanoides (see Boles 1995) from Riversleigh’s Faunal Zone B assemblages, which are interpreted to be early Miocene in age. Tarsometatarsi of honeyeaters (Meliphagidae) were also reported from Faunal Zone C (middle Miocene) assemblages and the Pliocene Rackham’s Roost Site at Riversleigh (Boles 2005) (see Systematic palaeontology section for explanation of Faunal Zones). The logrunner material described here is significant because it provides a late Oligocene minimum age for the divergence between Orthonychidae and Pomatostomidae. Recent molecular studies show that these two families form a sister clade to the crown group oscine assemblage (e.g., Hugall & Stuart-Fox 2012). Therefore, the new material can be used to calibrate molecular estimates of divergence times within the basal oscine radiation. Morphology and ecology To support the unusual sideways sweeping action of the legs, logrunners require specialized hind limb musculature and increased stabilization of the ankle (reviewed by Schodde & Mason 1999). This is also necessary for the supporting leg during scratching, which can be flexed such that the tarsometatarsus lies flat on the ground (Zusi 1978). Their hind limb myology has yet to be described, but the following structural features of the distal tibiotarsus and proximal tarsometatarsus may be associated with stabilization of the ankle during sweeping. The distal tibiotarsus tentatively referred to Orthonyx kaldowinyeri and those of other logrunners have an enlarged, protuberant epicondylus lateralis, which provides greater surface area for attachment of the lig. collateralis lateralis. The tarsometatarsi of Orthonyx kaldowinyeri, O. spaldingii and O. temminckii (see Description) have an enlarged impressio lig. collaterale lateralis, which similarly provides a larger surface area for attachment of this collateral ligament. These give increased structural support to the logrunner’s ankle for sweeping aside leaf litter in wide lateral arcs of 90° or more (Zusi 1978). Beneath the arcus extensorius, there is a very deep passage for the tendon of M. extensor digitorum longus, which flexes the tarsometatarsus upon the tibiotarsus. This deep
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groove may serve to hold the tendon deeper into the bone surface to prevent its displacement during vigorous sweeping. The unique foraging strategy of modern representatives of the Orthonychidae appears to have been utilized since at least the early Miocene and has brought about some of the diagnostic features of this family. Because of the specific habitat requirements of modern species, logrunners are useful environmental indicators. Extant logrunners are sedentary birds that inhabit areas with dense canopy, low light intensity and an accumulation of leaf litter (Boles 2007). Although logrunners are primarily found in rainforest, O. temminckii has been recorded in non-rainforest areas that fulfil their habitat requirements. Orthonyx temminckii has often been recorded in areas with low, dense thickets of non-rainforest plants, such as blackberry Rubus fruticosus Linnaeus, 1753a and lantana Lantana camara Linnaeus, 1753b, and very little canopy (e.g., McNamara 1937, Keast 1944, Boles 2007). Rare occurrences of O. temminckii in open eucalypt forest with an understorey of rainforest species have also been recorded (Slater 1995). The palaeohabitat of the late Pleistocene Green Waterhole Cave, from where Orthonyx hypsilophus is derived, is interpreted to be Casuarina-dominated woodlands (Dodson 1975, Baird 1985). Baird (1985) suggested that O. hypsilophus inhabited Melaleuca Linnaeus, 1767 thickets that bordered waterways, which would have provided sufficient cover. Altogether, this suggests that logrunners are not strictly rainforest specialists. The surviving relictual species of the Orthonychidae live in habitats that do not reflect the much more diverse habitats utilized by family members in the past. Palaeoecological implications Given the habitat structure requirements of extant logrunners, the presence of fossil logrunners in the Riversleigh deposits provides clues to the palaeoenvironmental structure at the time. The early Miocene Faunal Zone B and early middle Miocene Faunal Zone C assemblages are interpreted to represent a closed wet forest community because of their very high species diversity, abundance of sympatric arboreal folivores and frog communities, and lack of grazers (e.g., Archer et al. 1989, Black et al. 2012). These assemblages also include taxa that are characteristic of rainforest communities, such as lyrebirds, musky rat-kangaroos, striped possums and cuscuses (ibid). The holotype femur of Orthonyx kaldowinyeri is from Last Minute Site, which is part of the lower Faunal Zone C assemblage (Boles 1993). The new record of logrunners from sites representing these Faunal Zones does not preclude this palaeohabitat reconstruction. Initial palaeohabitat interpretations for the late Oligocene Faunal Zone A assemblages were not too dissimilar from that of Faunal Zone B (Archer et al. 1989,
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1997). Palaeoecological studies of the mammalian fauna suggested that the Faunal Zone A environment was heterogeneous with forest components (Myers 2002, Bassarova 2005). Cenogram and body mass distribution analyses of this fauna, however, suggest an open forest environment (Travouillon et al. 2009). Faunal Zone D assemblages have been interpreted to inhabit a drier, more open forest environment, with the gradual fragmentation and disappearance of rainforests during the late Miocene (Myers et al. 2001, Travouillon et al. 2009). However, extant logrunners can persist in small, isolated patches of suitable habitat within a larger expanse of a less suitable environment. Examples of these include wet gullies, riverine areas and small remnant patches in clearfelled forest (Howe et al. 1981). The discovery of Orthonyx kaldowinyeri in Riversleigh’s Faunal Zone A and D sites indicates the presence of sufficiently thick vegetation and leaf litter in these palaeoenvironments. Accordingly, we suggest here that the palaeoenvironments of these sites were structurally heterogeneous, which would have provided suitable habitat for logrunners, and supports Myers’ (2002) and Bassarova’s (2005) interpretation of Faunal Zone A sites. A heterogeneous palaeoenvironment may explain the presence of taxa characteristic of both closed and open forests in the Encore local fauna (Myers et al. 2001, Brewer et al. 2007, Travouillon et al. 2009). This lends further support to Baird’s (1985) hypothesis that logrunners were ecologically more diverse in the past than is currently the case.
Acknowledgements The authors are grateful to Jaynia Sladek (AM), who kindly provided access to comparative material and provided JMTN with a workspace. We also thank Wayne Longmore (NMV), Heather Janetzki (QM), Robert Palmer and Leo Joseph (ANWC) for facilitating access to additional reference material, and Travis Park (NMV) for measurements of Orthonyx wakefieldi. Thanks to Anna Gillespie (UNSW) for skilled preparation of specimens. Research at Riversleigh is supported by the Australian Research Council (LP100200486, DP1094569, DP130100197 DE130100467 grants to MA, SJH and K.H. Black at UNSW); XSTRATA Community Partnership Programme (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 & Margaret Pettit; the Carpentarian Land Council and the Waanyi people of northwestern Queensland. We are grateful for the field assistance of many volunteers at Riversleigh, as well as staff and postgraduate students of UNSW. JMTN is supported by grants from the Linnean Society of New South Wales (Betty Mayne Scientific Research Fund for Earth Sciences), BirdLife
ALCHERINGA Australia (Stuart Leslie Bird Research Award) and the CREATE Fund. Thanks to Sue Lindsay (Scanning Electron Microscopy and Microanalytical Unit, AM) for assistance with photographing specimens. Thanks to Simon Ho for providing helpful comments on an earlier draft of this manuscript. We thank the editors and reviewers Gerald Mayr and Storrs Olson for their constructive feedback.
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