Australian Field 2021, 38, 59–65 http://dx.doi.org/10.20938/afo38059065

Challenges for research on the Spotted Quail-thrush Cinclosoma punctatum in north Queensland

Leo Joseph

Australian National Wildlife Collection, National Research Collections Australia, CSIRO, G.P.O. Box 1700, Canberra ACT 2601, Australia Email: [email protected]

Abstract. A case is made for why researchers should consider the possibilities that the north Queensland population of Spotted Quail-thrush Cinclosoma punctatum discovered in 2008 may be neither geographically or genetically isolated nor taxonomically distinct. Field and museum work are clearly needed to address these questions as well as the biology of the north Queensland population itself.

Introduction Ford 1983, and eBird (https://ebird.org/), and Atlas of Living Australia (http://www.ala.org), both accessed 29 March The Spotted Quail-thrush Cinclosoma punctatum 2021]. Blackdown Tableland has long been the northern (Passeriformes: Cinclosomatidae) is widespread in specimen-based recorded limit of C. p. punctatum and the south-eastern Australia (see Figure 1; Higgins 2002). population(s) there and nearby at Carnarvon National Park Generally, the is moderately common in suitable including its Salvator Rosa section, for example, to date habitat. It is frequently observed despite being generally also appear isolated (Figure 1; Ford 1983; Higgins 2002). inconspicuous, cryptic and terrestrial. Overall, the species is of least conservation concern (e.g. Garnett et al. 2011; The ramifications of a population of Spotted Quail-thrush BirdLife International 2017) though it is extinct at the far having been first reported in north Queensland in 2008 in western extremity of its range in the Mount Lofty Ranges an area frequently visited by many resident ornithologists (subspecies C. p. anachoreta; see Figure 1; Paton et al. warrant careful consideration. Nielsen (2015) referred to 1994; Horton et al. 2013; van Weenen et al. in press). Two sightings from the 1970s of an unidentified quail-thrush subspecies are recognised among extant populations: Cinclosoma sp. at Mount Fox, 155 km south-east of nominotypical C. p. punctatum in mainland south- Nettle Creek, the southernmost locality where the newly eastern Australia, and C. p. dovei of Tasmania (Schodde discovered north Queensland population of Spotted Quail- & Mason 1999; Higgins 2002). In 2008, an apparently thrush has been recorded (Figure 1). These would almost isolated population of the species was discovered in certainly be of Spotted Quail-thrush but nothing can be north Queensland in eucalypt woodland and forest west deduced of their subspecific status. When these sightings of Atherton–Herberton–Ravenshoe (see Figure 1; Nielsen 2015). In this article, I elaborate on matters relevant to this are coupled with what Nielsen (2015, p. 302) termed the population as a further contribution to understanding the “rather incredible” 2008 report of the north Queensland of this family (e.g. Joseph 2021). , the likelihood arises that other populations of this often cryptic, terrestrial species (Higgins 2002) exist in the relatively under-surveyed and difficult-to-access area Spotted Quail-thrush in central and broadly indicated by ‘?’ in Figure 1. More precisely, I refer north Queensland here to an area bounded in the north and south by the Herbert and Burdekin Rivers, respectively, and in the Nielsen (2015, p. 302) reported that the north Queensland west and east by Kinrara and Girringun National Parks, population of the Spotted Quail-thrush had been found respectively. This region has extensive suitable habitat “through the ranges from Ravenshoe and west through of eucalypt woodland (e.g. Figure 2 in DEWR 2007). It is the drier country to Dimbulah and Almaden” (see visited far less regularly by birdwatchers than the north Figure 1). Nine sightings have been reported on eBird Queensland sites (see the dearth of records in eBird (https://ebird.org/; accessed 29 March 2021) in 2019, 2020 (https://ebird.org/hotspots; accessed 19 March 2021). and 2021 from four localities within the range stated by Certainly, it should not be assumed that the species is Nielsen (2015). The population’s known range is therefore not there. Should more populations be found, they may ~166,600 ha in extent bounded by Almaden in the west, indicate genetic connectivity, even if sporadic, among Dimbulah in the north, Watsonville–Irvinebank in the east, and Nettle Creek and ranges west of Ravenshoe in the south populations of C. punctatum between the Blackdown (Figure 1). Not all of this area will be suitable for quail- Tableland–Carnarvon National Park area (Figure 1) and thrushes. This area is ~750–800 km north of the nearest north Queensland. Consistent with this possibility is Ford’s confirmed, known populations of the species at and (1983) observation that the species avoids wet coastal near Blackdown Tableland National Park and Carnarvon rainforest and is more a of inland habitats in the National Park in central Queensland [Figure 1; see e.g. northern parts of its range. 60 Australian Field Ornithology L. Joseph

Figure 1. Map of eastern Australia showing (1) specimen records of Spotted Quail-thrush Cinclosoma punctatum derived from the Atlas of Living Australia (www.ala.org) and modified based on Ford (1983) to exclude misidentified old specimens, (2) the approximate distributions of the three subspecies recognised by Schodde & Mason (1999), and (3) the location of the recently discovered population from north Queensland west of the towns of Herberton, Ravenshoe and Atherton. Detail in the inset shows the limits of the population’s recorded range derived from Nielsen (2015) and sightings in eBird (https://ebird.org/). Localities mentioned in the text are also shown. Note that close examination of records in Ford (1983) and the Atlas of Living Australia from the region shown here as the Blackdown Tableland suggests the possibility of a number of isolated or partially isolated populations in that region. Question marks indicate gaps in recorded range for which further survey work is warranted to look for other populations. Variation in any such populations would help clarify understanding of the north Queensland population. Abbreviations: NSW – New South Wales, NT – Northern Territory, QLD – Queensland, SA – South Australia, TAS – Tasmania, VIC – Victoria.

Taxonomy of the Spotted Quail-thrush await scientific verification and detail if only because fabrications of data related to those precise attributes have in north Queensland been argued recently in Australian ornithology (see details in Olsen 2018; Menkhorst et al. 2020; Olsen & Menkhorst The next issue I address is whether the north Queensland 2020). Notably, photographs of north Queensland Spotted population can yet be differentiated as a distinct subspecies. Quail-thrushes are online [e.g. photographs taken by In two non-peer-reviewed sources, the population has Enoch Bultreys of live male and female birds (at least been noted as different in having an “overall rufous one of each) can be seen at https://ebird.org/checklist/ appearance” (Nielsen 2015, p. 302) and in vocalisations, S60990696; https://ebird.org/checklist/S58191497; https:// nest and male (Roberts 2021). The purported ebird.org/checklist/S58187638; all accessed 29 March differences, especially those cited in the latter source, 2021]. Citing conservation concerns about the newly Spotted Quail-thrush in north Queensland 61

discovered population, Roberts (2021) reported some It is helpful to understand and parse two key issues as vehement though far from unanimous opposition in the bird- separate but related. The first issue is one of the rules of observing community to the collection of any specimens zoological nomenclature and the second is one of science to determine its taxonomic status within C. punctatum. No and scientific practice. On the nomenclatural issue, Article specimens of the population have been collected. Where 73.1.4 of the International Code of Zoological Nomenclature does this leave research into the question of whether the (ICZN 2000) allowed that, north Queensland population is taxonomically distinct? Designation of an illustration of a single specimen as a Schodde & Mason (1999) compared 24 specimens of holotype is to be treated as designation of the specimen C. p. punctatum from north of the Hunter River, a minor illustrated; the fact that the specimen no longer exists or cannot be traced does not of itself invalidate the designation. biogeographical barrier in eastern Australia (Figure 1; Ford 1987; Bryant & Krosch 2016) and 68 from south of it. They This means that a new scientific name for a species or considered the 24 northern birds to be slightly paler olive- subspecies can be erected without a conventional museum brown dorsally and to ‘gain’ more of a faint russet wash over specimen. Later, ICZN (2017) issued a series of strong the back as one moves towards the northern geographical recommendations concerning the practice of not having a limit around the Blackdown Tableland. Schodde & Mason traditional museum specimen as a holotype. These include (1999) further remarked that the more northern specimens (1) that one provide (e.g. from near Gympie and Blackdown Tableland; detailed reasoning why at least one preserved specimen, Figure 1) become washed more richly cinnamon-rufous whether a complete individual organism or a part of such over the flanks. They could not determine whether there is an individual, was not used as the name-bearing for a step in variation at the Hunter River or smooth, latitudinal the new taxon and why the formal naming of the taxon is clines in these traits. Higgins (2002) called for further needed at a point in time when no preserved name-bearing research, noting the small number of samples available type will be available (p. 96), from more northern populations. Given the need argued and (2) that when so naming a new species or subspecies, above to search for other populations between north Queensland and the Blackdown Tableland, it is entirely steps taken by an author to capture and preserve a physical specimen of the new taxon and/or locate an existing possible at this stage that the north Queensland population preserved specimen in natural history collections should be could be the end-point of a geographically fragmented recounted (p. 96) (ICZN 2017). cline, whether for morphological or genetic characters. ICZN (2017, pp. 96–97) further stated: Relevant here is the phenomenon known as isolation Whenever feasible, new species-group taxa [i.e. a species by distance (Wright 1943). It describes step-wise genetic or subspecies] should be established on the basis of at differentiation among subpopulations connected by low least one preserved type specimen. Additional information but non-zero levels of dispersal and gene flow. If sampling representing diagnostic characters (e.g. illustrations, is incomplete, subpopulations at geographical ends of the DNA sequences, audio recording analyses, etc.) should distribution can appear genetically different when in fact accompany the description of a new species-group taxon they are genetically continuous. A recent example of exactly whenever possible, but well-preserved biological specimens this problem concerns recognition of subspecies in a high- (either as complete individuals, or parts of individuals) are widely regarded as representing the most generally reliable profile species, the Chimpanzee Pan troglodytes (see means for establishing the biological and scientific basis Lester et al. 2021). Similar reasoning applies to plumage for a species-group name. Establishing new species- and morphometric characters. Apparent differences in a group taxa without preserved name-bearing type material population at a species’ geographical end-point for one is permissible under the Code, but is discouraged unless or more characters may be an artefact of inadequate justified by special circumstances, such as when capture sampling. More complete sampling, however, could show or preservation of specimens is not feasible for technical other outcomes such as the characters varying in a smooth reasons or for conservation concerns, or when specimens must be destroyed to reliably diagnose a new species. gradation, i.e. a primary cline, or where the characters are While preserving a whole organism as the type specimen is truly stepped in their variation. The taxonomic relevance preferable and encouraged, in circumstances when whole of any purported differences between the northern and organism preservation is not feasible a portion (or portions) southern Queensland populations cannot be determined of the organism sufficient to allow the new species-group until these aspects have been properly studied. taxon to be reliably diagnosed should be preserved. The second issue is whether the research underpinning Why is a blood sample or a feather not the decision to erect a new name for a species or subspecies enough? satisfactorily shows that supposedly distinctive characters and variation within the ‘new’ species or subspecies fall beyond the range of variation expected in one or more Roberts (2021) voiced the view, without documented other taxa. support, that it is no longer necessary to collect museum specimens and that a blood sample or a feather will Given concerns in some parts of the ornithological suffice to “determine the taxonomy” of north Queensland community about taking a Spotted Quail-thrush specimen C. punctatum. He presumably meant that a DNA sequence from north Queensland, one could, and in a strictly derived from a blood sample or a feather can “determine nomenclatural sense, validly introduce a scientific name the taxonomy”. Complementing the questions raised for the north Queensland population of the Spotted above, I offer the following on why this view is ill-informed at Quail-thrush. Until the questions I have posed above are best. Bates et al. (2004) also discussed why permanently addressed, however, such an action would not be backed archived voucher specimens ultimately are needed over by good science, let alone the highest possible scientific an exhaustible blood sample. standards. 62 Australian Field Ornithology L. Joseph

Some examples of new species names introduced photographs and DNA data. Escalante et al. (2018) either largely or wholly on the basis of DNA and/or without later roundly debunked the science and showed that a conventional museum specimen are presented below. the procedures used in naming this taxon bordered The examples show why ICZN (2017) at best reluctantly on fraud; they argued that the birds involved are best permits description of a new species or subspecies without considered hybrids. a traditional specimen. 5. Athreya (2006) described a new species of babbler Brower (2010) used DNA sequence data to split what (Timaliidae), the Bugun Liocichla bugunorum. Though the holotype was stated to be a bird that was had long been considered to be one variable species of released, the description of the new species met butterfly into ten species. He described how each was the requirements of the Code at the time (i.e. ICZN unique solely from DNA sequences but he sequenced 2000). Feathers from the released bird, including 466 sacrificed individuals to deduce the ten groups that rectrices showing the distinctive tail markings and he could then name as species. Most importantly, he cited one secondary flight-feather, and photographs of a an archived whole butterfly specimen (and not one wing, living bird were lodged in the collection of the Bombay or a head, or an abdomen), as a holotype for every one Natural History Society, Mumbai, . Athreya (2006) of the ten species that he named and so complied with inferred the new Liocichla to be very rare and localised, the International Code of Zoological Nomenclature (ICZN which was supported by later modelling (Peterson & Papeş 2006). Athreya (2006, p. 92) noted that steps 2000, 2017). For very sensible reasons, the Code does “will be taken” to collect a specimen if a later census not allow a species or subspecies to be named from DNA found a larger population than the “three (breeding?) without a specimen or illustration as a holotype even if the pairs” (Athreya 2006, pp. 82, 84, 87 and 92) then specimen is not a traditional museum specimen (see ICZN known. It is unclear if this has happened. 2017; Krell & Marshall 2017). Imagine that one is observing a bird in the field or even a specimen in a museum. We Viewing the case of the today and how cannot see the DNA level characters that supposedly it relates to the north Queensland Spotted Quail-thrush, distinguish a species or subspecies so how can we name I suggest four other points worth noting. First, a different what we are looking at? Brower (2010) admitted in the and much more widely distributed species of Liocichla title of his paper that naming of the ten new species of babbler that is now known not to be the Bugun Liocichla’s butterfly in this way was setting a bad precedent but he did closest relative (Mays et al. 2015) is sympatric with it, i.e. nominate a holotype specimen for each one. co-occurs with it, and this is one of the gold standards for Next, I summarise five recent cases of new species- species status. Certainly, no other quail-thrush co-occurs group names (i.e. species or subspecies) for birds being with the north Queensland population of the Spotted Quail- introduced without specimens: thrush. Second, the closest relative of the Bugun Liocichla, the Emei Shan or Grey-faced Liocichla L. omeiensis, and 1. Schodde & Weatherly (1983) named a fairy- the Bugun Liocichla replace each other geographically, wren (Maluridae), Campbell’s Fairy-wren Malurus both occurring at altitudes around ~2500 m. They are campbelli, from a photograph but a specimen was soon collected (Schodde 1984). DNA sequence data separated by ~1000 km and by the ~7500-m-high now available (Driskell et al. 2011) are inconclusive Hengduan Mountains. In contrast, we cannot be sure about its taxonomic status because of still inadequate that north Queensland Spotted Quail-thrushes are truly population-level sampling, exactly akin to what I isolated (see above). Third, the Bugun Liocichla showed have argued above for the Spotted Quail-thrush. See several key differences from that are LeCroy & Diamond (1995), del Hoyo & Collar (2016), discrete, i.e. grey vs black crown, yellow vs red loral and and Gill et al. (2021) for unresolved debate on this post-ocular spots, or even presence vs absence of these taxon. spots. In contrast, current claims suggest that the north 2. Smith et al. (1991) described a new species of shrike Queensland Spotted Quail-thrush at best ‘differs’ from (Laniidae), the Bulo Burti Boubou Laniarius liberatus, other populations in more-difficult-to-measure,continuous , using DNA from a single blood sample of an individual and possibly clinally varying characters such as “overall bird that was released. Nguembock et al. (2008) rufous appearance” (Nielsen 2015, p. 302). Finally, and Finch et al. (2016) later addressed the kinds of phylogenetic analysis of mitochondrial DNA showed that questions of variation that I have argued should be Bugun and Emei Shan Liochichlas are closest relatives addressed for the north Queensland Spotted Quail- thrush. They showed that it had been a mistake to but deeply divergent relative to divergences among other name the ‘new’ species because it was indeed part (Mays et al. 2015). of the variation within a separate species, Manda In short, describing a new species or subspecies without Boubou L. nigerrimus. The name L. liberatus is now a specimen is a last resort and, almost certainly, far from not recognised even at the subspecies level. best practice. A DNA sequence certainly tells something 3. Olsen et al. (2002) described a hawk-owl (Strigidae), of the evolutionary history and distinctiveness of that DNA the Little Sumba Hawk-Owl Ninox sumbaensis, sequence relative to any other sequence of the same primarily from DNA data. Olsen et al. (2002) published gene in any other organism. However, without a voucher photographs of an individual that had nonetheless specimen and without other sequences for context, we learn been killed. Feathers from it were registered in an nothing more of the population from which the individual academic institution museum as the holotype. It is carrying the DNA sequence came and whether individuals tragically unfortunate that apparently nobody thought or knew how to preserve the killed bird as a specimen. in the population can be seen to be different from those in other populations. An example from the same part of 4. Silva et al. (2017) described a new parrot (Psittacidae), north Queensland as being discussed here will illustrate the Blue-winged Amazon Amazona gomezgarzai, that point and show that there are many downstream from the Yucatán Peninsula, Mexico, based on uses of specimens beyond “determining the taxonomy” Spotted Quail-thrush in north Queensland 63

(Roberts 2021) of a population. Their value does not begin specimens, while of course needing to be done sensibly, and end with whether a population can be named as a new sensitively and with checks and balances (Costello et al. species or subspecies. 2016), ultimately is what underpins the scientific order we Joseph & Moritz (1993) described a White-browed try to bring to understanding, conserving and managing Scrubwren frontalis, collected in eucalypt the populations that comprise biodiversity (Remsen 1997; woodland south of Herberton, that had mitochondrial Winker & Withrow 2017; Schmitt et al. 2018). The gap DNA (mtDNA) of an Atherton Scrubwren S. keri. Without between these two views may never be fully bridged. I a voucher specimen, we would be left with the puzzling hope, however, that each time the debate arises, and each possibility of a strictly rainforest bird, S. keri, occurring in time an article such as this is written, those on one side of eucalypt country kilometres away from rainforest. Other the debate can more clearly see the views of those on the explanations such as a mix-up of samples in the field or other. the laboratory, or that this showed that S. keri individuals disperse away from rainforest, were all eliminated through Acknowledgements reference to the museum specimen from which the DNA data came. It was deduced that a female S. keri and male I gratefully acknowledge the strong support of the leadership S. frontalis must have hybridised at some time in the past, of BirdLife North Queensland and the Queensland Department presumably at the rainforest–eucalypt ecotone where of Environment and Science for the securing of one or two specimens, although none have been collected, with which to the two species’ habitats essentially abut, and that the assess the taxonomic status of the north Queensland population mtDNA of S. keri is in the local population of S. frontalis. of Spotted Quail-thrush. I thank G. Harrington, S. Murphy and The proof provided by the specimen that the two species P. Webster for showing me the population in the field in October hybridise allows us to ask other research questions. Do 2019. I also acknowledge the sincere and courteous expressions they hybridise regularly? Is it always female S. keri mating of both support and opposition put to me when I spoke to BirdLife with male S. frontalis, or does the reciprocal apply? Why North Queensland’s October 2019 meeting about securing one or do they hybridise? Does this help explain difficult field two specimens of this population. Julian Teh prepared Figure 1. I identifications of scrubwrens in the area? thank two reviewers and the Editor for very helpful comments and additional references. The research needed to resolve the key questions posed in this article (i.e. Is the population of the Spotted Quail- thrush in north Queensland truly isolated? Is it consistently References and recognisably different?) would eventually be helped not hindered from a starting point of at least one specimen Athreya, R. (2006). A new species of Liocichla (Aves: Timaliidae) from the north Queensland population of the Spotted from Eaglenest Wildlife Sanctuary, Arunachal Pradesh, India. Quail-thrush. Searches for and specimens from any Indian Birds 2, 82–94. Bakker, F.T., Antonelli, A., Clarke. J.A., Cook, J.A., Edwards, S.V., other at-present-undiscovered populations would also be Ericson, P.G.P., Faurby, S., Ferrand, N., Gelang, M., Gillespie, critical. More generally, one specimen of any taxonomically R.G., Irestedt, M., Lundin, K., Larsson, E., Matos-Maraví, P., questionable population can provide extensive and useful Müller, J., von Proschwitz, T., Roderick, G.K., Schliep, A., DNA data about the population. It is also a first datum Wahlberg, N., Wiedenhoeft, J. & Källersjö, M. (2020). The concerning whether the population is within or beyond the Global Museum: Natural history collections and the future of range of phenotypic variation (i.e. external appearance) of evolutionary science and public education. PeerJ 8, e8225. other relevant populations. Conversely, DNA sequences Bates, J.M., Bowie, R.C.K., Willard, D.E., Voelker, G. & from a taxonomically questionable population but not Kahindo, C.A. (2004). A need for continued collecting of avian anchored to a specimen are just that: their value in voucher specimens in Africa: Why blood is not enough. Ostrich rigorously positioning the population in questions of 75, 187–191. variation and taxonomy is more limited. Eventually, Bekoff, M. & Elzanowski, A. (1997). Collecting birds: The a specimen would also bring helpful dimensions to importance of moral debate. Bird Conservation International 7, conservation and management of the Spotted Quail-thrush 357–361. that can only come from the many advantages of museum BirdLife International (2017). Cinclosoma punctatum (amended specimens (for extensive reviews see Remsen 1995; version of 2016 assessment). 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