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Home , Afroaves, Australaves, Bridled honeyeater, Carterornis, Charles Sibley, Chestnut teal

Journal & Proceedings of the Royal Society of , vol. 150, part 2, 2017, pp. 220–231. ISSN 0035-9173/17/020220-12

A guide to the and classification of Australian in 2017

Leo Joseph Australian National Wildlife Collection, National Research Collections , CSIRO GPO Box 1700 Canberra ACT 2601 Email: [email protected]

Abstract This article was first published in ‘The Australian Guide’ (CSIRO Publishing, 2017). It is reproduced here with minor updates and modifications. Scientific names of individual are listed in an Appendix except where pertinent to points in the main text.

he recently published Australian Bird related to each other that really animates their TGuide (Menkhorst et al., 2017) comes conversation. Chats are . “Well, at a time of tremendous and ongoing change yes, I suppose that’s nearly been suggested,” in our understanding of bird evolution and the 1975 worker says. Owlet-nightjars next how the classifications we adopt can best to swifts. “Surely, you modern people have summarize that understanding. been misled!” next to and Imagine two Australian ornithologists, not with other diurnal birds of prey. “Now one from 1935 and one from 1975, find- you’re delirious!” close relatives ing themselves perusing the new field guide of lorikeets, the Scrub-tits of whitefaces? in 2017 when this is being written. Both “Madness!” Shrewder, well-informed orni- recognize most but not all of the birds in it. thologists of any era would wryly note that Neither recognizes the Eungella hints of these relationships were often seen (but that 1962 photograph in Emu of a sup- in behaviour, , nests, , moults posed now makes more and . sense to our 1975 friend). The Grey Grass- The point here is that a revolution in wren, officially described in 1968 and thus , especially in systematics (the unknown to our 1935 colleague, was found scientific study of relationships and how to in 1975 in beyond where it classify according to those relationships), was first discovered around the western parts began in the late 1960s and today shows of the New South Wales- border. little sign of stopping because of the produc- Western Whipbirds, thought in 1935 to be tion of new field guides, such as the 2017 heading to in Western Australia, Bird Guide. Arguably, American ornitholo- had been confirmed in eastern Australia just gist Charles Sibley started it by studying bird two years earlier in 1933. systematics through molecules, first proteins Both our ornithologists can see how bird then DNA, and especially at the higher names have changed and find much to talk taxonomic levels of order and family. Since about in this book, but it’s the ways in which the 1980s, techniques to read and analyse we now understand groups of birds to be DNA sequences have improved such that

220 Journal & Proceedings of the Royal Society of New South Wales Joseph — A guide to the evolution and classification of Australian birds in 2017 we can now sample DNA from most of the At the tree’s trunk, living birds divide genome – the full complement of DNA in into Palæognathæ (ratites, ) and a cell. DNA provides new frameworks with Neognathæ (all others). The palæognaths which to understand how species are related continue to surprise. The flighted tinamous to each other in the avian evolutionary tree of the Americas are more closely related to of life, or phylogeny. A well-supported phyl- the extinct among the ogeny helps us assess how any aspect of bird flightless ratites (emus, cassowaries and so biology has changed during the various proc- on) than some of the latter are to each other. esses of evolution. Kiwis appear closer to the extinct elephant Charles Sibley placed the birds of Aus- birds of Madagascar (Mitchell et al., 2014). tralia front and centre in this global, orni- A corollary is that flight must have been lost thological revolution, along with those of multiple times in the evolution of palæog- and New Zealand (Sibley and nathous birds. Ahlquist, 1985). In looking at why this The Neognathæ, in turn, branches into happened, I hope the reader of today, if not the Galloanseres (waterfowls and chicken- our imaginary 1935 and 1975 friends as like birds) and the (all other birds). well, will embrace three ideas. First, science Research in the study of whole genomes constantly refines our understanding of the in 2015 challenged the view suggested by avian and how we use clas- similar research from 2014. It is fair to say sification from the Class ‘Aves’, right down that much of this debate centres on how, to the level of the species, to summarize that in the absence of a complete fossil record, understanding. There are mis-steps along the our genomic technologies can recover from way, for sure, but that is how science, and DNA any kind of signal of evolutionary people, work. Second, the 2017 Guide and events that happened a very long time ago. its successors should look very different from It is also fair to remind ourselves that while each other and from their predecessors in it is always tempting to think of a new study the species and groups they recognize. Third, of avian relationships as being the best or research in systematics can enliven the way final word on a topic to date (and maybe it one observes any bird. When observing a is), the next study will likely differ (and it bird, we are looking at the latest steps in did — see Burleigh et al. 2015!), however ongoing and open-ended evolution. That slightly, but, again, that is how science works. makes things far more interesting than if our So, while it is reassuring that the composi- understanding of the birds and the names tion of most of the major groups of birds we use all just stood still. seems to be settled, there is still uncertainty as to where some of them fit on the avian The Big Picture evolutionary tree relative to each other. For Observing a community of birds is also akin example, it is now not debated that swifts, to looking at different branches of the phy- nightjars, owlet-nightjars, and logenetic tree of birds. A brief summary of form a natural evolutionary our current understanding of the avian tree grouping. Research published at the end of of life from the roots to the tips will be help- 2014 suggested that that group is embedded ful (see details in Jarvis et al., 2014; Prum in the Neoaves whereas Prum et al., (2015) et al., 2015).

221 Journal & Proceedings of the Royal Society of New South Wales Joseph — A guide to the evolution and classification of Australian birds in 2017 placed it as the closest living relative of all South America’s three species of other Neoaves. Other differences are appar- being on another and then all other living ent among the two recent genomic studies anseriforms essentially making up a third but I am struck by the commonalities more and final “very bushy” branch. Next, Aus- than the differences. For example, Jarvis et tralia’s four smallest rails, the White-browed, al. (2014) recognized that most Neoaves are Spotted, Spotless and Baillon’s Crakes, far in what they called the , which has from being a cohesive evolutionary group, several main lineages, the two main ones apparently represent three different lineages being so-called “core” landbirds () (Garcia-R. et al., 2014). The White-browed and waterbirds (Aequornithia). The Tellu- appears to be most closely related to a simi- raves branches into (, larly odd African bird, the Striped Crake, parrots, falcons, South American ) and the bush-hens. The Spotted is closest to a and (kingfishers and relatives, , handful of similar Porzana species worldwide, eagles, woodpeckers, hornbills, ). whereas Spotless and Baillon’s are on a differ- Prum et al. (2015) retained the composi- ent branch as their own closest relatives. This tion and structure of the Telluraves, for is why they have recently been assigned to a example, especially its two enormous com- different , , which is not very ponent groups together in the same pattern closely related to Porzana. It also reminds us of relationships but differs from the earlier that a “body plan” like that of small crakes work in how the ever-mysterious South may not always be a good indicator of who American Hoatzin is related to them. I can is most closely related to whom. live with that kind of debate! Resolution of Parrots and passerines (the latter loosely these debates will depend on how well we termed ‘perching birds’), which turn out can analyse any signal of the deep evolu- to be each other’s closest relatives in a phy- tionary past of birds that is present in their logenetically surprising result, yielded still genomes. further phylogenetic surprises. In evolution, the first branching point led to A closer look New Zealand wrens in one lineage and all Considering some neognathous birds can other passerines in the other. Similarly in deepen one’s appreciation of Australia’s role parrots, the first branching point led to New in bird evolution. (mound- Zealand’s , kea and kakas in one line- builders, such as the familiar Australian age and to all other parrots in the other. Our Brush-), Plains-wanderer and Magpie understanding of passerine evolution has Goose each have their closest living phylo- advanced steadily to the point where remem- genetic connections in South America, the bering the detail of what we have learned is first being most closely related to a formidable task. In essence, the lineage in and guans, the second closest to seedsnipe, passerines that led to all species other than and the third to the marvellously named New Zealand wrens subdivided into sub- screamers. Indeed, the when oscines (represented on mainland Australia so considered is a very special anseriform only by pittas), and oscines or . bird (, geese, swans). It sits on its own The deepest lineages of the oscines are in branch in the anseriform phylogenetic tree, Australia and New Guinea (Australo-Papua),

222 Journal & Proceedings of the Royal Society of New South Wales Joseph — A guide to the evolution and classification of Australian birds in 2017

Figure 1: This figure is reproduced from a paper published in Science in 2014 (Jarvis et al. 2014). It is a summary or hypothesis of the relationships among the major groups of the world’s birds based on an extensive dataset of avian genomes. While not the final word in pattern of relationships, the different colours used for the names of different groups indicate various aspects of avian biology and how many times they may have evolved independently. Light green names, for example, applied to oscine passerines, parrots and hummingbirds show that the biological mechanisms for learning of vocalizations have evolved independently at least twice, and red names show that predation has evolved twice.

223 Journal & Proceedings of the Royal Society of New South Wales Joseph — A guide to the evolution and classification of Australian birds in 2017 this being our current understanding of a the 2017 Guide? Examples will show that, in finding Sibley first reported in 1985 (Sibley essence, much the same reasons apply. and Ahlquist, 1985). Examples include so Since 1975, several generations of ornithol- many birds characteristic of Australia (and ogists have become accustomed to some 20 New Guinea) that have no very close relatives species of honeyeaters making up the genus in the Northern Hemisphere despite some . Not until 2011 did the first having similar sounding English names: lyre- of several DNA studies tackle this and find birds, scrub-birds, bowerbirds, treecreepers, that Lichenostomus in Australia was made up whipbirds, logrunners, Australo-Papuan of at least seven different groups or lineages babblers (to distinguish them from scimitar (Nyari and Joseph 2011; Joseph et al., 2014); babblers and their allies), fairy-wrens, hon- the total number is eight, now that one key eyeaters, acanthizids (thornbills, gerygones New Guinean species, formerly known as and allies), whistlers, , butch- Oreornis chrysogenys, has finally been included erbirds, and more. Mostly beyond Australia, in DNA studies (Marki et al., 2017). Further, an even larger group of passerine species, at least six generic names were needed for the recently termed the Passerides, evolved and Australian species. Why? Five of the seven mostly in the Northern Hemisphere. One Australian groups were scattered throughout of its earliest branches, however, is the line- the full honeyeater phylogeny and were not age of Australo-Papuan robins, and could each other’s closest relatives, and so needed even be largely descended from an ancestor five generic names. The remaining two were of that group itself. Some Passerides have indeed each other’s closest relatives. They later (‘secondarily’) returned to and radiated could validly be placed in a single, sixth within Australo-Papua, grassfinches being genus or divided into a sixth and a seventh. a fine example. The true thrushes, white- The argument was made that because they are eyes, swallows and martins, reed-warblers, such distinctive lineages which diverged sev- cisticolas, and grassbirds are all in this cat- eral million years ago, assigning them to two egory. Many other Passerides are only con- genera and (“plumed” and vergently similar to birds in the Australian “fasciated” honeyeaters, respectively) made region. Thus, treecreepers and creepers are sense. Consider, for example, how different not closely related to each other, and fairy- are the “plumed” Yellow-plumed and “fas- wrens are not at all close to true wrens, for ciated” Singing Honeyeaters. The habit of example. The 1935 worker especially might using a broad Lichenostomus is dying hard, need to sit down at this point. but die it must. Incidentally, the New Gui- nean species that was last to be included in Down to genus DNA studies belongs in the genus Micropti- We have seen that at higher levels of bird lotis, itself split from (Joseph et al., classification, research most often clarifies 2014; Marki et al., 2017). And, last but not which clearly defined group is related to least, two species remain in Lichenostomus. which other clearly defined group, rather One is the species on which Lichenostomus than altering the membership of the groups was based, the Purple-gaped Honeyeater L. themselves. What of the genus-level? Why so cratitius, and the other is its closest relative, many perhaps unfamiliar generic names in the Yellow-tufted Honeyeater L. melanops.

224 Journal & Proceedings of the Royal Society of New South Wales Joseph — A guide to the evolution and classification of Australian birds in 2017

Still with honeyeaters, three species having the end of species-level for Aus- black-and-white (Pied, Black, tralian birds. Modern research documents Banded) were long placed in Certhionyx. the complex evolution still occurring at the They are not each other’s closest relatives and species-population level, within and among must be classified in three different genera species. We interpret this through ‘lenses’ (Joseph et al., 2014). of different biological characters (DNA Monarch flycatchers all look like mon- sequences, plumage, vocalizations, and arch flycatchers, so they belong in the genus more). Interpretation through one such lens . Right? No. DNA showed that coupled with one of many ways of defining a Monarcha, like Lichenostomus, fell into species might suggest two populations share groups or lineages of species that are not all a gene pool and that they should be treated each other’s closest relatives (Andersen et al., as one species. Another lens and another 2015). So, again, Monarcha is restricted to way of defining a species might suggest that the lineage that includes the single species on they are well and truly different lineages in which Monarcha was first based, the Black- the avian phylogeny and that two or more faced Monarch M. melanopsis. Monarcha, species should be recognized. Reconciling then, is more closely related to Pacific island these different lenses using a taxonomic genera such as Chasiempis and than system devised before Charles Darwin is to other Australian and New Guinean spe- akin to feeding rocket fuel to a horse pull- cies formerly placed in Monarcha. So, those ing a cart: things may collapse! By another other species need other genera and they analogy, seeing present-day diversity through now fall into two genera, two different lenses of biological characters and . And, finally here, the Aus- is akin to slicing a cake two different ways. tralian Magpie- and its close relative the Both have their validity but which will we Torrent-lark of New Guinea are a part of the follow? radiation. Fortunately, one category of problem The importance of all this is not that is simplest to resolve and again hinges on names have changed — that’s the least of ‘tree-thinking’. Until 2010, the White-naped it. It is that through systematics we grow to Honeyeater lunatus was thought appreciate the spectacular evolution of even to comprise eastern Australian M. l. lunatus our most familiar birds. Systematics directs and south-western Australian M. l. chlorop- taxonomic changes, the details of which are sis. DNA shows the eastern birds are more governed by the rules of zoological nomen- closely related to the Tasmanian endemic clature. All of this enhances what you see Black-headed Honeyeater M. affinis than to when observing a bird! the western birds (Toon et al., 2010). We infer that the white nape-band was present The species level — higher hanging in the ancestor of Melithreptus honeyeaters taxonomic fruit but has been lost in M. affinis. We could treat Discovery of the , the all three as one species (sensible? you decide!) last unquestioned species of Australian bird or the western birds must be a separate, third to be discovered and scientifically described species, M. chloropsis. In this case, the DNA (Longmore and Boles, 1983), did not mean evidence has in effect argued that it is time to

225 Journal & Proceedings of the Royal Society of New South Wales Joseph — A guide to the evolution and classification of Australian birds in 2017

correct and update an earlier way of thinking just ”? These seem thornier, not based on similarities and differences. so much because the various birds are again In Golden Whistlers, Pachycephala pecto- geographically isolated from each other, ralis, separate populations in Western Aus- but because debate about how many spe- tralia and, mostly, South Australia charac- cies there are has never really settled on any terized by cinnamon-bellied females were one prevailing view. Why? I suggest this is assigned to the subspecies Pachycephala pec- partly because there is often an unspoken toralis fuliginosa. The significance of subtle undercurrent of thinking in cases like the differences between them in plumage had -tits concerning how we should inter- been debated since the 1950s. DNA shows pret similarities and differences. That is, to that those two isolated or at best tenuously us they look so similar that the notion of connected populations are not each other’s them being separate species seems harder to closest relatives. Further, the eastern (mostly digest despite any differences and similari- South Australian) populations of “P. p. fuligi- ties in vocalizations or behaviour. If some nosa” are not genetically separate using the other differentiating character appears, such markers studied to date from other eastern as vocalizations, coupled with whether their Australian populations (Andersen et al. 2014; ranges overlap naturally or not, two species Joseph et al., 2014). Most critical of all, the may become accepted. Think of the very western populations of “P. p. fuliginosa” are similar-looking Chirruping and Chiming likely more closely related to another spe- Wedgebills, which sound so very different cies of whistler altogether and so may not and occupy different habitats where they even be the closest relative of eastern Aus- approach each other geographically that their tralian Golden Whistlers at all. Again, the recognition as two species now goes unre- cinnamon-belly of females in south-western marked. DNA evidence supports this (Toon and some parts of south-eastern Australia et al., 2013). Critically needed research on may be an ancestral character lost in some, isolated populations like the shrike-tits won’t but not all, present-day populations. Taxo- change the reality of their existence: bird- nomically, the south-west Western Austral- ers should want to see and hear them all! ian populations must become a separate spe- As with the Melithreptus honeyeaters and cies, which happily does have some subtly whistlers, research is needed to reveal one distinct plumage characteristics, and rules or other of a fairly small number of predict- of nomenclature dictate that it be known able patterns of relationships among them. as Pachycephala occidentalis. Further study The problem then is in interpreting patterns is needed to address how the re-defined P. among such isolated populations under the pectoralis fuliginosa mainly of South Australia Biological Species Concept, ornithology’s relates to other eastern Australian Golden dominant definition of a species since the Whistlers. mid-20th century. It requires that we venture What of cases like the Crested Shrike-tit’s to supposition at the edge of science. Does “Eastern”, “Northern” and “Western” forms it matter whether they could interbreed if where slight plumage differentiation (as we they came together, which clearly they aren’t perceive it) between isolated populations about to do? Does the degree of differentia- has long led ornithologists to say, “They are tion between some other closely related pair

226 Journal & Proceedings of the Royal Society of New South Wales Joseph — A guide to the evolution and classification of Australian birds in 2017 guide us? I ask, “Is this the best our science Silver-backed . Only in the can do today?” I hope not. Alternatively, in westernmost parts of Silver-backed’s range the case of such isolated populations about is the relevant piece of DNA still in its “pure” which there is taxonomic debate, we are Silver-backed form, and clos- interested in how they are related to each est to that of Black-backed Butcherbirds. other in the avian phylogeny, how similarities Silver-backed and Grey cannot be regarded and differences in other traits can be inter- as the same species. Hybridization and gene preted in a framework of well-understood flow has, we argue, occurred among species evolutionary relationships and, perhaps, a that are not closest relatives, that pattern of different species concept, whether gene flow relationships having been established much has ceased among them, and whether they earlier in butcherbird evolution. In other are continuing to diverge even if there is words, hybridization need not mean that gene flow, however occasional. We can test the birds involved are closest relatives. A that using data and analyses of those data classification reflecting evolutionary history that can be repeated. That is science. So, I is not achieved by making Grey and Silver- suggest that the species or subspecies ques- backed the same species because of past gene tion is interesting for very different reasons flow. A very similar example concerns the these days but, nonetheless, debate about Pale-headed, Northern, and Eastern Rosel- it won’t go away, especially where isolated las. Pale-headed and Northern are populations are concerned. I suggest that closest relatives and can be recognized as it will often be more interesting and useful two species. Genes from the Pale-headed to first examine how the isolated popula- appear to have extensively “infiltrated” East- tions are related to each other. Then we can ern Rosellas of the mainland but not Tas- ask whether some phenomenon like gene mania. Eastern Rosellas must nonetheless flow, say between two and not a third, is be regarded as a third species (Shipham et affecting their divergence. Finally, does all al., 2015, 2017). Recognition in the 2017 of that dictate an alternative approach to Guide of the Copper-backed Quail-thrush as classification? a species separate from the Chestnut Quail- An example that involves present-day iso- thrush arose from an example of this kind lated populations and past gene flow will of research into past gene flow having been help here. Debate has long been whether explored (Dolman and Joseph, 2016). Grey and Silver-backed Butcherbirds are By telling us about the phylogeny — the one or two species. We now know that evolutionary history or evolutionary foot- Silver-backed Butcherbirds are the closest print of a species or population — we can relatives of a third bird, the Black-backed learn something about biogeography: how Butcherbird. The Grey Butcherbird is, in species evolve as landscapes also evolve and turn, the closest relative of the other two. climates change. They tell us that the Tasma- Recent research (Kearns et al., 2014) gave a nian population of Eastern Ground Parrots, twist: genes from Grey Butcherbirds entered for example, still share genetic diversity with into eastern populations of Silver-backed mainland eastern Australian populations Butcherbirds, probably some 20,000 years despite current isolation by Bass Strait. The ago, and have now spread west through Western Ground , however, is weakly

227 Journal & Proceedings of the Royal Society of New South Wales Joseph — A guide to the evolution and classification of Australian birds in 2017 but consistently differentiated in plumage mitochondrial DNA, may not have “caught and shares no diversity with any eastern up” yet, as it evolves more slowly than, say, birds for the piece of DNA so far studied plumage and the genes controlling plum- (Murphy et al., 2011). Recognizing these as age differences. Certainly, in the teals and two species says that until we can demon- perhaps in the woodswallows, those genes strate genetic connections between western may well even be located on the sex chro- and eastern populations, we interpret the mosomes, whereas their pool of diversity for available data as favouring the idea (‘hypoth- mitochondrial DNA may still essentially be esis’) that they are two lineages not exchang- that of their common ancestor (Joseph et ing genes and that we should call them two al., 2006; Dhami et al., 2016). Alternatively, species. Further, we can suggest why they testing for a role of natural selection at the look so similar: strong natural selection for level of DNA itself can be critical. This camouflage to avoid predation. Indeed, the has, we believe, led to the best approach unquestionably different species, the Night to understanding some truly remarkable Parrot, is in many ways not so different in patterns of genetic diversity in the Eastern appearance, so its plumage, too, is probably Yellow Robin. Within that species, there is a under similar long-term evolutionary pres- geographically structured but extraordinarily sure. This reiterates potential traps of rely- deep genetic break in mitochondrial DNA ing on differences in plumage. Australian diversity between two groups of populations. populations of the most The magnitude of this break is more typical definitely are two genetic groups with respect of that seen between genera than within a to mitochondrial DNA, but these in no way species. We have argued, however, that it is match geographical structure in their plum- best interpreted as evidence of selection on age variation. Research is still in progress to mitochondrial DNA, and that there is no examine why this is so; the answer, I suspect, need to alter subspecies or even species-level will again involve how well we can under- classifications (Morales et al., 2015, 2017). stand intricacies of the population genetics Geographical overlap of migratory and of the species in its past. non-migratory populations also needs dis- Lessons have been learned about how entangling by field, museum and labora- important it is to understand what we might tory work. The mystery of how many spe- call DNA’s own natural history. DNA studies cies should be recognized in the Cicadabird, of two pairs of Australian birds well illustrate which by our current understanding also one potential interpretative trap. One pair occurs widely outside Australia, is a fine of species is the White-browed and Masked example (Pedersen et al., in press). Woodswallows, and the other is the Grey and I hope all this gives a taste of the complex- Chestnut Teals. What we have learned from ity of these species-level problems and why these two pairs concerns multiple species they will be around for a while yet. Each that have diverged from their most recent case will be different. Patterns of relation- common ancestor only very recently in evo- ships and what gene flow does or does not lutionary terms. It may be that we can see (or mean with regard to whether it is stopping hear) very clearly that they are distinct spe- divergence between two populations will be cies. Some of the DNA we study, especially critical. The 2017Guide is a treasure chest of

228 Journal & Proceedings of the Royal Society of New South Wales Joseph — A guide to the evolution and classification of Australian birds in 2017 the problems waiting for study. How many Eastern Platycercus eximius species are in the Purple Swamphen, Spinifex Eastern Ground Parrot wallicus Pigeon, Red-tailed Black-, Hel- Pezoporus flaviven- meted , Spectacled Monarch and tris Cicadabird, to name a few? Are the three Pol- Pezoporus occidentalis ytelis parrots (Superb, Regent and Princess) Grey Grasswren Amytornis barbatus more closely related to each other than one Eungella Honeyeater Bolemoreus hind- or two of them are to other parrots? While woodi reassuring our friends from 1935 and 1975 Bridled Honeyeater Bolemoreus frenatus that we have made strides, they’d delight in Yellow-plumed Honeyeater Ptilotula ornata reminding us of the old maxim — the more Gavicalis virescens we learn, the more questions we find. And Helmeted Friarbird Philemon buceroides that is as it should be. Scrub-tit Acanthornis magna Copper-backed Quail-thrush Cinclosoma Appendix clarum Scientific names of species mentioned but omit- Chestnut Quail-thrush Cinclosoma castano- ted in the text for clarity of reading tum Australian Brush-turkey Alectura lathami Western Whipbird recently advocated to be Plains-wanderer Pedionomus torquatus nigrogularis and Psophodes Magpie Goose Anseranas semipalmata leucogaster White-browed Crake Amaurornis cinerea Psophodes cristatus Spotted or Australian Crake Porzana flu- Psophodes occidentalis minea Crested Shrike-tit Falcunuclus frontatus Australian Spotless Crake Zapornia tabuensis Spectacled Monarch Symposiachrus trivir- (Porzana tabuensis in some texts) gatus Baillon’s Crake Zapornia pusilla (Porzana Cicadabird Edoliisoma tenuirostre pusilla in some texts) Grey Butcherbird torquatus gracilis Black-backed Butcherbird Cracticus menta- Chestnut Teal Anas castanea lis Australasian Purple Swamphen Porphyrio Silver-backed Butcherbird Cracticus argen- melanotus (Porphyrio porphyrio in some teus texts) White-browed Artamus super- Hoatzin Opisthocomus hoazin ciliosus Spinifex Pigeon Geophaps plumifera Masked Woodswallow Artamus personatus Melopsittacus undulatus Eastern Yellow Robin Eopsaltria australis Red-tailed Black-Cockatoo Calyptorhynchus Acknowledgements banksii I am very grateful to John Manger and Peter Superb Parrot swainsonii Menkhorst for their support in inviting me Polytelis anthopeplus to write the piece (Joseph, 2017) on which Princess Parrot Polytelis alexandrae the present article is based and for their per- Pale-headed Rosella Platycercus adscitus mission to modify it here. Dr Robert Marks Northern Rosella Platycercus venustus

229 Journal & Proceedings of the Royal Society of New South Wales Joseph — A guide to the evolution and classification of Australian birds in 2017 kindly suggested its appearance here and I Science, 346, 1320-1331. doi: 10.1126/ thank him for that support. J. Van Remsen science.1253451 offered comments on a draft, and I followed Joseph, L. 2017. A guide for birders to the evolution and classification of Australian some of them! birds. In Menkhorst, P., Rogers, D., Clarke, R., Davies, J., Marsack, P. & Franklin, K., References The Australian Bird Guide, CSIRO Publishing, Andersen, M.J., Nyári, A. S., Mason, I., Joseph, Melbourne, 23-29. L., Dumbacher, J.P., Filardi, C.E. Moyle, Joseph, L. and Buchanan, K. L. 2015. A R.G. 2014. Multi-locus phylogeography quantum leap in avian biology. Emu 115(1) of the world’s most polytypic bird: the 1-5. doi: 10.1071/MUv115n1_ED Pachycephala pectoralis/melanura species Joseph, L. Nyari, A. and Andersen, M. complex. Zoological Journal of the Linnean 2014. Taxonomic consequences of Society, 170, 566-588. doi: 10.1111/ cryptic speciation in the Golden Whistler zoj.12088. Pachycephala pectoralis complex in mainland Andersen, M.J., Hosner, P.A., Filardi, C.E., southern Australia. Zootaxa, 3900, 294–300. and Moyle, R.G. 2015. Phylogeny of Joseph, L., Toon, A., Nyári, Á. S., Trueman, J. the monarch flycatchers reveals extensive and Gardner, J. 2014. A new synthesis of the paraphyly and novel relationships within a molecular systematics and biogeography of major Australo-Pacific radiation. Molecular honeyeaters (Passeriformes: Meliphagidae) Phylogenetics and Evolution, 83, 118–136. highlights biogeographical complexity of a Burleigh, J.G., Kimball, R.T., and Braun, E.L. spectacular avian radiation. Zoologica Scripta, 2015. Building the avian tree of life using 43, 235-248. doi:10.111f1/zsc.12049 a large-scale, sparse supermatrix. Molecular Joseph, L., Wilke, T., Ten Have, J. and Chesser, Phylogenetics and Evolution, 84, 53-63. R.T. 2006. Implications of mitochondrial Dhami, K., Joseph, L., Roshier, D., and Peters, DNA polyphyly in two ecologically J. 2016. Recent speciation and elevated undifferentiated but morphologically distinct Z-chromosome differentiation between a migratory birds, the Masked and White- sexually dichromatic and monochromatic browed Woodswallows Artamus spp. of pair of Australian teals (: Anas). inland Australia. Journal of Avian Biology, 37, Journal of Avian Biolog,y 47, 92–102. doi: 625-636. 10.1111/jav.00693. Kearns, A., Joseph, L., Toon, A. and Dolman, G. and Joseph, L. 2016. Multi- Cook, L. 2014. Australia’s arid-adapted locus sequence data reveal Pleistocene butcherbirds experienced range expansions speciation in semi-arid southern Australian during Pleistocene glacial maxima. Nature birds (Cinclosoma spp.) was associated with Communications 5, 3994 doi: 10.1038/ increased genetic drift. BMC Evolutionary ncomms4994. Biology, 16, 226. doi: 10.1186/s12862-016- Longmore, N.W. and Boles, W.E. 1983. 0798-6. Description and systematics of the Eungella Garcia-R., J.-C., Gibb, G.C., and Trewick, S.A. Honeyeater Meliphaga hindwoodi, a new 2014. Deep global evolutionary radiation species of honeyeater from central eastern in birds: Diversification and trait evolution Queensland, Australia. Emu, 83, 59-65. in the cosmopolitan bird family Rallidae. Marki, P.Z.L., Jønsson K.A., Irestedt, and Evolution, 81, M., Nguyen, J.M.T., Rahbek, C., and 96–108. Fjeldså, J. 2017. Supermatrix phylogeny Jarvis, E.D., Mirarab, S., Aberer, A.J., Li, and biogeography of the Australasian B., Houde, P., [and 100 other authors]. Meliphagides radiation (Aves: Passeriformes). 2014. Whole-genome analyses resolve early Molecular Phylogenetics and Evolution, 107, branches in the tree of life of modern birds. 516–529.

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Menkhorst, P., Rogers, D., Clarke, R., Davies, Prum, R.O., Berv, J.S., Dornburg, A., Field, J., Marsack, P. and Franklin, K. 2017. The D.J., Townsend, J.P., Lemmon, E.M. and Australian Bird Guide. CSIRO Publishing, Lemmon, A.R. 2015. A comprehensive Melbourne. phylogeny of birds (Aves) using targeted next- Mitchell, K.J., Llamas, B., Soubrier, J., generation DNA sequencing. Nature, 526, Rawlence, N.J., Worthy, T.H., Wood, J., Lee, 569-573. doi:10.1038/nature15697. M.S.Y., and Cooper, A. 2014. Ancient DNA Shipham, A., Schmidt, D., Joseph, L. and reveals elephant birds and are sister taxa Hughes, J. 2015. Phylogenetic analysis of the and clarifies ratite bird evolution. Science, Australian rosella parrots (Platycercus) reveals 344, 898-900. discordance among molecules and plumage. Morales, H.E., Pavlova, A., Joseph, L., and Molecular Phylogenetics and Evolution, 91, Sunnucks, P. 2015. Positive and purifying 150–159. doi:10.1016/j.ympev.2015.05.012. selection in mitochondrial genomes of a bird Shipham, A., Joseph, L., Schmidt, D. and with mitonuclear discordance. Molecular Hughes, J.H. 2017. A genomic approach Ecology, 24, 2820-2837. doi: 10.1111/ reinforces a hypothesis of mitochondrial mec.13203. capture in eastern Australian rosellas. The Auk, Morales, H., Pavlova, A., Joseph, L., and 134, 181-192. doi: 10.1642/AUK-16-31.1. Sunnucks, P. 2017. Perpendicular axes Sibley, C.G. and Ahlquist, J.E. 1985. The of incipient speciation generated by phylogeny and classification of the Australo- mitochondrial introgression. Online early 19 Papuan passerine birds. Emu, 85, 1–14. April 2017. Molecular Ecology, doi: 10.1111/ http://dx.doi.org/10.1071/mu9850001 mec.14114. Toon, A., Hughes, J. and Joseph, L. 2010. Murphy, S., Joseph, L., Burbidge, A. and Multilocus analysis of honeyeaters (Aves: Austin, J. 2011. A cryptic and critically Meliphagidae) highlights the spatio- endangered species revealed by mitochondrial temporal heterogeneity in the influence DNA analyses – the Western Ground Parrot. of biogeographic barriers in the Australian Conservation Genetics, 12, 595-600. doi: monsoonal zone. Molecular Ecology, 10.1007/s10592-010-0161-1 19, 2980-2994. doi: 10.1111/j.1365- Nyari, A. and Joseph, L. 2011. Systematic 294X.2010.04730.x dismantlement of Lichenostomus improves Toon, A., Joseph, L. and Burbidge, A. 2013. the basis for understanding relationships Genetic analysis of the Australian whipbirds within the honeyeaters (Meliphagidae) and and wedgebills illuminates the evolution historical development of Australo-Papuan of their plumage and vocal diversity. Emu, bird communities. Emu, 111, 202-211. 113, 359-366. http://dx.doi.org/10.1071/ Pedersen, M.P., Irestedt, M., Joseph, L., MU13005. Rahbek, C., and Jønsson, K.A. In review. Phylogeography of a “great speciator” (Aves: Edolisoma tenuirostre) reveals complex dispersal and diversification dynamics across the Indo-Pacific. Accepted Journal of Biogeography, 4 December 2017, doi: 10.1111/jbi.13182

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