<<

AllThe in the Family Family Tree The deep roots of the parrot family tree have long been a mysterious affair. With their stocky bodies, fleshy ceres, strongly curved bills and zygodactyl feet (two toes forward and two back) the (Order Psittaciformes) are easily distinguished from all other orders of . However, the physical similarities shared across the parrots has made it difficult for taxonomists to agree on the ordering of relationships among different genera and . In some cases it has been difficult to decide whether different populations of a species represent distinct species or may simply be the result of the variation within a single species. The identification of such cryptic species is vital for effective conservation. If genetic evidence shows that a small sub-population of a widespread species is actually a distinct species, then saving this rare new species becomes a conservation priority. Such a discovery might also suggest modifications of avicultural practices by zoos and private breeders. In addition to these practical issues, resolving the evolutionary history of a group is valuable for biologists who want to better understand the evolution of the very traits that make the parrots so interesting, such as their long lifespans, colourful , keen intelligence and striking vocal abilities. by Timothy F. Wright and Erin E. Schirtzinger, Biology Department, New Mexico State University

Photo Credits: Crimson Rosella © Aaardvaark/Flickr.com, Mitred Conure © Mike Bowles, © Steve Milpacher, Hyacinth Macaw © Shutterstock, Black-masked © Steve Martin, © Shutterstock, Green-cheeked Amazon © Mike Bowles, Sulphur-crested Cockatoo © Shutterstock, African Grey © Shutterstock, Kea © Ron Hoff

May 2009 PsittaScene 9 o shed light on the mysterious relationships within parrots, we have worked for the last several years to create an Tevolutionary family tree of parrots (a phylogeny) using genetic data collected with modern molecular techniques. Agapornis roseicollis, Peach-faced In order to obtain a broad sample of Loriculus galgulus, Blue-crowned Bolbopsittacus lunulatus, parrot species, we collaborated with an Neophema elegans, Elegant Parrot Neopsephotus bourkii, Bourke’s Parrot international team of scientists and Barnardius zonarius, Port Lincoln Parrot Platycercus adscitus, Pale-headed Rosella veterinarians (see Special thanks below). Psephotus varius, Mulga Parrot The legal complexities of transporting Purpureicephalus spurius, Red-capped Parrot Northiella haematogaster, Bluebonnet parrot samples (generally preserved Cyanoramphus auriceps, Uellow-fronted Parakeet Eunymphicus uvaeensis, Ouvéa Parakeet blood or frozen tissue) meant that in the Prosopeia tabuensis, Red duivenbodei, end, one of us, (Erin) actually travelled reticulata, Blue-streaked Lory to each of these foreign countries in haematodus, Rainbow Lorikeet Pseudoeos fuscata, order to perform the lab work necessary Glossopsitta porphyrocephala, Purple-crowned Lorikeet albidinucha, White-naped Lory to collect the genetic data. musschenbroekii, Musschenbroek’s Lorikeet papou, Phigys solitarius, Collared Lorikeet australis, Blue-crowned Lorikeet Our globe-trotting efforts resulted in a Melopsittacus undulatus, large family tree (Figure 1) which diophthalma, Double-eyed edwardsii, Edwards’s Fig Parrot includes representative species of 69 of amboinensis, Moluccan King Parrot Aprosmictus erythropterus, Red-winged Parrot the 82 recognized genera. Polytelis alexandrae, Princess Parrot Eclectus roratus, heteroclitus, Singing Parrot Interpreting phylogenetic trees can be columboides, Malabar Parakeet lucionensis, Blue-naped Parrot confusing to the uninitiated, so here we Prioniturus luconensis, Breen Racquet-tailed parrot Micropsitta finschii, Finch’s answer some basic questions about the Amazona viridigenalis, Green-cheeked Amazon Pionus menstruus, Blue-headed Parrot phylogeny and some of the evolutionary Hapalopsittaca amazonina, Rusty-faced Parrot patterns it shows. Pionopsitta caica, Caica Parrot Forpus passerinus, Green-rumped Brotogeris jugularis, Orange-chinned Parakeet Myiopsitta monachus, Monk Parakeet panychlora, Touit batavica, Lilac-tailed Parrotlet Aratinga pertinax, Brown-throated Conure Ara macao, Scarlet Macaw Orthopsittaca manilata, Red-bellied Macaw Cyanopsitta spixii, Spix’s Macaw Diopsittaca nobilis, Red-shouldered Macaw Guaruba guarouba, Golden Conure Leptosittaca branickii, Golden-plumed Conure Cyanoliseus patagonus, Patagonian Conure Deroptyus accipitrinus, Hawk-headed Parrot Pionites melanocephala, Black-capped Parrot Pyrrhura picta, Painted Conure Rhynchopsitta pachyrhyncha, Thick-billed Parrot Enicognathus leptorhynchus, Slender-billed Conure Nandayus nenday, Nanday Conure robustus, Brown-necked Parrot Psittacus erithacus, Grey Parrot Psittrichas fulgidus, Pesquet’s Parrot Coracopsis vasa, Cacatua sulphurea, Yellow-crested Cockatoo Eolophus roseicapilla, Galah Probosciger aterrimus, Palm Cockatoo Nymphicus hollandicus, Cockatiel Calyptorhynchus funereus, Yellow-tailed Black Cockatoo Nestor notabilis, Kea Strigops habroptilus, Kakapo Falco peregrinus, Peregrine Falcon Serinus canarius, Canary Colius colius, White-backed Mousebird Picus canus, Grey-headed Woodpecker Columbina passerina, Common Ground Dove Otus sunia, Oriental Scops-owl Tockus flavirostris, Eastern Yellow-billed Hornbill Figure 1. A phylogeny of the parrots Coccyzus americanus, Yellow-billed Cuckoo constructed using genetic data. Colours denote 50 changes groups of parrot species discussed in the text

Questions? If you have further questions For more information consult our more Special thanks to the Smithsonian about the parrot family tree, please send them detailed paper published in the journal Molecular Institution, the Universidade de São Paulo in to Joanna Eckles, PsittaScene Editor, at Biology and Evolution (Wright et al 2008, 25(10) , Victoria University at Wellington in New [email protected]. We look forward 2141-2156), a version of which is posted at our Zealand, Loro Parque and the National Institute to answering them in a future issue. website http://biology-web.nmsu.edu/twright/. of Toxicology and Forensic Science in the Canary Islands, Spain.

10 PsittaScene May 2009 How do you read a phylogeny? Furthermore, other recently published Why do some of these groups Phylogenies can be thought of as a family phylogenies that have sampled more widely seem jumbled up? 1 tree with descendants branching outward across avian orders have also found 7 There are some distinct surprises in from a single ancestral species. The tips of the contradictory results regarding who the closest the parrot family tree. One is that the tree (on the right in Figure 1) are the species that relatives of parrots might be. This continuing Budgerigar is not closely related to the )are currently in existence. The places where two confusion suggests that parrots are indeed a very rosellas,) bluebonnets and other Platycercine branches join are called nodes, which represent ancient order of birds that split from the parrots of . Instead it is a member the last common ancestor of those two living ancestors of other modern birds a very long of a clade that includes the lorikeets and Fig species. For example, at the top of the time ago, perhaps as long as 80-90 million years Parrots (teal clade). Also, the African parrots phylogeny, the place where the branches leading ago during the Cretaceous Period. The identity do not form a single clade; instead the from the Peach-faced Lovebird (Agapornis of their closest relative remains a mystery African Grey and parrots of the roseicollis) and the Blue-crowned Hanging Parrot, awaiting further investigation. Poicephalus are closely related to each other, (Loriculus galgulus), meet is the node and while the Vasa Parrot of Madagascar represents the last common ancestor of those Who are the most ancient parrots? diverged earlier in the tree and the lovebirds two species. All the branches coming out of a If you work your way along the tree of the genus Agapornis are in an entirely node towards the tips on the right side of the 4 from the basal node connecting the different clade that also includes the tree are the descendents of that ancestor, and parrots to the non-parrots, you will see that Hanging Parrots of (genus the species at the ends of these branches are the first parrot group to split off from the Loriculus) and the odd Guaiabero found only thus more closely related to each other than to )common ancestor of parrots is a group (or in the (top clade in magenta). those coming from a different node. The length 'clade') of New Zealand parrots that includes This pattern suggests that Africa may have of the branches separating a tip from a node, or the Kea and the Kakapo (red clade in Figure 1). been colonized by several different lineages two nodes from each other, represents the This split indicates they are the group that is of parrots at different times. amount of evolution between those nodes. In most-distantly related to the remainder of the our case the length of the branches represents parrots and has interesting implications for the Is nothing of the old the amount of genetic change seen between the geographic origins of parrots, as New Zealand is classifications sacred? sampled DNA sequences. one of first landforms to split from the ancient 8 Some historically recognized groups supercontinent of Gondwana, around 82 million are strongly supported by our tree. In How did you choose the years ago. It supports a long-standing hypothesis addition to the cockatoos and lories (albeit species in your tree? that the ancestors of the modern parrots with) a surprising Budgerigar relative) already 2 Our species were chosen to represent as many originated on this continent and that present mentioned, a core group of the Psittaculine of the existing parrot genera as possible. In distribution of parrots in Australia, South parrots of Australasia that includes the general our choice of which species to use America, southeast Asia and Africa can largely Eclectus Parrot, the Singing Parrot, King )within a genus was guided by availability of be explained by the subsequent breakup of this Parrot and the Princess Parrot received samples appropriate for genetic analysis, either a supercontinent into these modern continents. strong support (dark green clade). A second tissue sample in a museum or a live in a zoo well-supported clade is the Platycercine collection from which we could obtain a blood I thought cockatoos were parrots that include the Australian rosellas, sample. the most ancestral parrots? the bluebonnet, Mulga Parrot and Port 5 Cockatoos have been thought by many Lincoln Parrot, and some species found in Who are the closest relatives to represent the most ancient lineage of New Zealand and as far away as (blue of the parrots? parrots based on some unique anatomical clade). A third strongly supported clade is The non-parrots in our tree include an owl, a )characteristics (including their erectile crests), the Neotropical parrots of Mexico, Central 3 songbird, a hornbill, a woodpecker, a cuckoo, a but this hypothesis was not supported by our America, South America and the West mousebird, a falcon and a dove. These orders genetic data. They were the next group to split Indies, which are all more closely related to )were chosen because at one time or another they off after the New Zealand species and are each other than they are to any other parrots had been suggested as the closest relatives to strongly supported as a distinct group by the (largest clade in pale green). parrots either by morphological or genetic genetic data (orange clade). evidence. They are shaded in grey and branch What's next? outward from the most basal node, which What comes next in the tree? We are continuing our work by represents the common ancestor of all these If you follow the tree up from the node 9 filling in missing genera and by starting avian orders. In this particular tree the falcon and 6 that leads to cockatoos, you will see a to amass data for all species in certain the songbird species branch from the same node large number of nodes separated by short clades. Next up, as part of Erin's PhD that leads to all of the parrots, suggesting that branches, suggesting that there was a rapid dissertation,) is a species-level phylogeny of they are the closest relatives of parrots. This )diversification of parrots that led to the modern the Neotropical parrots. Stay tuned for result, however, was not consistently found in groups such as the lories, the Neotropical more exciting revelations about the other trees we constructed from subsets of the parrots, the African parrots, the various groups parrot family tree! data or with different tree building methods. found in Australia and Asia.

May 2009 PsittaScene 11