The question of when modern birds (Neornithes) first diversified has generated much debate among avian systematists. Fossil evidence generally supports a Tertiary diversification, whereas estimates based on molecular dating favor an earlier diversification in the Cretaceous period. In this study, we used an alternate approach, the inference of historical biogeographic patterns, to test the hypothesis that the initial radiation of the Order Psittaciformes (the parrots and cockatoos) originated on the Gondwana supercontinent during the Cretaceous. We utilized broad taxonomic sampling (representatives of 69 of the 82 extant genera and 8 outgroup taxa) and multilocus molecular character sampling (3,941 bp from mitochondrial DNA (mtDNA) genes cytochrome oxidase I and NADH dehydrogenase 2 and nuclear introns of rhodopsin intron 1, tropomyosin alpha-subunit intron 5, and transforming growth factor ss-2) to generate phylogenetic hypotheses for the Psittaciformes. Analyses of the combined character partitions using maximum parsimony, maximum likelihood, and Bayesian criteria produced well-resolved and topologically similar trees in which the New Zealand taxa Strigops and Nestor (Psittacidae) were sister to all other psittaciforms and the cockatoo clade (Cacatuidae) was sister to a clade containing all remaining parrots (Psittacidae). Within this large clade of Psittacidae, some traditionally recognized tribes and subfamilies were monophyletic (e.g., Arini, Psittacini, and Loriinae), whereas several others were polyphyletic (e.g., Cyclopsittacini, Platycercini, Psittaculini, and Psittacinae). Ancestral area reconstructions using our Bayesian phylogenetic hypothesis and current distributions of genera supported the hypothesis of an Australasian origin for the Psittaciformes. Separate analyses of the timing of parrot diversification constructed with both Bayesian relaxed-clock and penalized likelihood approaches showed better agreement between geologic and diversification events in the chronograms based on a Cretaceous dating of the basal split within parrots than the chronograms based on a Tertiary dating of this split, although these data are more equivocal. Taken together, our results support a Cretaceous origin of Psittaciformes in Gondwana after the separation of Africa and the India/Madagascar block with subsequent diversification through both vicariance and dispersal. These well-resolved molecular phylogenies will be of value for comparative studies of behavior, ecology, and life history in parrots.

Behavioral flexibility is an important adaptive response to changing environments for many animal species. Such plasticity may also promote the invasion of novel habitats by introduced species by providing them with the ability to expand or change their ecological niche, a longstanding idea with recent empirical support. At the individual level, flexibility may arise through innovation, in which an individual invents a new behavior, or through social learning, in which an individual adopts a behavior used by others. There is increasing evidence that the adaptive value of these two modes of learning, and the overall expression of behavioral flexibility, may vary with social and environmental context. In this paper, we propose that invasive species may change the degree to which they express behavioral flexibility in an adaptive manner during the different stages of invasion. Specifically, the "adaptive flexibility hypothesis" predicts that the expression of behavioral flexibility, and thus the diversity of behaviors observed in a population, will be high during the initial stage of introduction into a novel environment due to innovation, followed by a decline in behavioral diversity during the establishment and growth of a founding population due to social learning of successful behavioral variants. We discuss several alternatives to this hypothesis and suggest empirical and theoretical tests of these hypotheses. This "adaptive flexibility hypothesis" suggests that a more nuanced approach to the study of the behaviors employed by individuals in populations at different invasion stages could generate new insight into the importance of such flexibility during species invasions, and the evolution of behavioral plasticity in general.

We report a duplication and rearrangement of the mitochondrial genome involving the control region of parrots in the genus Amazona. This rearrangement results in a gene order of cytochrome b/tRNA(Thr)/pND6/pGlu/CR1/tRNA(Pro)/NADH dehydrogenase 6/tRNA(Glu)/CR2/tRNA(Phe)/12s rRNA, where CR1 and CR2 refer to duplicate control regions, and pND6 and pGlu indicate presumed pseudogenes. In contrast to previous reports of duplications involving the control regions of birds, neither copy of the parrot control region shows any indications of degeneration. Rather, both copies contain many of the conserved sequence features typically found in avian control regions, including the goose hairpin, TASs, the F, C, and D boxes, conserved sequence box 1 (CSB1), and an apparent homolog to the mammalian CSB3. We conducted a phylogenetic analysis of homologous portions of the duplicate control regions from 21 individuals representing four species of Amazona (A. ochrocephala, A. autumnalis, A. farinosa, and A. amazonica) and Pionus chalcopterus. This analysis revealed that an individual's two control region copies (i.e., the paralogous copies) were typically more closely related to one another than to corresponding segments of other individuals (i.e., the orthologous copies). The average sequence divergence of the paralogous control region copies within an individual was 1.4%, versus a mean value of 4.1% between control region orthologs representing nearest phylogenetic neighbors. No differences were found between the paralogous copies in either the rate or the pattern in which the two copies accumulated base pair changes. This pattern suggests concerted evolution of the two control regions, perhaps through occasional gene conversion events. We estimated that gene conversion events occurred on average every 34,670 +/- 18,400 years based on pairwise distances between the paralogous control region sequences of each individual. Our results add to the growing body of work indicating that under some circumstances duplicated mitochondrial control regions are retained through evolutionary time rather than degenerating and being lost, presumably due to selection for a small mitochondrial genome.

Although the poaching of nestlings for the pet trade is thought to contribute to the decline of many species of parrots, its effects have been poorly demonstrated. We calculated rates of mortality due to nest poaching in 23 studies of Neotropical parrots, representing 4024 nesting attempts in 21 species and 14 countries. We also examined how poaching rates vary with geographic region, presence of active protection programs, conservation status and economic value of a species, and passage of the U.S. Wild Bird Conservation Act. The average poaching rate across all studies was 30% of all nests observed. Thirteen studies reported poaching rates of ≥20%, and four reported rates openface> 70%. Only six studies documented no nest poaching. Of these, four were conducted on islands in the Caribbean region, which had significantly lower poaching rates than the mainland Neotropics. The other two studies that showed no poaching were conducted on the two species with the lowest economic value in our sample ( U.S. retail price). In four studies that allowed direct comparison between poaching at sites with active nest protection versus that at unprotected sites, poaching rates were significantly lower at protected sites, suggesting that active protection efforts can be effective in reducing nest poaching. In those studies conducted both before and after the passage of the U.S. Wild Bird Conservation Act, poaching rates were found to be significantly lower following its enactment than in the period before. This result supports the hypothesis that the legal and illegal parrot trades are positively related, rather than inversely related as has been suggested by avicultural interests. Overall, our study indicates that poaching of parrot nestlings for economic gain is a widespread and biologically significant source of nest mortality in Neotropical parrots.