Invasion history and demographic processes associated with rapid morphological changes in the Red‐whiskered bulbul established on tropical islands

The Red‐whiskered bulbul is a very successful invasive bird species. Morphological differences have been reported among individuals inhabiting the humid and dry coasts of Reunion Island, in a 30‐year‐old population. This suggests a capacity for rapid local adaptation which could explain the general invasive success of this species. However, the origin and invasion history of this population is unknown. It is therefore not possible to establish with certainty the cause of these morphological differences. Here, we investigated the invasion history of populations of Red‐whiskered bulbul established on Reunion Island, Mauritius and Oahu (three geographically similar tropical islands) to assess the link between invasion history and morphological changes in these populations. We first assessed the source(s) of the invasive populations. We then compared the morphology of the individuals between the invasive and native populations and between the dry and humid coasts of invaded islands. Finally, we inferred the invasion history of the invasive populations to investigate the role of neutral processes (e.g. founder effect and drift) on morphology. We found that the invasive populations have a similar origin and that the morphology of the individuals in these populations has diverged in a similar way from the native range, suggesting a convergent adaptation to tropical islands. Like on Reunion, we found differences in morphology between the dry and humid coasts on Mauritius. These morphological differences can be explained by invasion history on Reunion but not on Mauritius. Both neutral evolution and adaptation thus shape the morphology of invasive Red‐whiskered bulbuls.     

Nuclear DNA from old collections of avian study skins reveals the evolutionary history of the Old World suboscines (Aves, Passeriformes)

Museum study skins represent an invaluable source of DNA for phylogenetics, phylogeography and population genetics. This becomes evident by comparing the number of study skins of birds housed in museums worldwide ( c . 10 million) with the corresponding number of tissue samples (probably fewer than 500 000). While the laboratory techniques used hitherto have primarily allowed PCR-based studies of mitochondrial genes from museum skins, we present here the first avian phylogeny based on a large number of nuclear sequences. The targeted fragment sizes and the properties of the primers used are important contributory factors to obtain good amplification results. In this study we routinely amplified fragments of c. 350 bp nuclear DNA. This advance in methodology opens up a new avenue for the use of avian museum skins, as nuclear DNA is especially useful when studying ancient patterns of diversification. The phylogenetic hypothesis of the Old World suboscines (Eurylaimides) presented herein strongly supports a monophyletic origin of the pittas (Pittidae). The phylogeny further suggests that pittas could be divided into three major groups, in agreement with the external morphological variation found in this group. The broadbills (Eurylaimidae) as currently defined are, on the other hand, found to be a paraphyletic family, as both Sapayoa aenigma and the asities (Philepittidae) are nested among them. Based on the phylogenetic results we suggest a revised classification of the Old World suboscines (Eurylaimides).     

Diversification of Neoaves: integration of molecular sequence data and fossils

Patterns of diversification and timing of evolution within Neoaves, which includes almost 95% of all bird species, are virtually unknown. On the other hand, molecular data consistently indicate a Cretaceous origin of many neoavian lineages and the fossil record seems to support an Early Tertiary diversification. Here, we present the first well-resolved molecular phylogeny for Neoaves, together with divergence time estimates calibrated with a large number of stratigraphically and phylogenetically well-documented fossils. Our study defines several well-supported clades within Neoaves. The calibration results suggest that Neoaves, after an initial split from Galloanseres in Mid-Cretaceous, diversified around or soon after the K/T boundary. Our results thus do not contradict palaeontological data and show that there is no solid molecular evidence for an extensive pre-Tertiary radiation of Neoaves.     

The phylogenetic relationships and generic limits of finches (Fringillidae)

Phylogenetic relationships among the true finches (Fringillidae) have been confounded by the recurrence of similar plumage patterns and use of similar feeding niches. Using a dense taxon sampling and a combination of nuclear and mitochondrial sequences we reconstructed a well resolved and strongly supported phylogenetic hypothesis for this family. We identified three well supported, subfamily level clades: the Holoarctic genus Fringilla (subfamly Fringillinae), the Neotropical Euphonia and Chlorophonia (subfamily Euphoniinae), and the more widespread subfamily Carduelinae for the remaining taxa. Although usually separated in a different family-group taxon (Drepanidinae), the Hawaiian honeycreepers are deeply nested within the Carduelinae and sister to a group of Asian Carpodacus. Other new relationships recovered by this analysis include the placement of the extinct Chaunoproctus ferreorostris as sister to some Asian Carpodacus, a clade combining greenfinches (Carduelis chloris and allies), Rhodospiza and Rhynchostruthus, and a well-supported clade with the aberrant Callacanthis and Pyrrhoplectes together with Carpodacus rubescens. Although part of the large Carduelis-Serinus complex, the poorly known Serinus estherae forms a distinct lineage without close relatives. The traditionally delimited genera Carduelis, Serinus, Carpodacus, Pinicola and Euphonia are polyphyletic or paraphyletic. Based on our results we propose a revised generic classification of finches and describe a new monotypic genus for Carpodacus rubescens.     

Phylogenetic relationships among Palearctic–Oriental starlings and mynas (genera Sturnus and Acridotheres: Sturnidae)

We used nuclear and mitochondrial genes to generate a phylogenetic hypothesis for the Palearctic–Oriental starlings and mynas (genera Sturnus , Acridotheres , Leucopsar , Creatophora and Fregilupus : Sturnidae). Our results indicate that the group has undergone a rapid diversification in Asia since the late Miocene. A reassessment of the morphological and ecological characters used in previous taxonomic revisions shows that all characters are highly homoplastic. We suggest that the elevated morphological and ecological plasticity favoured the adaptation of starlings and mynas to the local environment, the exploitation of all niches and their successful radiation in south-east Asia. Under the current limits the genera Sturnus and Acridotheres are not monophyletic, and we propose a revised phylogenetic taxonomy for the entire clade. We confirm that the extinct Fregilupus varius is a starling and it colonized Réunion Island (Mascarenes) by transoceanic dispersal from Asia.     

Convergent evolution of morphological and ecological traits in the open-habitat chat complex (Aves, Muscicapidae: Saxicolinae)

Snails in the closely related trochid genera Phorcus Risso, 1826 and Osilinus Philippi, 1847 are ecologically important algal grazers in the intertidal zone of the northeastern Atlantic Ocean and Mediterranean Sea. Here we present the first complete molecular phylogeny for these genera, based on the nuclear 28S rRNA gene and the mitochondrial 16S rRNA and COI genes, and show that the current classification is erroneous. We recognize nine species in a single genus, Phorcus: estimated by BEAST analysis, this arose 30 (±10) Ma; it consists of two subgenera, Phorcus and Osilinus, which we estimate diverged 14 (±4.5) Ma. Osilinus kotschyi, from the Arabian and Red Seas, is not closely related and is tentatively referred to Priotrochus Fischer, 1879. Our phylogeny allows us to address biogeographical questions concerning the origins of the Mediterranean and Macaronesian species of this group. The former appear to have evolved from Atlantic ancestors that invaded the Mediterranean on several occasions after the Zanclean Flood, which ended the Messinian Salinity Crisis 5.3 Ma; whereas the latter arose from several colonizations of mainland Atlantic ancestors within the last 3 (±1.5) Ma.     

Higher-level phylogeny and morphological evolution of tyrant flycatchers, cotingas, manakins, and their allies (Aves: Tyrannida)

Despite increased understanding of higher-level relationships in passerine birds in the last 15 years, the taxonomic boundaries and phylogenetic interrelationships of the major groups of the Tyrannida (including the cotingas, manakins, tityrines, and tyrant flycatchers) remain unclear. Here, we present an analysis of DNA sequence data obtained from two nuclear exons, three introns, and one mitochondrial gene for 26 genera of Tyrannida and 6 tracheophone outgroups. The analysis resulted in well-supported hypotheses about the earliest evolution within Tyrannida. The Cotingidae, Pipridae, Tityrinae (sensu) [Prum, R.O., Rice, N.H., Mobley, J.A., Dimmick, W.W., 2000. A preliminary phylogenetic hypothesis for the cotingas (Cotingidae) based on mitochondrial DNA. Auk 117, 236-241], Tyrannidae, and the tyrannid subfamiles Tyranninae and Pipromorphinae (sensu) [Sibley, C.G., Monroe, B. L. Jr., 1990. Distribution and Taxonomy of Birds of the World. Yale University Press, New Haven, CT] were all found to be reciprocally monophyletic (given the present taxon sampling). The Cotingidae and Pipridae form a clade that is the sister group to a well-supported clade including Oxyruncus, the Tityrinae, Piprites, and the Tyrannidae. Oxyruncus is the sister group to the Tityrinae, and Piprites is placed as the sister group to the Tyrannidae. The tyrannid subfamilies Tyranninae and Pipromorphinae are monophyletic sister taxa, but the relationships of Platyrinchus mystaceus to these two clades remains ambiguous. The presence of medial (=internal) cartilages in the syrinx is a synapomorphy for the Oxyruncus-Tityrinae-Piprites-Tyrannidae clade. Although morphological synapomorphies currently support the monophyly of both the Pipridae and the Cotingidae, convergences and/or reversals in morphological character states are common in Tyrannida. The relationship between Oxyruncus and the Tityrinae is congruent with additional syringeal synapomorphies and allozyme distance data. Accordingly, we propose the recognition the family Tityridae within the Tyrannida to include the genera Schiffornis, Laniisoma, Laniocera, Iodopleura, Xenopsaris, Pachyramphus, Tityra, and Oxyruncus.     

Rapid morphological changes,admixture and invasive success in populations of Ring-necked parakeets (<Emphasis Type="Italic">Psittacula krameri</Emphasis>) established in Europe

The Ring-necked parakeet (Psittacula krameri), native of Asia and Africa, is a very successful invasive species in Europe: it has been present there for over 50 years. A recent study showed that European invasive populations occupy a colder climatic niche than in their native range but the establishment of this tropical species in temperate regions remains unexplained. Two main hypotheses may explain the success of Ring-necked parakeet in Europe: admixture between individuals from different origins and/or rapid adaptation to new environmental conditions. In this study, we investigated with molecular data the origin of European populations of Ring-necked parakeets to assess whether these populations result from admixture between individuals from different source populations. We also investigated the morphology of individuals from European populations and from the native range to assess whether the invasive populations have morphologically diverged from their source and could have become adapted to European conditions. We found evidence of admixture in some of the European populations but not all of them. Admixture between individuals from different origins within European populations thus cannot explain alone their invasive success. Conversely, we found that the morphology of the individuals from European populations has diverged from the morphology of native individuals, in a similar direction. Rapid adaptation to European environmental conditions via phenotypic plasticity or natural selection could thus be a factor explaining the invasive success of Ring-necked parakeets in Europe.