Miocene diversification of an open‐habitat predatorial passerine radiation,the shrikes (Aves: Passeriformes: Laniidae)

Diversification of avifaunas associated with savannah and steppes appears to correlate with open habitats becoming available, starting in the Miocene. Few comparative analyses exist for families for which all species are predominantly adapted to these habitats. One such group is Laniidae (Passeriformes), which are small‐ to medium‐sized predatory passerines known for their distinctive behaviour of impaling prey. We used multispecies coalescent‐based and concatenation methods to provide the first complete species‐level phylogeny for this group, as well as an estimate of the timing of diversification. Our analyses indicate that Laniidae as currently delimited is not monophyletic, as the genus Eurocephalus is not closely related to the remaining species. The two species currently assigned to the monotypic genera Urolestes and Corvinella are part of the same clade as the Lanius species, and we propose that they are included in the genus Lanius, making Laniidae monogeneric. The initial diversification of the clade is inferred to have occurred very rapidly, starting about 7.2–9.1 million years ago, timing depending on calibration method, but in either case coinciding with the expansion of C4 grasses. An African origin is inferred in the biogeographic analysis. In the redefined Laniidae, cooperative breeding is inferred to be restricted to a single clade, characterized by gregarious behaviour and rallying. Migratory behaviour evolved multiple times within the family.     

Phylogeny and biogeography of the parrot genus Prioniturus (Aves: Psittaciformes)

The parrot genus Prioniturus occurs in the oceanic Philippines, Palawan and Wallacea, a geologically dynamic region with a complex history of land and sea. The described taxa of Prioniturus have been variously placed in different assemblages, and different numbers of species have been recognized. However, a phylogenetic framework is so far lacking. This would be the prerequisite to reconstructing dispersal and colonization patterns of Prioniturus across and within Wallacea and the Philippines. Following our robustly supported phylogenetic hypothesis based on two mitochondrial genes, we propose to treat Prioniturus mindorensis comb. nov. as well as Prioniturus montanus and Prioniturus waterstradti as separate species. In Prioniturus discurus discurus and Prioniturus discurus whiteheadi, further studies using additional data and specimens are necessary to clarify their taxonomic status. This result is congruent with other studies demonstrating that alpha diversity of the Philippine avifauna is strongly underestimated. According to our biogeographic reconstruction, Prioniturus has diversified by a complex combination of colonization of islands and subsequent divergence in allopatry among and within island groups. Dispersal between Sulawesi/Wallacea and the Philippines occurred twice and documents a rare case of faunal exchange between these two regions.     

The Hawaiian Archipelago is a stepping stone for dispersal in the Pacific: an example from the plant genus Melicope (Rutaceae)

Aim Pacific biogeographical patterns in the widespread plant genus Melicope J.R. Forst. & G. Forst. (Rutaceae) were examined by generating phylogenetic hypotheses based on chloroplast and nuclear ribosomal sequence data. The aims of the study were to identify the number of colonization events of Melicope to the Hawaiian Islands and to reveal the relationship of Hawaiian Melicope to the Hawaiian endemic genus Platydesma H. Mann. The ultimate goal was to determine if the Hawaiian Islands served as a source area for the colonization of Polynesia. Location Nineteen accessions were sampled in this study, namely eight Melicope species from the Hawaiian Islands, four from the Marquesas Islands, one species each from Tahiti, Australia and Lord Howe Island, two Australian outgroups and two species of the Hawaiian endemic genus Platydesma. To place our results in a broader context, 19 sequences obtained from GenBank were included in an additional analysis, including samples from Australia, Papua New Guinea, New Zealand, Southeast Polynesia and Asia. Methods DNA sequences were generated across 19 accessions for one nuclear ribosomal and three chloroplast gene regions. Maximum parsimony analyses were conducted on separate and combined data sets, and a maximum likelihood analysis was conducted on the combined nuclear ribosomal and chloroplast data set. A broader nuclear ribosomal maximum parsimony analysis using sequences obtained from GenBank was also performed. Geographic areas were mapped onto the combined chloroplast and nuclear ribosomal tree, as well as onto the broader tree, using the parsimony criterion to determine the dispersal patterns. Results Phylogenetic analyses revealed that Platydesma is nested within Melicope and is sister to the Hawaiian members of Melicope. The Hawaiian Melicope + Platydesma lineage was a result of a single colonization event, probably from the Austral region. Finally, Marquesan Melicope descended from at least one, and possibly two, colonization events from the Hawaiian Islands. Main conclusions These data demonstrate a shifting paradigm of Pacific oceanic island biogeography, in which the patterns of long‐distance dispersal and colonization in the Pacific are more dynamic than previously thought, and suggest that the Hawaiian Islands may act as a stepping stone for dispersal throughout the Pacific.     

Historical biogeography of an Indo‐Pacific passerine bird family (Pachycephalidae): different colonization patterns in the Indonesian and Melanesian archipelagos

Aim We use molecular‐based phylogenetic methods and ancestral area reconstructions to examine the systematic relationships and biogeographical history of the Indo‐Pacific passerine bird family Pachycephalidae (whistlers). Analysed within an explicit spatiotemporal framework, we elucidate distinct patterns of diversification across the Melanesian and Indonesian archipelagos and explore whether these results may be explained by regional palaeogeological events. We further assess the significance of upstream colonization and its role in species accumulation within the region. Location The Indo‐Pacific region, with an emphasis on the archipelagos on either side of the Australo‐Papuan continent. Methods We used three nuclear and two mitochondrial markers to construct a molecular phylogenetic hypothesis of the Pachycephalidae by analysing 35 of the 49 species known to belong to the family. The programs diva and Mr Bayes were used to reconstruct ancestral area relationships and to examine biogeographical relationships across the family, and beast was implemented to assess the timing of dispersal events. Results We constructed a molecular phylogenetic hypothesis for the Pachycephalidae and estimated divergence times and ancestral area relationships. Different colonization patterns are apparent for the Pachycephalidae in the Indonesian and the Melanesian archipelagos. The Indonesian archipelago was colonized numerous times, whereas one or two colonizations of the Melanesian archipelagos account for the entire diversity of that region. After initial colonization of the Melanesian archipelagos some whistler species recolonized Australia and may have commenced a second round of colonization into Melanesia. Main conclusions The contrasting dispersal patterns of whistlers in archipelagos on either side of the Australo‐Papuan continent are congruent with the arrangement and history of islands in each of the regions and demonstrate that knowledge of palaeogeography is important for an understanding of evolutionary patterns in archipelagos. We also highlight that recolonization of continents from islands may be more common than has previously been assumed.     

Increased taxon and character sampling reveals novel intergeneric relationships in the Cracidae (Aves: Galliformes)

The Cracidae is one of the most endangered bird families in the World. Several studies have been published recently on the evolution and conservation of cracids. Phylogenetic analyses using a fragment of 661 bp of the mitochondrial cytochrome b gene for 39 different species of cracids corroborated most relationships found in previous studies. The present work attempts to refine the former phylogenetic hypothesis by increasing taxon sampling and combining molecular with osteological, integumentary and behavioural characters using Maximum Parsimony (MP) and Bayesian analyses. We present both separate and combined total evidence analyses with our molecular data, 152 osteological and 74 integumentary + behavioural characters. While supporting most aspects of the molecular-based hypotheses, the tree based on the combined matrix suggests several modifications of the generic composition for each of the two subfamilies: Penelopinae and Cracinae, and supports the merging of the genera Pipile with Aburria and Mitu with Pauxi . These results suggest that increased taxon and character sampling from a diversity of sources may be at least as important as increased sampling of only one type. Besides, of a total of 891 characters we had 437 parsimony-informative sites (almost half of the analyzable sites) proving the efficiency of a total-evidence approach.     

Multiple cenozoic invasions of Africa by penguins (Aves, Sphenisciformes)

Africa hosts a single breeding species of penguin today, yet the fossil record indicates that a diverse array of now-extinct taxa once inhabited southern African coastlines. Here, we show that the African penguin fauna had a complex history involving multiple dispersals and extinctions. Phylogenetic analyses and biogeographic reconstructions incorporating new fossil material indicate that, contrary to previous hypotheses, the four Early Pliocene African penguin species do not represent an endemic radiation or direct ancestors of the living Spheniscus demersus (blackfooted penguin). A minimum of three dispersals to Africa, probably assisted by the eastward-flowing Antarctic Circumpolar and South Atlantic currents, occurred during the Late Cenozoic. As regional sea-level fall eliminated islands and reduced offshore breeding areas during the Pliocene, all but one penguin lineage ended in extinction, resulting in today's depleted fauna.     

The colubrid radiation in Africa (Serpentes: Golubridae): phylogenetic relationships and evolutionary patterns based on immunological data

Phylogenetic relationships among genera of African colubrids were evaluated using estimates of divergence among serum albumins compared by microcomplement fixation. Representatives of about half of the extant genera of African colubrids, as well as the Elapidae, Atractaspis and the Madagascan colubrid Leioheterodon, were analysed. The tree of best fit to the data has an unresolved basal polychotomy comprising at least five lineages of colubrids, as well as Elapidae and Atractaspis; thus, colubrids were not demonstrably monophyletic with these data. Two cosmopolitan clades, colubrines and natricines, are represented in Africa by series of closely related genera, but divergence among other genera is relatively great. Rate tests show that this is apparently not due to higher rates of albumin evolution in these, relative to other colubrids. Among the other associations supported by the immunological data are: (1) Psammophis-(Rhamphiophis-Dipsina)-Malpolon-Psammophylax; (2) Amblyodipsas-Macrelaps; (3) (Lycodonomorphus-Lamprophis)-Mehelya; and (4) Colubrinae-Natricinae. Grayia is questionably associated with the colubrine-natricine lineage. Prosymna and Lycodon are clearly members of the colubrine clade, and Amplorhinus possibly associates with Leioheterodon. Gonionotophis, Duherria, Lycophidion and Pseudaspis show no strong association with any other genera, and represent other basal or near-basal clades within the colubrid/elapid radiation. The immunological data do not support a clade comprising the Elapidae, Atractaspis and some ‘aparallactines’ relative to Viperidae and other colubrids. The basal colubrid-elapid-Atractaspis divergence occurred more than 30 Myr ago, and the fossil record of colubrids in Africa greatly underestimates both the age and clade diversity of this group. In contrast to the pattern of radiation in the neotropics, where most colubrids belong to one of three major clades, in Africa only the colubrine lineage comprises a substantial portion of the extant generic diversity; most other genera stem from relatively ancient cladogenetic events and have few living representatives.     

Short-faced bats (Phyllostomidae: Stenodermatina): a Caribbean radiation of strict frugivores

Aim  To test the hypothesis that Caribbean Short-faced bats descended from a single recent ancestor that originated in the continental Neotropics (Mexico, Central America and/or South America).
Location  The Neotropics, including the West Indies.
Methods  New mitochondrial cytochrome b and nuclear Rag2 sequences were combined with published molecular data to estimate phylogenetic relationships and sequence divergence among Short-faced bats. The resulting phylogenies were compared with those compatible with the single-origin hypothesis using two model-based statistical tests. Confidence limits on sequence divergence were estimated using a parametric bootstrap.
Results  All molecular phylogenies revealed two independent Caribbean lineages and showed that continental Short-faced bats share a recent common ancestor. Morphology-based trees compatible with the single-origin hypothesis were significantly worse at explaining the molecular data than any molecular phylogeny.
Main conclusions  The ancestor of all Short-faced bats reached the Antilles in the Miocene, too recently to have used a proposed Oligocene land bridge, and well before the Pleistocene glaciations that are thought to have facilitated dispersal for many bats. After a long period of isolation, Short-faced bats diversified quickly on the Caribbean islands. A single Short-faced lineage then reached the continent and subsequently expanded its range and diversified into the four extant genera. Among bats, independent lineages of aerial insectivores and nectarivores have also recolonized the continent after evolving in the West Indies. The evidence for an insular origin of the short-faced frugivorous radiation completes a dynamic model of Caribbean biogeography that encompasses an entire biological community.     

Ancient origins of Indo-Pacific coral reef fish biodiversity: a case study of the leopard wrasses (Labridae: Macropharyngodon)

Temporal origins of reef fishes in the Indo-Australian Archipelago were examined using wrasses in the genus Macropharyngodon. The genus was selected as it is morphologically and ecologically distinct, with strongly reef-associated species exhibiting discrete distributions across the Indo-Pacific. Phylogenetic relationships were explored using COI, 16S, and 12S rRNA mitochondrial sequences. Monophyly of the genus was supported by congruent Bayesian, maximum likelihood, and maximum parsimony trees. Estimates of lineage ages based on fossil-calibrated reef fish divergences suggest that Macropharyngodon had an extensive evolutionary history starting in the early Miocene. Repeated divergences of Indian Ocean-Pacific Ocean lineages appear to have occurred over at least 19 million years. Regional endemics represent both old and young clades. Our estimates of early Miocene origins, and mid-Miocene to Pliocene diversifications of Macropharyngodon are supported by recent studies of other reef fish genera, and emphasise the importance of pre-Pleistocene events in generating Indo-Pacific coral reef fish biodiversity.     

Biogeography of Plagiochila (Hepaticae): natural species groups span several floristic kingdoms

Aim This paper presents a synthesis of our recent results regarding the biogeography of Plagiochila using a molecular approach, and documents intercontinental ranges within this largest genus of the hepatics. Methods A maximum likelihood analysis of sixty‐one nrITS sequences of Plagiochila was performed and the molecular topology obtained was compared with morphological, phytochemical and geographical data. Results Our molecular data set allowed the identification of eleven Plagiochila sections, the majority of which cover at least two floristic kingdoms. Seven sections have species in Europe (sect. Arrectae, Carringtoniae, Fuscoluteae, Glaucescentes, Plagiochila, Rutilantes, Vagae). Plagiochila species from Atlantic Europe are usually close to or conspecific with neotropical taxa, whereas species widespread in Europe are closely related to Asian ones and not to those in the Neotropics. Plagiochila sect. Arrectae represents a neotropical – Atlantic European clade. The section is not closely related – as has often been suggested – to the morphologically similar sect. Zonatae from Asia and western North America. Sequence data show that the African P. integerrima and the neotropical P. subplana are members of the Asian sect. Cucullatae (sect. Ciliatae, syn. nov.), which becomes pantropical in distribution. An ITS sequence of P. boryana from Uganda confirms the Afro‐American range of the primarily neotropical sect. Hylacoetes. Similarities in sporophyte morphology between the sect. Cucullatae and sect. Hylacoetes are the result of parallel evolution. Main conclusions Our results indicate that intercontinental ranges at section and species level are common in Plagiochila. Carl's (1931) subdivision of Plagiochila into sections restricted to one floristic kingdom is outdated. Biogeographical patterns in Plagiochila are not dissimilar to those of other groups of bryophytes but elucidation of the geographical ranges of the taxa requires a molecular approach. Contrary to earlier belief, most Plagiochila species from Atlantic Europe do not have close relatives in Asia but are conspecific with or closely related to species from tropical America.     

Turnover of passerine birds on islands in the Aegean Sea (Greece)

Aim We wish to determine the effect of migratory status on turnover rates in island birds. Because turnover is influenced by factors other than migratory status, we also considered the influence of body size and physical characteristics of the islands inhabited on the probabilities of extinction and immigration. Location The Mediterranean islands of Delos, Astypalea, Paros, Naxos and Lesvos in the Aegean Sea, Greece. Methods The passerine birds of these islands were surveyed between 1954 and 1961 by G.E. Watson, and were resurveyed between 1988 and 1992. The effects of migratory status and body size on the probabilities of extinction and immigration were examined by G‐tests of linear trend in proportion, and analysis of variance, respectively. A combined analysis of migratory status, body size and physical characteristics of the islands was carried out using logistic regressions of the probabilities of extinction and immigration on these factors. Results Species number on each island changed little between surveys, with no island's species number changing by more than one species. Twelve population extinctions and 11 immigrations were recorded. The smallest island, Delos (6 km²), had the highest annualized relative turnover rate (1.08), while the four larger islands (96–1614 km²) had lower and mutually similar rates (0.21–0.27). Populations on higher elevation islands were less likely to go extinct. There is no evidence for an effect of body size on the probabilities of extinction or immigration. Migratory status affected extinction and immigration probabilities differently: migratory species were more likely to immigrate, but less likely to go extinct. Main conclusions The position of the Aegean islands along a major north–south flyway may account for the observed effects of migratory status. The annual passage of large numbers of migrants may, via the rescue effect, decrease the chances of extinction, while at the same time increasing the chances of colonization of unoccupied islands. The likelihood of both extinction and immigration involves a complex interaction between life‐history traits and island characteristics. The effects of migratory status will depend not only on consideration of vagility, vulnerability and stochasticity identified by previous authors, but also upon the location of the islands in relationship to migratory pathways.     

Ancient differentiation in the single-island avian radiation of endemic Hispaniolan chat-tanagers (Aves: Calyptophilus)

The simple geographic structure of island systems often makes them tractable for studies of the patterns and processes of biological diversification. The Calyptophilus chat-tanagers of Hispaniola are of general evolutionary interest because their multiple lineages might have arisen on a single island, of conservation concern because several isolated populations are nearly extinct, and taxonomically ambiguous because they have been variously lumped or split into one to four species. To explore the context of diversification of the seven extant Calyptophilus populations, we conducted a multilocus coalescent analysis based on sequences of mitochondrial ND2 and three nuclear intron loci. We then compared patterns of phylogeographic genetic variation with the morphological differences that distinguish these populations. Mitochondrial haplotypes formed two reciprocally monophyletic groups separated by a large magnitude of nucleotide divergence. Intron structure largely paralleled the geographic grouping pattern of the mitochondrial DNA (mtDNA), but these groups were only reciprocally monophyletic at one of the three introns. Also, the magnitude of between-group divergence was much lower in the introns than mtDNA genealogies. Multilocus coalescent analyses inferred a nonzero divergence time between these two major geographic groups, but suggested that they have experienced a low level of gene flow. All four markers showed substantial allele sharing within each of the two groups, demonstrating that many now separated montane populations do not have long histories of isolation. Considered in concert, our multilocus phylogeographic reconstructions support the recognition of two species within the Calyptophilus complex, and raise the possibility that these taxa differentiated prior to the fusion of the two palaeo-islands that form present-day Hispaniola.     

A new Transantarctic relationship: morphological evidence for a Rheidae–Dromaiidae–Casuariidae clade (Aves,Palaeognathae, Ratitae)

Although ratites have been studied in considerable detail, avian systematists have been unable to reach a consensus regarding their relationships. Morphological studies indicate a basal split separating Apterygidae from all other extant ratites, and a sister‐group relationship between Rheidae and Struthionidae. Molecular studies have provided evidence for the paraphyly of the Struthionidae and Rheidae, with respect to a clade of Australasian extant ratites. The position of the extinct Dinornithidae and Aepyornithidae also remains hotly debated. A novel pattern of diversification of ratites is presented herein. The phylogenetic analysis is based on 17 taxa and 129 morphological characters, including 77 new characters. The resultant tree yields a sister‐group relationship between New Zealand ratites (Apterygidae plus Dinornithidae) and all other ratites. Within this clade, the Aepyornithidae and Struthionidae are successive sister taxa to a new, strongly supported clade comprising the Rheidae, Dromaiidae, and Casuariidae. The link between South American and Australian biotas proposed here is congruent with numerous studies that have evidenced closely related taxa on opposite sides of the Southern Pacific. These repeated patterns of area relationships agree with current knowledge on Gondwana break‐up, which indicates that Australia and South America remained in contact across Antarctica until the earliest Tertiary. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 641–663.     

Multiple origins of Locusta migratoria (Orthoptera: Acrididae) in the Japanese Archipelago and the presence of two major clades in the world: evidence from a molecular approach

Phylogenetic relationships among migratory locust (Locusta migratoria) populations in different climatic regions were analysed by sequencing four mitochondrial DNA regions, with special reference to the origin of Japanese populations. The populations are clearly separated into two clades: one consists of individuals from temperate and cold‐temperate areas of Japan and the Chinese continent, and the other comprises those from subtropical islands of Japan, Hainan Island in China, Timol Leste, Australia, Ethiopia, France, and some individuals from Tsushima Island and Honshu of Japan. The divergence time between the two clades is estimated to be 0.86–1.89 Mya. The phylogenetic analysis revealed that Japanese L. migratoria populations were composed of individuals of six different origins: (1) Hokkaido populations possibly from the Russian continent; (2) Honshu–Kyushu populations from the Chinese continent; (3) Southwest Island populations from Hainan Island or adjacent areas; (4) Ogasawara populations that might have originated from Micronesia; (5) part of the Tsushima population that originated from somewhere in the Asian tropics; and (6) a possible relict population of ancient southern haplotypes that exists in western areas of northern Honshu. The Tsugaru Straits and Tokara Straits have acted as effective geographical barriers, as in other organisms, isolating locust populations for a few thousand years. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 570–581.     

Systematic placement of an enigmatic Southeast Asian taxon Eupetes macrocerus and implications for the biogeography of a main songbird radiation, the Passerida

Biogeographic connections between Australia and other continents are still poorly understood although the plate tectonics of the Indo-Pacific region is now well described. Eupetes macrocerus is an enigmatic taxon distributed in a small area on the Malay Peninsula and on Sumatra and Borneo. It has generally been associated with Ptilorrhoa in New Guinea on the other side of Wallace's Line, but a relationship with the West African Picathartes has also been suggested. Using three nuclear markers, we demonstrate that Eupetes is the sister taxon of the South African genus Chaetops, and their sister taxon in turn being Picathartes, with a divergence in the Eocene. Thus, this clade is distributed in remote corners of Africa and Asia, which makes the biogeographic history of these birds very intriguing. The most parsimonious explanation would be that they represent a relictual basal group in the Passerida clade established after a long-distance dispersal from the Australo-Papuan region to Africa. Many earlier taxonomic arrangements may have been based on assumptions about relationships with similar-looking forms in the same, or adjacent, biogeographic regions, and revisions with molecular data may uncover such cases of neglect of ancient relictual patterns reflecting past connections between the continents.     

Adaptive radiation in miniature: the minute salamanders of the Mexican highlands (Amphibia: Plethodontidae: Thorius)

The small size and apparent external morphological similarity of the minute salamanders of the genus Thorius have long hindered evolutionary studies of the group. We estimate gene and species trees within the genus using mitochondrial and nuclear DNA from nearly all named and many candidate species and find three main clades. We use this phylogenetic hypothesis to examine patterns of morphological evolution and species coexistence across central and southern Mexico and to test alternative hypotheses of lineage divergence with and without ecomorphological divergence. Sympatric species differ in body size more than expected after accounting for phylogenetic relationship, and morphological traits show no significant phylogenetic signal. Sympatric species tend to differ in a combination of body size, presence or absence of maxillary teeth, and relative limb or tail length, even when they are close relatives. Sister species of Thorius tend to occupy climatically similar environments, which suggests that divergence across climatic gradients does not drive species formation in the genus. Rather than being an example of cryptic species formation, Thorius more closely resembles an adaptive radiation, with ecomorphological divergence that is bounded by organism‐level constraints. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 622–643.