Breeding system evolution influenced the geographic expansion and diversification of the core Corvoidea (Aves: Passeriformes)

Birds vary greatly in their life‐history strategies, including their breeding systems, which range from brood parasitism to a system with multiple nonbreeding helpers at the nest. By far the most common arrangement, however, is where both parents participate in raising the young. The traits associated with parental care have been suggested to affect dispersal propensity and lineage diversification, but to date tests of this potential relationship at broad temporal and spatial scales have been limited. Here, using data from a globally distributed group of corvoid birds in concordance with state‐dependent speciation and extinction models, we suggest that pair breeding is associated with elevated speciation rates. Estimates of transition between breeding systems imply that cooperative lineages frequently evolve biparental care, whereas pair breeders rarely become cooperative. We further highlight that these groups have differences in their spatial distributions, with pair breeders overrepresented on islands, and cooperative breeders mainly found on continents. Finally, we find that speciation rates appear to be significantly higher on islands compared to continents. These results imply that the transition from cooperative breeding to pair breeding was likely a significant contributing factor facilitating dispersal across tropical archipelagos, and subsequent world‐wide phylogenetic expansion among the core Corvoidea.     

Origin and evolution of endemic plants of the Bonin (Ogasawara) Islands

The Bonin Islands are typical oceanic islands, located at the western part of the North Pacific Ocean and approximately 1,000 km south of mainland Japan. This archipelago consists of about 20 small islands. Although floristic diversity is low due to the small area and limited environmental diversity, the Bonin Islands harbor unique endemic flora as in other well-known oceanic islands. This paper presents a brief summary of the results obtained from recent studies on the endemic flora of the Bonin Islands. The results are reviewed in relation to the four stages of the evolution of endemic flora in the oceanic islands; migration, establishment, enlargement and diversification. The ancestors of the flora originated mostly from tropical and subtropical Southeast Asia or mainland Japan by rare events of long distance dispersal. The proportion of bird-dispersed species is relatively high as for other oceanic islands. Genetic data sets obtained from allozyme variation in some endemic species suggest that migration occurred several million years ago and genetic diversity is correlated with current population size. At the time of establishment, self-compatible plants are expected to have an advantage. However, the percentage of dioecious plants is relatively high. This is partly due to evolutionary changes from hermaphroditic ancestors to dioecy which occurred in two genera in the Bonin Islands. In addition, there are some examples of evolutionary changes from herbaceous ancestors to woody endemics. Adaptive radiation is found in some genera, although the number of congeneric endemic species is less than five. Studies of allozyme variation inPittosporum, Symplocos andCrepidiastrum showed that genetic identity is generally very high between congeneric species in spite of their distinct morphologies. This result suggests that divergence of these species occurred rather recently and distinct morphological differences are based on a limited number of genetic changes.     

Molecular phylogenetics of the Macaronesian-endemic genus Bystropogon (Lamiaceae): palaeo-islands, ecological shifts and interisland colonizations

Abstract A molecular phylogenetic study of Bystropogon L'Her. (Lamiaceae) is presented. We performed a cladistic analysis of nucleotide sequences of the internal transcribed spacers (ITS), of the nuclear ribosomal DNA, and of the trnL gene and trnL-trnF intergenic spacer of the chloroplast DNA. Bystropogon odoratissimus is the only species endemic to the Canary Islands that occurs in the three palaeo-islands of Tenerife. This species is not part of an early diverging lineage of Bystropogon and we suggest that it has a recent origin. This phylogenetic pattern is followed by most of the species endemic to the palaeo-islands of Tenerife. The two sections currently recognized in Bystropogon form two monophyletic groups. Taxa belonging to the section Bystropogon clade show interisland colonization limited to the Canary Islands with ecological shifts among three ecological zones. Taxa from the section Canariense clade show interisland colonization both within the Canary Islands and between the Canary Islands and Madeira. Speciation events within this clade are mostly limited to the laurel forest. The genus has followed a colonization route from the Canaries towards Madeira. This route has also been followed by at least five other plant genera with species endemic to Macaronesia. Major incongruences were found between the current infrasectional classification and the molecular phylogeny, because the varieties of Bystropogon origanifolius and Bystropogon canariensis do not form two monophyletic groups. The widespread B. origanifolius appears as progenitor of the other species in section Bystropogon with a more restricted distribution.     

Persistent inter‐ and intraspecific gene exchange within a parallel radiation of caterpillar hunter beetles (Calosoma sp.) from the Galápagos

When environmental gradients are repeated on different islands within an archipelago, similar selection pressures may act within each island, resulting in the repeated occurrence of ecologically similar species on each island. The evolution of ecotypes within such radiations may either result from dispersal, that is each ecotype evolved once and dispersed to different islands where it colonized its habitat, or through repeated and parallel speciation within each island. However, it remains poorly understood how gene flow during the divergence process may shape such patterns. In the Galápagos islands, three phenotypically similar species of the beetle genus Calosoma occur at higher elevations of different islands, while lowlands are occupied by a fourth species. By genotyping all major populations within this radiation for two nuclear and three mitochondrial gene fragments and seven microsatellite markers, we found strong support that the oldest divergence separates the highland species of the oldest island from the remaining species. Despite their morphological distinctness, highland species of the remaining islands were genetically closely related to the lowland population on each island and within the same magnitude as lowland populations sampled at different islands. Repeated evolution of highland ecotypes out of the lowland species appears the most likely scenario and estimates of geneflow rates revealed extensive admixture among ecotypes within islands, as well as between islands. These findings indicate that gene exchange among the different populations and species may have shaped the phylogenetic relationships and the repeated evolution of these ecotypes.     

Patterns of genetic variation in the adaptive radiation of New World crossbills (Aves: Loxia)

Incipient species groups or young adaptive radiations such as crossbills (Aves: Loxia) present the opportunity to investigate directly the processes occurring during speciation. New World crossbills include white-winged crossbills (Loxia leucoptera), Hispaniolan crossbills (Loxia megaplaga), and red crossbills (Loxia curvirostra complex), the last of which is comprised of at least nine morphologically and vocally differentiated forms ('call types') where divergent natural selection for specialization on different conifer resources has been strongly implicated as driving diversification. Here we use amplified fragment length polymorphism (AFLP) markers to investigate patterns of genetic variation across populations, call types, and species of New World crossbills. Tree-based analyses using 440 AFLP loci reveal strongly supported clustering of the formally recognized species, but did not separate individuals from the eight call types in the red crossbill complex, consistent with recent divergence and ongoing gene flow. Analyses of genetic differentiation based on inferred allele frequency variation however, reveal subtle but significant levels of genetic differentiation among the different call types of the complex and indicate that between call-type differentiation is greater than that found among different geographic locations within call types. Interpreted in light of evidence of divergent natural selection and strong premating reproductive isolation, the observed genetic differentiation suggests restricted gene flow among sympatric call types consistent with the early stages of ecological speciation.     

Specialization and generalization in the diversification of phytophagous insects: tests of the musical chairs and oscillation hypotheses

Evolutionary biologists have often assumed that ecological generalism comes at the expense of less intense exploitation of specific resources and that this trade-off will promote the evolution of ecologically specialized daughter species. Using a phylogenetic comparative approach with butterflies as a model system, we test hypotheses that incorporate changes in niche breadth and location into explanations of the taxonomic diversification of insect herbivores. Specifically, we compare the oscillation hypothesis, where speciation is driven by host-plant generalists giving rise to specialist daughter species, to the musical chairs hypothesis, where speciation is driven by host-plant switching, without changes in niche breadth. Contrary to the predictions of the oscillation hypothesis, we recover a negative relationship between host-plant breadth and diversification rate and find that changes in host breadth are seldom coupled to speciation events. By contrast, we present evidence for a positive relationship between rates of host switching and butterfly diversification, consonant with the musical chairs hypothesis. These results suggest that the costs of trophic generalism in plant-feeding insects may have been overvalued and that transitions from generalists to ecological specialists may not be an important driver of speciation in general.     

Concerted genetic, morphological and ecological diversification in Nacella limpets in the Magellanic Province

Common inhabitants of Antarctic and Subantarctic rocky shores, the limpet genus Nacella, includes 15 nominal species distributed in different provinces of the Southern Ocean. The Magellanic Province represents the area with the highest diversity of the genus. Phylogenetic reconstructions showed an absence of reciprocal monophyly and high levels of genetic identity among nominal species in this Province and therefore imply a recent diversification in southern South America. Because most of these taxa coexist along their distribution range with clear differences in their habitat preferences, Nacella is a suitable model to explore diversification mechanisms in an area highly affected by recurrent Pleistocene continental ice cap advances and retreats. Here, we present genetic and morphological comparisons among sympatric Magellanic nominal species of Nacella. We amplified a fragment of the COI gene for 208 individuals belonging to seven sympatric nominal species and performed geometric morphometric analyses of their shells. We detected a complete congruence between genetic and morphological results, leading us to suggest four groups of Nacella among seven analysed nominal species. Congruently, each of these groups was related to different habitat preferences such as bathymetric range and substrate type. A plausible explanation for these results includes an ecologically based allopatric speciation process in Nacella. Major climatic changes during the Plio-Pleistocene glacial cycles may have enhanced differentiation processes. Finally, our results indicate that the systematics of the group requires a deep revision to re-evaluate the taxonomy of Nacella and to further understand the Pleistocene legacy of the glacial cycles in the southern tip of South America.     

Rapid evolution of cold tolerance in stickleback

Climate change is predicted to lead to increased average temperatures and greater intensity and frequency of high and low temperature extremes, but the evolutionary consequences for biological communities are not well understood. Studies of adaptive evolution of temperature tolerance have typically involved correlative analyses of natural populations or artificial selection experiments in the laboratory. Field experiments are required to provide estimates of the timing and strength of natural selection, enhance understanding of the genetics of adaptation and yield insights into the mechanisms driving evolutionary change. Here, we report the experimental evolution of cold tolerance in natural populations of threespine stickleback fish (Gasterosteus aculeatus). We show that freshwater sticklebacks are able to tolerate lower minimum temperatures than marine sticklebacks and that this difference is heritable. We transplanted marine sticklebacks to freshwater ponds and measured the rate of evolution after three generations in this environment. Cold tolerance evolved at a rate of 0.63 haldanes to a value 2.5°C lower than that of the ancestral population, matching values found in wild freshwater populations. Our results suggest that cold tolerance is under strong selection and that marine sticklebacks carry sufficient genetic variation to adapt to changes in temperature over remarkably short time scales.     

Linking divergent selection on vegetative traits to environmental variation and phenotypic diversification in the Iberian columbines (Aquilegia)

Divergent selection is a key in the ecological theory of adaptive radiation. Most evidence on its causes and consequences relies on studies of pairs of populations or closely related taxa. However, adaptive radiation involves multiple taxa adapted to different environmental factors. We propose an operational definition of divergent selection to explore the continuum between divergent and convergent selection in multiple populations and taxa, and its links with environmental variation and phenotypic and taxonomic differentiation. We apply this approach to explore phenotypic differentiation of vegetative traits between 15 populations of four taxa of Iberian columbines (Gen. Aquilegia). Differences in soil rockiness impose divergent selection on inflorescence height and the number of flowers per inflorescence, likely affecting the processes of phenotypic and, in the case of inflorescence height, taxonomic diversification between taxa. Elevational variation imposes divergent selection on the number of leaves; however, the current pattern of divergent selection on this trait seems related to ecotypic differentiation within taxa but not to their taxonomic diversification.     

The perils of single gene trees — mitochondrial versus single-copy nuclear DNA variation in white-eyes (Aves: Zosteropidae)

Phylogenetic relationships among animal populations and species commonly have been inferred from patterns of variation observed within a single gene system, most often the mitochondrial genome. Analysis of restriction site variation in the mitochondrial DNA of two species of white-eye ( Zosterops lateralis and Z. lutea ) in Australia produced a single gene tree that does not accurately represent the organismal tree. In contrast, patterns of variation at two anonymous, single-copy nuclear DNA loci revealed a phylogeography consistent with traditional classification of the species. Discordance between mitochondrial DNA and single-copy nuclear DNA variation is probably the result of past hybridization between Z. lateralis and Z. lutea , evidence of which has been lost from the nuclear genome by recombination. This study provides a clear empirical demonstration that single gene genealogies cannot be assumed to accurately represent the true phylogenies, and emphasizes the need for composite genetic analyses.     

Niche evolution and diversification in a Neotropical radiation of birds (Aves: Furnariidae)

Rapid diversification may be caused by ecological adaptive radiation via niche divergence. In this model, speciation is coupled with niche divergence and lineage diversification is predicted to be correlated with rates of niche evolution. Studies of the role of niche evolution in diversification have generally focused on ecomorphological diversification but climatic‐niche evolution may also be important. We tested these alternatives using a phylogeny of 298 species of ovenbirds (Aves: Furnariidae). We found that within Furnariidae, variation in species richness and diversification rates of subclades were best predicted by rate of climatic‐niche evolution than ecomorphological evolution. Although both are clearly important, univariate regression and multivariate model averaging more consistently supported the climatic‐niche as the best predictor of lineage diversification. Our study adds to the growing body of evidence, suggesting that climatic‐niche divergence may be an important driver of rapid diversification in addition to ecomorphological evolution. However, this pattern may depend on the phylogenetic scale at which rate heterogeneity is examined.     

Rapid diversification of Tragopogon and ecological associates in Eurasia

Tragopogon comprises approximately 150 described species distributed throughout Eurasia from Ireland and the UK to India and China with a few species in North Africa. Most of the species diversity is found in Eastern Europe to Western Asia. Previous phylogenetic analyses identified several major clades, generally corresponding to recognized taxonomic sections, although relationships both among these clades and among species within clades remain largely unresolved. These patterns are consistent with rapid diversification following the origin of Tragopogon, and this study addresses the timing and rate of diversification in Tragopogon. Using BEAST to simultaneously estimate a phylogeny and divergence times, we estimate the age of a major split and subsequent rapid divergence within Tragopogon to be ~2.6 Ma (and 1.7–5.4 Ma using various clock estimates). Based on the age estimates obtained with BEAST (HPD 1.7–5.4 Ma) for the origin of crown group Tragopogon and 200 estimated species (to accommodate a large number of cryptic species), the diversification rate of Tragopogon is approximately 0.84–2.71 species/Myr for the crown group, assuming low levels of extinction. This estimate is comparable in rate to a rapid Eurasian radiation in Dianthus (0.66–3.89 species/Myr), which occurs in the same or similar habitats. Using available data, we show that subclades of various plant taxa that occur in the same semi‐arid habitats of Eurasia also represent rapid radiations occurring during roughly the same window of time (1.7–5.4 Ma), suggesting similar causal events. However, not all species‐rich plant genera from the same habitats diverged at the same time, or at the same tempo. Radiations of several other clades in this same habitat (e.g. Campanula, Knautia, Scabiosa) occurred at earlier dates (45–4.28 Ma). Existing phylogenetic data and diversification estimates therefore indicate that, although some elements of these semi‐arid communities radiated during the Plio‐Pleistocene period, other clades sharing the same habitat appear to have diversified earlier.     

Systematics and evolution of the Disa draconis complex (Orchidaceae)

The systematic status of Disa draconis (L.f.) Sw. is revised following extensive field studies of population variation in the Western Cape, South Africa. Principal component and cluster analyses revealed clear distinctions between populations from sandplain, semi-arid and montane environments. Diagnostic characters were found in each of the population clusters indicating the existence of a species complex, rather than a single taxon as in the current taxonomy. We propose, therefore, that the name D. draconis (L.f.) Sw. be restricted to the individuals of the sandplain populations from which the type was collected. We reinstate an earlier name, Disa harveiana Lindl., to describe the montane populations which possess several autapomorphic characters. We also show that the geographical variation in spur length and flowering time within this species can be partitioned into two geographically distinct subspecies: D. harveiana subsp. harveiana and D. harveiana subsp. longicalcarata Johnson & Linder. The populations from the semi-arid Karoo region were recognized as a distinct new species, Disa karooica Johnson & Linder, on the basis of their peculiar petal structure. A cladistic analysis indicated that the three species forming the D. draconis complex form a monophyletic and relatively specialized lineage within Disa sect. Coryphaea. The revised classification also has important conservation implications as D. draconis , previously considered a common species, is now restricted to a few highly threatened populations on the west coast near Cape Town.     

Phylogenomics of pike cichlids (Cichlidae: Crenicichla): the rapid ecological speciation of an incipient species flock

The rapid rise of phenotypic and ecological diversity in independent lake‐dwelling groups of cichlids is emblematic of the East African Great Lakes. In this study, we show that similar ecologically based diversification has occurred in pike cichlids (Crenicichla) throughout the Uruguay River drainage of South America. We collected genomic data from nearly 500 ultraconserved element (UCEs) loci and >260 000 base pairs across 33 species, to obtain a phylogenetic hypothesis for the major species groups and to evaluate the relationships and genetic structure among five closely related, endemic, co‐occurring species (the Uruguay River species flock; URSF). Additionally, we evaluated ecological divergence of the URSF based on body and lower pharyngeal jaw (LPJ) shape and gut contents. Across the genus, we recovered novel relationships among the species groups. We found strong support for the monophyly of the URSF; however, relationships among these species remain problematic, likely because of the rapid and recent evolution of this clade. Clustered co‐ancestry analysis recovered most species as well delimited genetic groups. The URSF species exhibit species‐specific body and LPJ shapes associated with specialized trophic roles. Collectively, our results suggest that the URSF consists of incipient species that arose via ecological speciation associated with the exploration of novel trophic roles.     

Links to rare climates do not translate into distinct traits for island endemics

Current models of island biogeography treat endemic and non-endemic species as if they were functionally equivalent, focussing primarily on species richness. Thus, the functional composition of island biotas in relation to island biogeographical variables remains largely unknown. Using plant trait data (plant height, leaf area and flower length) for 895 native species in the Canary Islands, we related functional trait distinctiveness and climate rarity for endemic and non-endemic species and island ages. Endemics showed a link to climatically rare conditions that is consistent with island geological change through time. However, functional trait distinctiveness did not differ between endemics and non-endemics and remained constant with island age. Thus, there is no obvious link between trait distinctiveness and occupancy of rare climates, at least for the traits measured here, suggesting that treating endemic and non-endemic species as functionally equivalent in island biogeography is not fundamentally wrong.     

Parallel evolution of ecomorphological traits in the European whitefish Coregonus lavaretus (L.) species complex during postglacial times

The extensive phenotypic polymorphism in the European whitefish has triggered evolutionary research in order to disentangle mechanisms underlying diversification. To illuminate the ecological distinctiveness in polymorphic whitefish, and evaluate taxonomic designations, we studied nine Norwegian lakes in three watercourses, which each harboured pairs of divergent whitefish morphs. We compared the morphology and life history of these morphs, documented the extent of genetic differentiation between them, and contrasted the niche use of sympatric morphs along both the habitat and resource axes. In all cases, sympatric morphs differed in the number of gill rakers, a highly heritable trait related to trophic utilization. Individual growth rate, age and size at maturity, diet and habitat use also differed between morphs within lakes, but were remarkably similar across lakes within the same morph. Microsatellite analyses confirmed for all but one pair that sympatric morphs were significantly genetically different, and that similar morphs from different lakes likely have a polyphyletic origin. These results are most compatible with the process of parallel evolution through recurrent postglacial divergence into pelagic and benthic niches in each of these lakes. We propose that sparsely and densely rakered whitefish sympatric pairs may be a likely case of ecological speciation, mediated in oligotrophic lakes with few trophic competitors.     

Rapid radiation of Rheum (Polygonaceae) and parallel evolution of morphological traits

In this study, we examined diversification history of Rheum and tested the hypothesis that morphological traits related to plant 'body-plans' evolved in parallel in this genus. We sequenced eight chloroplast DNA fragments (representing more than 8000 bps of the sequence for each species) of 34 species from the genus and 13 species from closely related genera. Phylogenetic analyses indicate that all species of Rheum form a monophyletic lineage sister to the two-species genus Oxyria, indicating that radiative diversifications have occurred in its evolutionary history. Our dating analyses further suggest that these radiations largely coincided with the extensive uplifts of the Qinghai-Tibetan Plateau (QTP). Ancestral state reconstruction and likelihood sensitivity tests strongly indicate that both decumbent and 'glasshouse-like' body-plan traits evolved in parallel in different clades. Our findings highlight the importance of the uplift of the QTP in promoting species diversification and the parallel evolution of morphological traits that are putatively adaptive during such an evolutionary history.     

Historical biogeography and the evolution of the latitudinal gradient of species richness in the Papionini (Primata: Cercopithecidae)

We apply historical biogeography techniques to the macaques, baboons and their relatives (Primata: Papionini) and relate the inferred history of range shifts, and associated evolutionary events, to the latitudinal distribution of extant species, which is strongly tropical. The results of reversible parsimony, weighted ancestral area and dispersal-vicariance analyses all agree that Central Africa was part of the range of the ancestor of the tribe. Tropical regions with high current species richness (Central Africa, South-east Asia, Indonesia) have: (1) had a relatively long history of occupation, (2) experienced both a greater number and a greater average rate of speciation events and (3) given rise to more dispersal events to other regions. However, nested sister-taxon comparisons across the tribe show no overall association between differences in latitude and differences in rates of cladogenesis. Our historical reconstructions are largely consistent with previous hypotheses and fossil data, and suggest that both the passage of time since colonization and rates of cladogenesis have enhanced tropical species richness. Historical biogeography may thus considerably aid understanding of this and other spatial problems in macroecology.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 235–246.