Driver Of Speciation Research Articles

Genetic and morphological divergence in the warm-water planktonic foraminifera genus Globigerinoides.

The planktonic foraminifera genus Globigerinoides provides a prime example of a species-rich genus in which genetic and morphological divergence are uncorrelated. To shed light on the evolutionary processes that lead to the present-day diversity of Globigerinoides, we investigated the genetic, ecological and morphological divergence of its constituent species. We assembled a global collection of single-cell barcode sequences and show that the genus consists of eight distinct genetic types organized in five extant morphospecies. Based on morphological evidence, we reassign the species Globoturborotalita tenella to Globigerinoides and amend Globigerinoides ruber by formally proposing two new subspecies, G. ruber albus n.subsp. and G. ruber ruber in order to express their subspecies level distinction and to replace the informal G. ruber "white" and G. ruber "pink", respectively. The genetic types within G. ruber and Globigerinoides elongatus show a combination of endemism and coexistence, with little evidence for ecological differentiation. CT-scanning and ontogeny analysis reveal that the diagnostic differences in adult morphologies could be explained by alterations of the ontogenetic trajectories towards final (reproductive) size. This indicates that heterochrony may have caused the observed decoupling between genetic and morphological diversification within the genus. We find little evidence for environmental forcing of either the genetic or the morphological diversification, which allude to biotic interactions such as symbiosis, as the driver of speciation in Globigerinoides.

Chromosome-level genome assembly for giant panda provides novel insights into Carnivora chromosome evolution

BackgroundChromosome evolution is an important driver of speciation and species evolution. Previous studies have detected chromosome rearrangement events among different Carnivora species using chromosome painting strategies. However, few of these studies have focused on chromosome evolution at a nucleotide resolution due to the limited availability of chromosome-level Carnivora genomes. Although the de novo genome assembly of the giant panda is available, current short read-based assemblies are limited to moderately sized scaffolds, making the study of chromosome evolution difficult.ResultsHere, we present a chromosome-level giant panda draft genome with a total size of 2.29 Gb. Based on the giant panda genome and published chromosome-level dog and cat genomes, we conduct six large-scale pairwise synteny alignments and identify evolutionary breakpoint regions. Interestingly, gene functional enrichment analysis shows that for all of the three Carnivora genomes, some genes located in evolutionary breakpoint regions are significantly enriched in pathways or terms related to sensory perception of smell. In addition, we find that the sweet receptor gene TAS1R2, which has been proven to be a pseudogene in the cat genome, is located in an evolutionary breakpoint region of the giant panda, suggesting that interchromosomal rearrangement may play a role in the cat TAS1R2 pseudogenization.ConclusionsWe show that the combined strategies employed in this study can be used to generate efficient chromosome-level genome assemblies. Moreover, our comparative genomics analyses provide novel insights into Carnivora chromosome evolution, linking chromosome evolution to functional gene evolution.

Endless forms of sexual selection.

In recent years, the field of sexual selection has exploded, with advances in theoretical and empirical research complementing each other in exciting ways. This perspective piece is the product of a “stock-taking” workshop on sexual selection and sexual conflict. Our aim is to identify and deliberate on outstanding questions and to stimulate discussion rather than provide a comprehensive overview of the entire field. These questions are organized into four thematic sections we deem essential to the field. First we focus on the evolution of mate choice and mating systems. Variation in mate quality can generate both competition and choice in the opposite sex, with implications for the evolution of mating systems. Limitations on mate choice may dictate the importance of direct vs. indirect benefits in mating decisions and consequently, mating systems, especially with regard to polyandry. Second, we focus on how sender and receiver mechanisms shape signal design. Mediation of honest signal content likely depends on integration of temporally variable social and physiological costs that are challenging to measure. We view the neuroethology of sensory and cognitive receiver biases as the main key to signal form and the ‘aesthetic sense’ proposed by Darwin. Since a receiver bias is sufficient to both initiate and drive ornament or armament exaggeration, without a genetically correlated or even coevolving receiver, this may be the appropriate ‘null model’ of sexual selection. Thirdly, we focus on the genetic architecture of sexually selected traits. Despite advances in modern molecular techniques, the number and identity of genes underlying performance, display and secondary sexual traits remains largely unknown. In-depth investigations into the genetic basis of sexual dimorphism in the context of long-term field studies will reveal constraints and trajectories of sexually selected trait evolution. Finally, we focus on sexual selection and conflict as drivers of speciation. Population divergence and speciation are often influenced by an interplay between sexual and natural selection. The extent to which sexual selection promotes or counteracts population divergence may vary depending on the genetic architecture of traits as well as the covariance between mating competition and local adaptation. Additionally, post-copulatory processes, such as selection against heterospecific sperm, may influence the importance of sexual selection in speciation. We propose that efforts to resolve these four themes can catalyze conceptual progress in the field of sexual selection, and we offer potential avenues of research to advance this progress.

A topoclimate model for Quaternary insular speciation

AbstractAimUnderstanding the drivers of speciation within islands is key to explain the high levels of invertebrate diversification and endemism often observed within islands. Here, we propose an insular topoclimate model for Quaternary diversification (ITQD), and test the general prediction that, within a radially eroded conical island, glacial climate conditions facilitate the divergence of populations within species across valleys.LocationGran Canaria, Canary Islands.TaxonThe Laparocerus tessellatus beetle species complex (Coleoptera, Curculionidae).MethodsWe characterize individual‐level genomic relationships using single nucleotide polymorphisms produced by double‐digest restriction site associated DNA sequencing (ddRAD‐seq). A range of parameter values were explored in order to filter our data. We assess individual relatedness, species boundaries, demographic history and spatial patterns of connectivity.ResultsThe total number of ddRAD‐seq loci per sample ranges from 4,576 to 512, with 11.12% and 4.84% of missing data respectively, depending on the filtering parameter combination. We consistently infer four genetically distinct ancestral populations and two presumed cases of admixture, one of which is largely restricted to high altitudes. Bayes factor delimitation support the hypothesis of four species, which is consistent with the four inferred ancestral gene pools. Landscape resistance analyses identified genomic relatedness among individuals in two out of the four inferred species to be best explained by annual precipitation during the last glacial maximum rather than geographic distance.Main conclusionsOur data reveal a complex speciation history involving population isolation and admixture, with broad support for the ITQD model here proposed. We suggest that further studies are needed to test the generality of our model, and enrich our understanding of the evolutionary process in island invertebrates. Our results demonstrate the power of ddRAD‐seq data to provide a detailed understanding of the temporal and spatial dynamics of insular biodiversity.

Phylogeny and biogeography of the African Bathyergidae: a review of patterns and processes.

BackgroundWe review genealogical relationships, biogeographic patterns and broad historical drivers of speciation within the Bathyergidae, a group of endemic African rodents, as well as identify key taxa which need further research.MethodsWe sourced comparable cytochrome b sequence data (comparable data available for all members for the Family) and geographic information for all six genera of the African subterranean rodent. This information was combined into the most comprehensive and geographically representative evolutionary study for the Bathyergidae to date.ResultsSpecies richness within the Bathyergidae appears to be underestimated, with undescribed taxa in five of the six genera. Biogeographic patterns suggest large historical distributions, which were repeatedly fragmented by major landscape changes (especially rifting, uplift and drainage evolution) since the Miocene. Aside from vicariant events, other factors (ecological specialization, population-level responses and climatic change) may have been instrumental in driving divergences in the Bathyergidae. As such, adaptive differences may exist among both populations and species across their discrete ranges, driving independent evolutionary trajectories among taxa. In addition, highly fragmented distributions of divergent (and often relict) lineages indicates the possibility of narrow endemics restricted to diminishing suitable habitats. From this, it is clear that a systematic revision of the Bathyergidae is necessary; such a revision should include comprehensive sampling of all putative taxa, the addition of genomic information to assess adaptive differences, as well as ecological information.

Weak warning signals can persist in the absence of gene flow

Aposematic organisms couple conspicuous warning signals with a secondary defense to deter predators from attacking. Novel signals of aposematic prey are expected to be selected against due to positive frequency-dependent selection. How, then, can novel phenotypes persist after they arise, and why do so many aposematic species exhibit intrapopulation signal variability? Using a polytypic poison frog (Dendrobates tinctorius), we explored the forces of selection on variable aposematic signals using 2 phenotypically distinct (white, yellow) populations. Contrary to expectations, local phenotype was not always better protected compared to novel phenotypes in either population; in the white population, the novel phenotype evoked greater avoidance in natural predators. Despite having a lower quantity of alkaloids, the skin extracts from yellow frogs provoked higher aversive reactions by birds than white frogs in the laboratory, although both populations differed from controls. Similarly, predators learned to avoid the yellow signal faster than the white signal, and generalized their learned avoidance of yellow but not white. We propose that signals that are easily learned and broadly generalized can protect rare, novel signals, and weak warning signals (i.e., signals with poor efficacy and/or poor defense) can persist when gene flow among populations, as in this case, is limited. This provides a mechanism for the persistence of intrapopulation aposematic variation, a likely precursor to polytypism and driver of speciation.

Evolution of aposematism and mimicry in butterflies: Causes, consequences and paradoxes

Evolution of aposematism and mimicry in butterflies: Causes, consequences and paradoxes

Divergence in drought-response traits between sympatric species of Mimulus.

Differential adaptation to local environmental conditions is thought to be an important driver of speciation. Plants, whose sedentary lifestyle necessitates fine‐tuned adaptation to edaphic conditions such as water availability, are often distributed based on these conditions. Populations occupying water‐limited habitats may employ a variety of strategies, involving numerous phenotypes, to prevent and withstand desiccation. In sympatry, two closely related Mimulus species—M. guttatus and M. nasutus—occupy distinct microhabitats that differ in seasonal water availability. In a common garden experiment, we characterized natural variation within and between sympatric M. guttatus and M. nasutus in the ability to successfully set seed under well‐watered and drought conditions. We also measured key phenotypes for drought adaptation, including developmental timing, plant size, flower size, and stomatal density. Consistent with their microhabitat associations in nature, M. nasutus set seed much more successfully than M. guttatus under water‐limited conditions. This divergence in reproductive output under drought was due to differences in mortality after the onset of flowering, with M. nasutus surviving at a much higher rate than M. guttatus. Higher seed set in M. nasutus was mediated, at least in part, by a plastic increase in the rate of late‐stage development (i.e., fruit maturation), consistent with the ability of this species to inhabit more ephemeral habitats in the field. Our results suggest adaptation to water availability may be an important factor in species maintenance of these Mimulus taxa in sympatry.

Advances in the studies of plant diversity and ecological adaptation in the subnival ecosystem of the Qinghai-Tibet Plateau

<p indent=0mm>The alpine subnival belt, located at the front of the alpine life zone, very close to the snowline at high elevations, is the highest habitat of all terrestrial ecosystems. Due to various harsh environmental factors, the alpine subnival belt offers the most hostile conditions for the survival, growth and development of plant species. However, due to its highly varied climates, extremely diverse types of microhabitats, and unique corridor or isolation barrier for plant species, the Qinghai-Tibet Plateau (QTP) hosts the richest alpine subnival plant biodiversity in the world; and the driver for speciation, as well as the mechanisms of sustaining plant diversity in the alpine subnival ecosystem in this region are areas of interest for many researchers. Many recent studies have revealed: Highly specialized morphological features, such as large semi-translucent bracts, dense hairy inflorescences, and nodding flowers that appear in various systematically distant plant species, have multifold functions to protect their vegetative and reproductive organs from high intensity UV radiation, low temperatures, and high humidity, therefore improving the survival rate of these plant species in such a hostile environment. Numerous other researches have reported how advantageous reproductive strategies that have evolved in subnival plant species of the QTP, including the hermaphroditism system, dry fruits, delayed germination behavior and persistent seed bank, promote the probability of reproductive success. In addition, some studies have highlighted the critical role of plant-plant interactions, in particular the facilitation of other plant species by keystone species (e.g. <italic>Arenaria polytrichoides</italic>, <italic>Potentilla articulata </italic>with cushion morphology), and unique plant-pollinator interactions between flagship species (e.g. <italic>Rheum nobile</italic> with large semi-translucent bracts covering inflorescence) and insects on the composition and sustainability of plant diversity in the subnival ecosystem of the QTP. On the other hand, as a region extremely sensitive to climate changes, the subnival habitat has experienced great changes in appearance over the past decades. Recent reports have stated that as a result of warmer climates, the expansion of areas for plant migration and colonization has brought increasing species richness in some subnival areas of the QTP (e.g. the Hengduan Mountains, SE QTP). Yet at the same time, by taking advantage of the continuous rise in temperature, many lowland and middle-elevation plant species will be able to invade this cold-edge ecosystem, causing in a great compression in the niche of some typical and/or endemic subnival plant species and finally resulting in a dramatic decrease in the diversity and functions of the subnival ecosystem. Undoubtedly, the findings obtained from numerous studies on all aspects of terrestrial evolution in recent decades have improved our understanding of ecological adaptation and sustainability of plant diversity in the subnival ecosystem of the QTP. In the future, we still need to focus on: (1) Unveiling the adaptive mechanisms of the striking morphological characteristics of subnival plant species from a molecular and/or genome level; (2) exploring how the buffer effects of the safety net produced by keystone species (e.g. cushion plant) influence global warming on the plant diversity of subnival ecosystem; (3) examining the beneficiary feedback effect on benefactor species and changes in the plant-pollinator co-evolutionary interaction, under global warming and human activity disturbance; (4) investigating how species richness, phenological pattern, and the functional traits of subnival plant species respond to global warming; and (5) testing the importance of plant diversity on the functions and evolution of the subnival ecosystem, and the potential influence of plant diversity loss on the stability and integrity of the subnival ecosystem, through field experiments in permanent plots and historical records of plant collections and vegetation surveys.

Natural selection and repeated patterns of molecular evolution following allopatric divergence.

Although geographic isolation is a leading driver of speciation, the tempo and pattern of divergence at the genomic level remain unclear. We examine genome-wide divergence of putatively single-copy orthologous genes (POGs) in 20 allopatric species/variety pairs from diverse angiosperm clades, with 16 pairs reflecting the classic eastern Asia-eastern North America floristic disjunction. In each pair, >90% of POGs are under purifying selection, and <10% are under positive selection. A set of POGs are under strong positive selection, 14 of which are shared by 10-15 pairs, and one shared by all pairs; 15 POGs are annotated to biological processes responding to various stimuli. The relative abundance of POGs under different selective forces exhibits a repeated pattern among pairs despite an ~10 million-year difference in divergence time. Species divergence times are positively correlated with abundance of POGs under moderate purifying selection, but negatively correlated with abundance of POGs under strong purifying selection.

Cryptic species and independent origins of allochronic populations within a seabird species complex (Hydrobates spp.)

Humans are inherently biased towards naming species based on morphological differences, which can lead to reproductively isolated species being mistakenly classified as one if they are morphologically similar. Recognising cryptic diversity is needed to understand drivers of speciation fully, and for accurate estimates of global biodiversity and assessments for conservation. We investigated cryptic species across the range of band-rumped storm-petrels (Hydrobates spp.): highly pelagic, nocturnal seabirds that breed on tropical and sub-tropical islands in the Atlantic and Pacific Oceans. In many breeding colonies, band-rumped storm-petrels have sympatric but temporally isolated (allochronic) populations; we sampled all breeding locations and allochronic populations. Using mitochondrial control region sequences from 754 birds, cytochrome b sequences from 69 birds, and reduced representation sequencing of the nuclear genomes of 133 birds, we uncovered high levels of genetic structuring. Population genomic analyses revealed up to seven unique clusters, and phylogenomic reconstruction showed that these represent seven monophyletic groups. We uncovered up to six independent breeding season switches across the phylogeny, spanning the continuum from genetically undifferentiated temporal populations to full allochronic species. Thus, band-rumped storm-petrels encompass multiple cryptic species, with non-geographic barriers potentially comprising strong barriers to gene flow.

Genetic and Genomic Analyses Reveal Boundaries between Species Closely Related to Cryptococcus Pathogens.

Speciation is a central mechanism of biological diversification. While speciation is well studied in plants and animals, in comparison, relatively little is known about speciation in fungi. One fungal model is the Cryptococcus genus, which is best known for the pathogenic Cryptococcus neoformans/Cryptococcus gattii species complex that causes >200,000 new human infections annually. Elucidation of how these species evolved into important human-pathogenic species remains challenging and can be advanced by studying the most closely related nonpathogenic species, Cryptococcus amylolentus and Tsuchiyaea wingfieldii However, these species have only four known isolates, and available data were insufficient to determine species boundaries within this group. By analyzing full-length chromosome assemblies, we reappraised the phylogenetic relationships of the four available strains, confirmed the genetic separation of C. amylolentus and T. wingfieldii (now Cryptococcus wingfieldii), and revealed an additional cryptic species, for which the name Cryptococcus floricola is proposed. The genomes of the three species are ∼6% divergent and exhibit significant chromosomal rearrangements, including inversions and a reciprocal translocation that involved intercentromeric ectopic recombination, which together likely impose significant barriers to genetic exchange. Using genetic crosses, we show that while C. wingfieldii cannot interbreed with any of the other strains, C. floricola can still undergo sexual reproduction with C. amylolentus However, most of the resulting spores were inviable or sterile or showed reduced recombination during meiosis, indicating that intrinsic postzygotic barriers had been established. Our study and genomic data will foster additional studies addressing fungal speciation and transitions between nonpathogenic and pathogenic Cryptococcus lineages.IMPORTANCE The evolutionary drivers of speciation are critical to our understanding of how new pathogens arise from nonpathogenic lineages and adapt to new environments. Here we focus on the Cryptococcus amylolentus species complex, a nonpathogenic fungal lineage closely related to the human-pathogenic Cryptococcus neoformans/Cryptococcus gattii complex. Using genetic and genomic analyses, we reexamined the species boundaries of four available isolates within the C. amylolentus complex and revealed three genetically isolated species. Their genomes are ∼6% divergent and exhibit chromosome rearrangements, including translocations and small-scale inversions. Although two of the species (C. amylolentus and newly described C. floricola) were still able to interbreed, the resulting hybrid progeny were usually inviable or sterile, indicating that barriers to reproduction had already been established. These results advance our understanding of speciation in fungi and highlight the power of genomics in assisting our ability to correctly identify and discriminate fungal species.

Trait‐based range expansion aided in the global radiation of Crocodylidae

AbstractAimAlmost half of the extant species of Crocodylia (nine genera, sensu Benton &amp; Clark) belong to the genus Crocodylus, which originated in the Miocene. Today, this genus has a circumtropical distribution, with representatives found in Asia, Oceania, Africa and the Neotropics. However, its geographical origin and the historical events behind its rapid diversification and global radiation are still debated. Here, we inferred the evolution of a set of life traits that aid in dispersal and how they influenced the biogeography of Crocodylus.LocationGlobal.Time periodCretaceous to the present.Major taxa studiedCrocodylidae.MethodsWe estimated biogeographical history on a published phylogeny using probabilistic biogeography models. Next, we identified four life traits likely to promote range expansion and used a trait‐based dispersal model jointly to infer evolution of these traits and their influence on the dispersal of crocodilians. Finally, we used diversification analyses to identify shifts in diversification rates.ResultsAn Asian origin was reconstructed for Crocodylus. Despite the small size of the phylogeny, statistical model comparison reports substantially improved the model fit of trait‐dependent over trait‐independent dispersal models. In individual tests, the four life traits considered (salt tolerance, large size, large clutches and habitat generalism) appear to be correlated statistically with higher dispersal probabilities. However, the traits are likely to be correlated with each other, and we hypothesize that they all reflect the gradual evolution of a range‐expansion phenotype in early Crocodylus that resulted in increased diversification of the clade.Main conclusionsIncreased dispersal in the genus Crocodylus was associated with the gradual evolution of a range‐expansion phenotype. Interestingly, the evolution of the range‐expansion phenotype was also associated with the diversification of the genus in a period of global crocodilian extinction, indicating that range expansion might have served as a potential driver of speciation. This suggests that the concept of “spatial sorting”, normally applied at the population genetic level at the leading edge of an expanding population, might also act on a macroevolutionary scale.

Genome Sequences Provide Insights into the Reticulate Origin and Unique Traits of Woody Bamboos

Polyploidization is a major driver of speciation and its importance to plant evolution has been well recognized. Bamboos comprise one diploid herbaceous and three polyploid woody lineages, and are members of the only major subfamily in grasses that diversified in forests, with the woody members having a tree-like lignified culm. In this study, we generated four draft genome assemblies of major bamboo lineages with three different ploidy levels (diploid, tetraploid, and hexaploid). We also constructed a high-density genetic linkage map for a hexaploid species of bamboo, and used a linkage-map-based strategy for genome assembly and identification of subgenomes in polyploids. Further phylogenomic analyses using a large dataset of syntenic genes with expected copies based on ploidy levels revealed that woody bamboos originated subsequent to the divergence of the herbaceous bamboo lineage, and experienced complex reticulate evolution through three independent allopolyploid events involving four extinct diploid ancestors. A shared but distinct subgenome was identified in all polyploid forms, and the progenitor of this subgenome could have been critical in ancient polyploidizations and the origin of woody bamboos. Important genetic clues to the unique flowering behavior and woody trait in bamboos were also found. Taken together, our study provides significant insights into ancient reticulate evolution at the subgenome level in the absence of extant donor species, and offers a potential model scenario for broad-scale study of angiosperm origination by allopolyploidization.

Hybridisation, introgression events and cryptic speciation in Stipa (Poaceae): A case study of the Stipa heptapotamica hybrid-complex

Hybridisation, considered one of the major drivers of speciation in plants, commonly occurs among many genera of grasses, including the genus Stipa L. one of the largest in the family Poaceae in the Old World. In the course of research on Stipa, we studied S. richteriana, S. lessingiana, and their putative hybrid S. heptapotamica, all of which grow in south-eastern Kazakhstan. Stipa heptapotamica is treated as a local endemic, known only from the eastern part of the country. We used an integrative approach, combining morphological and molecular data, to validate the status of the hybrid taxon and present for the first time clear data documenting natural hybridisation in Stipa. The macro- and micromorphological analyses presented here show S. heptapotamica to be intermediate to its parental species in the characters studied; pollen analysis demonstrates low viability of pollen grains (below 50% compared to 94% and 87% in S. lessingiana and S. richteriana, respectively); molecular analyses based on inter simple sequence repeat markers (ISSR) and next-generation sequencing (NGS) data prove that S. heptapotamica originated strictly from hybridisation events involving S. richteriana and S. lessingiana. ISSR markers reveal possible introgression events between S. lessingiana and S. heptapotamica, whereas NGS data analyses confirm the maternal inheritance of a plastome from S. lessingiana in S. heptapotamica and support the finding of cryptic lineages within S. richteriana.

Recurrent changes in cichlid dentition linked to climate‐driven lake‐level fluctuations

AbstractClimate‐induced habitat change has often been invoked as an important driver of speciation and evolutionary radiation in cichlid fishes, yet studies linking morphological change directly to long‐term environmental fluctuations are scarce. Here, we track changes through time in the oral dentition of Oreochromis hunteri, the endemic and only indigenous fish species inhabiting the East African crater Lake Chala (Kenya/Tanzania), in relation to climate‐driven lake‐level fluctuations during the last 25,000 yr. Fossil teeth were recovered from six periods representing alternating high and low lake‐level stands associated with late‐Pleistocene and Holocene trends in rainfall and drought. To construct a reference framework for morphological variation in fossil assemblages, we first analyzed the variability in oral tooth morphology within the modern‐day fish population. These analyses established that, like other cichlids, O. hunteri gradually replace their bicuspid/tricuspid oral dentition with more unicuspid teeth as they grow. In the fossil assemblages, we found systematic and recurrent changes in the relative abundance of these oral tooth types, with a higher proportion of unicuspids during low‐stands. Moreover, O. hunteri living during low‐stands systematically developed unicuspid dentition at a smaller body size, compared with conspecifics living during high‐stands. Considering that low‐stand conditions created a sizable area of oxygenated soft‐bottom habitat that is lacking during high‐stands, we propose that the associated change in oral dentition reflects the fishes’ exploitation of this new habitat either for feeding or for reproduction. The recurrent nature of the observed shifts provides evidence for the ability of O. hunteri to systematically adapt to local habitat change, and strongly suggests that morphological change in oral dentition promoted its long‐term population persistence in an aquatic ecosystem presenting the dual challenge of being both sensitive to climate change (creating constantly shifting selective pressures) and isolated from surrounding populations (limiting gene flow). In Lake Chala, ancient climate‐driven lake‐level fluctuations did not directly promote speciation in O. hunteri, but the population bottlenecks they may have caused potentially contributed to its divergence from sister species Oreochromis jipe.

Tracking temporal shifts in area, biomes, and pollinators in the radiation of Salvia (sages) across continents: leveraging anchored hybrid enrichment and targeted sequence data.

Premise of the StudyA key question in evolutionary biology is why some clades are more successful by being widespread geographically, biome diverse, or species‐rich. To extend understanding of how shifts in area, biomes, and pollinators impact diversification in plants, we examined the relationships of these shifts to diversification across the mega‐genus Salvia.MethodsA chronogram was developed from a supermatrix of anchored hybrid enrichment genomic data and targeted sequence data for over 500 of the nearly 1000 Salvia species. Ancestral areas and biomes were reconstructed using BioGeoBEARS. Pollinator guilds were scored, ancestral pollinators determined, shifts in pollinator guilds identified, and rates of pollinator switches compared.Key ResultsA well‐resolved phylogenetic backbone of Salvia and updated subgeneric designations are presented. Salvia originated in Southwest Asia in the Oligocene and subsequently dispersed worldwide. Biome shifts are frequent from a likely ancestral lineage utilizing broadleaf and/or coniferous forests and/or arid shrublands. None of the four species diversification shifts are correlated to shifts in biomes. Shifts in pollination system are not correlated to species diversification shifts, except for one hummingbird shift that precedes a major shift in diversification near the crown of New World subgen. Calosphace. Multiple reversals back to bee pollination occurred within this hummingbird clade.Conclusions Salvia diversified extensively in different continents, biomes, and with both bee and bird pollinators. The lack of tight correlation of area, biome, and most pollinator shifts to the four documented species diversification shifts points to other important drivers of speciation in Salvia.

Mitogenomes Uncover Extinct Penguin Taxa and Reveal Island Formation as a Key Driver of Speciation.

The emergence of islands has been linked to spectacular radiations of diverse organisms. Although penguins spend much of their lives at sea, they rely on land for nesting, and a high proportion of extant species are endemic to geologically young islands. Islands may thus have been crucial to the evolutionary diversification of penguins. We test this hypothesis using a fossil-calibrated phylogeny of mitochondrial genomes (mitogenomes) from all extant and recently extinct penguin taxa. Our temporal analysis demonstrates that numerous recent island-endemic penguin taxa diverged following the formation of their islands during the Plio-Pleistocene, including the Galápagos (Galápagos Islands), northern rockhopper (Gough Island), erect-crested (Antipodes Islands), Snares crested (Snares) and royal (Macquarie Island) penguins. Our analysis also reveals two new recently extinct island-endemic penguin taxa from New Zealand's Chatham Islands: Eudyptes warhami sp. nov. and a dwarf subspecies of the yellow-eyed penguin, Megadyptes antipodes richdalei ssp. nov. Eudyptes warhami diverged from the Antipodes Islands erect-crested penguin between 1.1 and 2.5Ma, shortly after the emergence of the Chatham Islands (∼3Ma). This new finding of recently evolved taxa on this young archipelago provides further evidence that the radiation of penguins over the last 5 Ma has been linked to island emergence. Mitogenomic analyses of all penguin species, and the discovery of two new extinct penguin taxa, highlight the importance of island formation in the diversification of penguins, as well as the extent to which anthropogenic extinctions have affected island-endemic taxa across the Southern Hemisphere's isolated archipelagos.

Sphagnum ×lydiae, the first allotriploid peatmoss in the northern hemisphere

In plants, hybridization between interspecific taxa leading to chromosome doubling seems to happen regularly and may be an important driver of speciation. Allopolyploid species often have morphological traits from both parents, broader ecological niches, and are hypothesized to be more common going northwards. About 20% of all Sphagnum species have been inferred to be allopolyploids. Here we describe the first allotriploid Sphagnum species in the northern hemisphere, using molecular data, Feulgen densitometry, and morphological examination. Our molecular data also supports the finding of the first polyploid species with inter-subgeneric origin between Acutifolia and Subsecunda. Morphological examination further supports this finding. The Acutifolia progenitor is likely the maternal parent of the new species, named S. ×lydiae, and it is probably S. incundum, S. ×arcticum or S. ×olafii. The Subsecunda parent could not be identified based on molecular and morphological characters. The currently known distribution of S. ×lydiae is confined to regions with arctic and subarctic climate and tundra vegetation in northern Russia.

Ecological divergence plays an important role in strong but complex reproductive isolation in campions (Silene).

New species arise through the evolution of reproductive barriers between formerly interbreeding lineages. Yet, comprehensive assessments of potential reproductive barriers, which are needed to make inferences on processes driving speciation, are only available for a limited number of systems. In this study, we estimated individual and cumulative strengths of seven prezygotic and six postzygotic reproductive barriers between the recently diverged taxa Silene dioica (L.) Clairv. and S. latifolia Poiret using both published and new data. A combination of multiple partial reproductive barriers resulted in near-complete reproductive isolation between S. dioica and S. latifolia, consistent with earlier estimates of gene flow between the taxa. Extrinsic barriers associated with adaptive ecological divergence were most important, while intrinsic postzygotic barriers had moderate individual strength but contributed only little to total reproductive isolation. These findings are in line with ecological divergence as driver of speciation. We further found extensive variation in extrinsic reproductive isolation, ranging from sites with very strong selection against migrants and hybrids to intermediate sites where substantial hybridization is possible. This situation may allow for, or even promote, heterogeneous genetic divergence.