The role of dispersal for shaping phylogeographical structure of flightless beetles from the Andes.
ABSTRACT: Background:Páramo is a tropical alpine ecosystem present in the northern Andes. Its patchy distribution imposes limits and barriers to specialist inhabitants. We aim to assess the effects of this habitat distribution on divergence across two independently flightless ground beetle lineages, in the genera Dyscolus and Dercylus. Methods:One nuclear and one mitochondrial gene from 110 individuals from 10 sites across the two lineages were sequenced and analyzed using a combination of phylogenetics, population genetic analyses, and niche modeling methods. Results:The two lineages show different degrees of population subdivision. Low levels of gene flow were found in Dyscolus alpinus, where one dominant haplotype is found in four out of the six populations analyzed for both molecular markers. However, complete population isolation was revealed in species of the genus Dercylus, where high levels of differentiation exist at species and population level for both genes. Maximum entropy models of species in the Dercylus lineage show overlapping distributions. Still, species distributions appear to be restricted to small areas across the Andes. Conclusion:Even though both beetle lineages are flightless, the dispersal ability of each beetle lineage appears to influence the genetic diversity across fragmented páramo populations, where Dyscolus alpinus appears to be a better disperser than species in the genus Dercylus.
Project description:The ant-loving beetle genus Panabachia Park 1942 is a poorly studied beetle lineage from the new world tropics. We recently collected Panabachia from several previously unrecorded locations in the páramo biome of the high Ecuadorian Andes, with males exhibiting great morphological variation in the distribution of the foveae and depressions in the pronotum, as well as aspects of the male genitalia. Here, we employ phylogenetic and species delimitation methods with mitochondrial (COI) and nuclear protein-coding (wingless) gene sequences to examine the concordance of morphological characters and geography with hypothesized species boundaries. Three methods of species delimitation (bPTP, GMYC and Stacey) were used to estimate the number of species, and divergence times between putative species using molecular clock calibration. Phylogenetic analysis revealed two parallel radiations, and species delimitation analyses suggest there are between 17 and 22 putative species. Based on clade support and concordance across species delimitation methods we hypothesize 17 distinct clusters, with allopatric speciation consistent with most geographic patterns. Additionally, a widespread species appears to be present in northern páramo sites, and some sister species sympatry may indicate other diversification processes have operated on certain lineages of Panabachia. Divergence time estimates suggest that Panabachia originated in the Miocene, but most species analyzed diverged during the Pliocene and Pleistocene (5.3-0.11 Mya), contemporaneous with the evolution of páramo plant species.
Project description:The recent uplift of the tropical Andes (since the late Pliocene or early Pleistocene) provided extensive ecological opportunity for evolutionary radiations. We test for phylogenetic and morphological evidence of adaptive radiation and convergent evolution to novel habitats (exposed, high-altitude páramo habitats) in the Andean fern genera Jamesonia and Eriosorus. We construct time-calibrated phylogenies for the Jamesonia-Eriosorus clade. We then use recent phylogenetic comparative methods to test for evolutionary transitions among habitats, associations between habitat and leaf morphology, and ecologically driven variation in the rate of morphological evolution. Páramo species (Jamesonia) display morphological adaptations consistent with convergent evolution in response to the demands of a highly exposed environment but these adaptations are associated with microhabitat use rather than the páramo per se. Species that are associated with exposed microhabitats (including Jamesonia and Eriorsorus) are characterized by many but short pinnae per frond whereas species occupying sheltered microhabitats (primarily Eriosorus) have few but long pinnae per frond. Pinnae length declines more rapidly with altitude in sheltered species. Rates of speciation are significantly higher among páramo than non-páramo lineages supporting the hypothesis of adaptation and divergence in the unique Páramo biodiversity hotspot.
Project description:The evolution of flight is a key innovation that may enable the extreme diversification of insects. Nonetheless, many species-rich, winged insect groups contain flightless lineages. The loss of flight may promote allopatric differentiation due to limited dispersal power and may result in a high speciation rate in the flightless lineage. Here we show that loss of flight accelerates allopatric speciation using carrion beetles (Coleoptera: Silphidae). We demonstrate that flightless species retain higher genetic differentiation among populations and comprise a higher number of genetically distinct lineages than flight-capable species, and that the speciation rate with the flightless state is twice that with the flight-capable state. Moreover, a meta-analysis of 51 beetle species from 15 families reveals higher genetic differentiation among populations in flightless compared with flight-capable species. In beetles, which represent almost one-fourth of all described species, repeated evolution of flightlessness may have contributed to their steady diversification since the Mesozoic era.
Project description:Nebria (Falcinebria) reflexa Bates (Carabidae), a Japanese endemic flightless ground beetle, is geographically polytypic and was thought to be composed of four subspecies (including nominotypical subspecies). Populations from Honshû recognized as three subspecies were taxonomically revised based primarily on the shape of the endophallus, a membranous inner sac everted from the aedeagus. Three known taxa, Nebria reflexa, Nebria niohozana Bates, and Nebria uenoi Nakane, are redefined based on endophallus morphology, and the latter two are described as distinct species rather than subspecies of N. reflexa. Seven new species are described: N. sagittata sp. nov., N. iidesana sp. nov., N. furcata sp. nov., N. pisciformis sp. nov., N. kuragadakensis sp. nov., N. dichotoma sp. nov., and N. chugokuensis sp. nov. Comparative morphology, primarily of the endophallus, indicated that N. furcata, N. pisciformis, N. kuragadakensis, and N. uenoi form basal lineages, and the remaining six species form a clade in which N. niohozana and N. dichotoma are sister taxa. Species phylogeny and known distributions suggest that the initial diversification of these species occurred in the western Chûbu and eastern Kinki regions of Japan.
Project description:The páramos, high-elevation Andean grasslands ranging from ca. 2800 m to the snow line, harbor one of the fastest evolving biomes worldwide since their appearance in the northern Andes 3-5 million years (Ma) ago. Hypericum (St. John's wort), with over 65% of its Neotropical species, has a center of diversity in these high Mountain ecosystems. Using nuclear rDNA internal transcribed spacer (ITS) sequences of a broad sample of New World Hypericum species we investigate phylogenetic patterns, estimate divergence times, and provide the first insights into diversification rates within the genus in the Neotropics. Two lineages appear to have independently dispersed into South America around 3.5 Ma ago, one of which has radiated in the páramos (Brathys). We find strong support for the polyphyly of section Trigynobrathys, several species of which group within Brathys, while others are found in temperate lowland South America (Trigynobrathys s.str.). All páramo species of Hypericum group in one clade. Within these páramo Hypericum species enormous phenotypic evolution has taken place (life forms from arborescent to prostrate shrubs) evidently in a short time frame. We hypothesize multiple mechanisms to be responsible for the low differentiation in the ITS region contrary to the high morphological diversity found in Hypericum in the páramos. Amongst these may be ongoing hybridization and incomplete lineage sorting, as well as the putative adaptive radiation, which can explain the contrast between phenotypic diversity and the close phylogenetic relationships.
Project description:With the impending threat of climate change, greater understanding of patterns of species distributions and richness and the environmental factors driving them are required for effective conservation efforts. Species distribution models enable us to not only estimate geographic extents of species and subsequent patterns of species richness, but also generate hypotheses regarding environmental factors determining these spatial patterns. Projected changes in climate can then be used to predict future patterns of species distributions and richness. We created distribution models for most of the flightless ground beetles (Carabidae) within the Wet Tropics World Heritage Area of Australia, a major component of regionally endemic invertebrates. Forty-three species were modelled and the environmental correlates of these distributions and resultant patterns of species richness were examined. Flightless ground beetles generally inhabit upland areas characterised by stable, cool and wet environmental conditions. These distribution and richness patterns are best explained using the time-stability hypothesis as this group's primary habitat, upland rainforest, is considered to be the most stable regional habitat. Projected changes in distributions indicate that as upward shifts in distributions occur, species currently confined to lower and drier mountain ranges will be more vulnerable to climate change impacts than those restricted to the highest and wettest mountains. Distribution models under projected future climate change suggest that there will be reductions in range size, population size and species richness under all emission scenarios. Eighty-eight per cent of species modelled are predicted to decline in population size by over 80%, for the most severe emission scenario by the year 2080. These results suggest that flightless ground beetles are among the most vulnerable taxa to climate change impacts so far investigated in the Wet Tropics World Heritage Area. These findings have dramatic implications for all other flightless insect taxa and the future biodiversity of this region.
Project description:<h4>Background</h4>The páramo is a high-elevation biogeographical province in the northern Andes, known for its great biodiversity and ecosystem services. Because there have been very few biogeographic studies encompassing the entire province to date, this study aimed at conducting a phytogeographical regionalisation of the páramo. Specifically, (1) clustering analyses were conducted to identify the main phytogeographical units in the three altitudinal belts: sub-páramo, mid-páramo and super-páramo, and examine their diagnostic flora, (2) an ordination complemented the geo-climatic characterization of the obtained units and (3) a hierarchical classification transformation was obtained to evaluate the relationships between units.<h4>Methods</h4>The study area included the entire Andean páramo range in northern Peru, Ecuador, Colombia and Venezuela. The analyses were based on 1,647 phytosociological plots from the <i>VegPáramo</i> database. The K-means non-hierarchical clustering technique was used to obtain clusters identifiable as phytogeographical units, and the Ochiai fidelity index was calculated to identify their diagnostic species. A principal component analysis was conducted to obtain the geo-climatic characterization of each unit. Finally, the relationships between clusters were traced using a hierarchical plot-based classification.<h4>Results</h4>Fifteen clusters were obtained, 13 natural and two artificial, of which two represented the sub-páramo, nine the mid-páramo and four the super-páramo. Even though data representativeness was a potential limitation to segregate certain sub-páramo and super-páramo units, the overall bioregionalisation was robust and represented important latitudinal, altitudinal and climatic gradients.<h4>Discussion</h4>This study is the first to bioregionalise the páramo province based on a substantial widely distributed biological dataset, and therefore provides important novel scientific insight on its biogeography. The obtained phytogeographical units can be used to support further research on the páramo at smaller scale and on the humid Neotropical high-elevation ecosystems at broader-scale. Finally, several units were highlighted in our results as particularly worthy of further scientific and conservation focus.
Project description:Changes in growth forms frequently accompany plant adaptive radiations, including páramo-a high-elevation treeless habitat type of the northern Andes. We tested whether diverse group of Senecio inhabiting montane forests and páramo represented such growth form changes. We also investigated the role of Andean geography and environment in structuring genetic variation of this group. We sampled 108 populations and 28 species of Senecio (focusing on species from former genera Lasiocephalus and Culcitium) and analyzed their genetic relationships and patterns of intraspecific variation using DNA fingerprinting (AFLPs) and nuclear DNA sequences (ITS). We partitioned genetic variation into environmental and geographical components. ITS-based phylogeny supported monophyly of a Lasiocephalus-Culcitium clade. A grade of herbaceous alpine Senecio species subtended the Lasiocephalus-Culcitium clade suggesting a change from the herbaceous to the woody growth form. Both ITS sequences and the AFLPs separated a group composed of the majority of páramo subshrubs from other group(s) comprising both forest and páramo species of various growth forms. These morphologically variable group(s) further split into clades encompassing both the páramo subshrubs and forest lianas, indicating independent switches among the growth forms and habitats. The finest AFLP genetic structure corresponded to morphologically delimited species except in two independent cases in which patterns of genetic variation instead reflected geography. Several morphologically variable species were genetically admixed, which suggests possible hybrid origins. Latitude and longitude accounted for 5%-8% of genetic variation in each of three AFLP groups, while the proportion of variation attributed to environment varied between 8% and 31% among them. A change from the herbaceous to the woody growth form is suggested for species of high-elevation Andean Senecio. Independent switches between habitats and growth forms likely occurred within the group. Hybridization likely played an important role in species diversification.
Project description:Two new species of Dendrophthora Eichler (Viscaceae) from northwestern Venezuela are described and illustrated. Both mistletoes, D. apiculata Canelón et al., sp. nov. and D. coronata Canelón et al., sp. nov., are confined to subpáramo and páramo ecosystems of the Venezuelan Andes and are, at present, only known from Guaramacal National Park (Portuguesa and Trujillo states). Ecological aspects and possible taxonomic affinities are discussed. A distribution map also is presented.
Project description:Understanding the processes that cause speciation is a key aim of evolutionary biology. Lineages or biomes that exhibit recent and rapid diversification are ideal model systems for determining these processes. Species rich biomes reported to be of relatively recent origin, i.e., since the beginning of the Miocene, include Mediterranean ecosystems such as the California Floristic Province, oceanic islands such as the Hawaiian archipelago and the Neotropical high elevation ecosystem of the Páramos. Páramos constitute grasslands above the forest tree-line (at elevations of c. 2800-4700 m) with high species endemism. Organisms that occupy this ecosystem are a likely product of unique adaptations to an extreme environment that evolved during the last three to five million years when the Andes reached an altitude that was capable of sustaining this type of vegetation. We compared net diversification rates of lineages in fast evolving biomes using 73 dated molecular phylogenies. Based on our sample, we demonstrate that average net diversification rates of Páramo plant lineages are faster than those of other reportedly fast evolving hotspots and that the faster evolving lineages are more likely to be found in Páramos than the other hotspots. Páramos therefore represent the ideal model system for studying diversification processes. Most of the speciation events that we observed in the Páramos (144 out of 177) occurred during the Pleistocene possibly due to the effects of species range contraction and expansion that may have resulted from the well-documented climatic changes during that period. Understanding these effects will assist with efforts to determine how future climatic changes will impact plant populations.