Reconciling phylogeography and ecological niche models for New Zealand beetles: Looking beyond glacial refugia

Mitochondrial DNA (cox1) sequence data and recently developed coalescent phylogeography models were used to construct geo-spatial histories for the New Zealand fungus beetles Epistranus lawsoni and Pristoderus bakewelli (Zopheridae). These methods utilize continuous-time Markov chains and Bayesian stochastic search variable selection incorporated in BEAST to identify historical dispersal patterns via ancestral state reconstruction. Ecological niche models (ENMs) were incorporated to reconstruct the potential geographic distribution of each species during the Last Glacial Maximum (LGM). Coalescent analyses suggest a North Island origin for E. lawsoni, with gene flow predominately north-south between adjacent regions. ENMs for E. lawsoni indicated glacial refugia in coastal regions of both main islands, consistent with phylogenetic patterns but at odds with the coalescent dates, which implicate much older topographic events. Dispersal matrices revealed patterns of gene flow consistent with projected refugia, suggesting long-term South Island survival with population vicariance around the Southern Alps. Phylogeographic relationships are more ambiguous for P. bakewelli, although long-term survival on both main islands is evident. Divergence dates for both species are consistent with the topographic evolution of New Zealand over the last 10Ma, whereas the signature of the LGM is less apparent in the time-scaled phylogeny.     

Glacial refugia,recolonization patterns and diversification forces in Alpine‐endemic Megabunus harvestmen

The Pleistocene climatic fluctuations had a huge impact on all life forms, and various hypotheses regarding the survival of organisms during glacial periods have been postulated. In the European Alps, evidence has been found in support of refugia outside the ice shield (massifs de refuge) acting as sources for postglacial recolonization of inner‐Alpine areas. In contrast, evidence for survival on nunataks, ice‐free areas above the glacier, remains scarce. Here, we combine multivariate genetic analyses with ecological niche models (ENMs) through multiple timescales to elucidate the history of Alpine Megabunus harvestmen throughout the ice ages, a genus that comprises eight high‐altitude endemics. ENMs suggest two types of refugia throughout the last glacial maximum, inner‐Alpine survival on nunataks for four species and peripheral refugia for further four species. In some geographic regions, the patterns of genetic variation are consistent with long‐distance dispersal out of massifs de refuge, repeatedly coupled with geographic parthenogenesis. In other regions, long‐term persistence in nunataks may dominate the patterns of genetic divergence. Overall, our results suggest that glacial cycles contributed to allopatric diversification in Alpine Megabunus, both within and at the margins of the ice shield. These findings exemplify the power of ENM projections coupled with genetic analyses to identify hypotheses about the position and the number of glacial refugia and thus to evaluate the role of Pleistocene glaciations in driving species‐specific responses of recolonization or persistence that may have contributed to observed patterns of biodiversity.     

The demographic history of the New Zealand short-tailed bat Mystacina tuberculata inferred from modified control region sequences

Short-tailed bats Mystacina tuberculata were widespread throughout the forest that dominated prehuman New Zealand, but extensive deforestation has restricted them to scattered populations in forest fragments. In a previous study, the species' intraspecific phylogeny was investigated using multiple mitochondrial gene sequences. Six phylogroups were identified with estimated divergences of 0.93-0.68 Ma. In the current study, the phylogeographical structure and demographic history of the phylogroups were investigated using control region sequences modified by removing homoplasic sites. Phylogeographical structure in the North Island was generally consistent with an isolation-by-distance dispersal model. Coalescent-based analyses (i.e. mismatch distributions, skyline plots, lineage dispersal analysis and nested clade analysis) indicated that the three phylogroups found in central and southern North Island expanded before the last glacial maximum, presumably during interstadials when Nothofagus forest was most extensive. Genetic structure within a central North Island hybrid zone was consistent with range expansion from separate refugia following reforestation after catastrophic volcanic eruptions. Phylogeographical structure in the South Island was consistent with southern populations originating during rapid southward range expansion from refugia in northern South Island following postglacial reforestation of the South Island 10-9 kya.     

Glacial refugia in a maritime temperate climate: Cicada (Kikihia subalpina) mtDNA phylogeography in New Zealand

Understanding the biological significance of Pleistocene glaciations requires knowledge of the nature and extent of habitat refugia during glacial maxima. An opportunity to examine evidence of glacial forest refugia in a maritime, Southern Hemisphere setting is found in New Zealand, where the extent of Pleistocene forests remains controversial. We used the mitochondrial phylogeography of a forest-edge cicada ( Kikihia subalpina ) to test the hypothesis that populations of this species survived throughout South Island during the Last Glacial Maximum. We also compared mitochondrial DNA phylogeographic patterns with male song patterns that suggest allopatric divergence across Cook Strait. Cytochrome oxidase I and II sequences were analyzed using network analysis, maximum-likelihood phylogenetic estimation, Bayesian dating and Bayesian skyline plots. K. subalpina haplotypes from North Island and South Island form monophyletic clades that are concordant with song patterns. Song divergence corresponds to approximately 2% genetic divergence, and Bayesian dating suggests that the North Island and South Island population-lineages became isolated around 761 000 years bp . Almost all South Island genetic variation is found in the north of the island, consistent with refugia in Marlborough Sounds, central Nelson and northwest Nelson. All central and southern South Island and Stewart Island haplotypes are extremely similar to northern South Island haplotypes, a 'northern richness/southern purity' pattern that mirrors genetic patterns observed in many Northern Hemisphere taxa. Proposed southern South Island forest habitat fragments may have been too small to sustain populations of K. subalpina , and/or they may have harboured ecological communities with no modern-day analogues.     

Idiosyncratic responses to climate‐driven forest fragmentation and marine incursions in reed frogs from Central Africa and the Gulf of Guinea Islands

Organismal traits interact with environmental variation to mediate how species respond to shared landscapes. Thus, differences in traits related to dispersal ability or physiological tolerance may result in phylogeographic discordance among co‐distributed taxa, even when they are responding to common barriers. We quantified climatic suitability and stability, and phylogeographic divergence within three reed frog species complexes across the Guineo‐Congolian forests and Gulf of Guinea archipelago of Central Africa to investigate how they responded to a shared climatic and geological history. Our species‐specific estimates of climatic suitability through time are consistent with temporal and spatial heterogeneity in diversification among the species complexes, indicating that differences in ecological breadth may partly explain these idiosyncratic patterns. Likewise, we demonstrated that fluctuating sea levels periodically exposed a land bridge connecting Bioko Island with the mainland Guineo‐Congolian forest and that habitats across the exposed land bridge likely enabled dispersal in some species, but not in others. We did not find evidence that rivers are biogeographic barriers across any of the species complexes. Despite marked differences in the geographic extent of stable climates and temporal estimates of divergence among the species complexes, we recovered a shared pattern of intermittent climatic suitability with recent population connectivity and demographic expansion across the Congo Basin. This pattern supports the hypothesis that genetic exchange across the Congo Basin during humid periods, followed by vicariance during arid periods, has shaped regional diversity. Finally, we identified many distinct lineages among our focal taxa, some of which may reflect incipient or unrecognized species.     

Phylogeography and ecological niche modelling implicate coastal refugia and trans‐alpine dispersal of a New Zealand fungus beetle

The Last Glacial Maximum (LGM) severely restricted forest ecosystems on New Zealand’s South Island, but the extent of LGM distribution for forest species is still poorly understood. We used mitochondrial DNA phylogeography (COI) and ecological niche modelling (ENM) to identify LGM refugia for the mycophagous beetle Agyrtodes labralis (Leiodidae), a forest edge species widely distributed in the South Island. Both the phylogenetic analyses and the ENM indicate that A. labralis refuged in Kaikoura, Nelson, and along much of the South Island’s west coast. Phylogeography of this species indicates that recolonization of the largely deforested east and southeast South Island occurred in a west–east direction, with populations moving through the Southern Alps, and that the northern refugia participated little in interglacial population expansion. This contradicts published studies of other New Zealand species, in which recolonization occurs in a north–south fashion from many of the same refugia.     

Resilient forest faunal communities in South Africa: a legacy of palaeoclimatic change and extinction filtering?

Aim To examine the influence of climatic extinction filtering during the last glacial maximum (LGM; c. 18,000 yr bp ) and of the subsequent recolonization of forest faunas on contemporary assemblage composition in southern African forests. Location South Africa, Mozambique, Swaziland, Zimbabwe. Methods Data comprised presence/absence by quarter‐degree grid cell for forest‐dependent and forest‐associated birds, non‐volant mammals and frogs. Twenty‐one forest subregions were assigned to one of three previously identified forest types: Afrotemperate, scarp, and Indian Ocean coastal belt. Differences among forest types were examined through patterns and gradients of species richness and endemism, assemblage similarity, species turnover, and coefficients of species dispersal direction. The influence of contemporary environment on assemblage composition was investigated using partial canonical correspondence analysis. Several alternative biogeographical hypotheses for the recolonization of forest faunas were tested. Results Afrotemperate faunas are relatively species‐poor, have low species turnover, and are unsaturated and infiltrated by generalist species. In northern and central regions, communities are supplemented by recolonization from scarp forest refugia, and among frogs by autochthanous speciation in localized refugia. Scarp faunas are relatively species‐rich, contain many forest‐dependent species, have high species turnover, and overlap with coastal and Afrotemperate faunas. Coastal forests are relatively species‐rich with high species turnover. Main conclusions Afrotemperate communities were affected most by climatic extinction filtering events. Scarp forests were Afrotemperate refugia during the LGM and are a contemporary overlap zone between Afrotemperate and coastal forest. Coastal faunas derive from post‐LGM colonization along the eastern seaboard from tropical East African refugia. The greatest diversity is achieved in scarp and coastal forest faunas in northern KwaZulu–Natal province. This historical centre of diversity has influenced the faunal diversity of nearly all other forests in South Africa. The response of vertebrate taxa to large‐scale, historical processes is dependent on their relative mobility: forest birds best illustrate patterns resulting from post‐glacial faunal dispersal, while among mammals and frogs the legacy of climatic extinction filtering remains stronger.     

Identifying glacial refugia in a geographic parthenogen using palaeoclimate modelling and phylogeography: the New Zealand stick insect Argosarchus horridus (White)

We have used phylogeographic analysis of mitochondrial DNA (COI and COII genes) and ecological niche modelling (ENM) to reconstruct the population history of Argosarchus horridus (White), a widespread species of New Zealand stick insect. These data were used to address outstanding questions on the role of glacial refugia in determining the distribution and genetic structure of New Zealand species. Phylogeographic analysis shows a general pattern of high diversity in upper North Island and reduced diversity in lower North Island and South Island. The ENM indicates that during the last glacial maximum, A. horridus was largely restricted to refugia around coastal areas of North Island. The ENM also suggests refugia on the northeast coast of South Island and southeast coast of North Island and this prediction is verified by phylogeographic analysis, which shows a clade restricted to this region. Argosarchus horridus is also most likely a geographic parthenogen where males are much rarer at higher latitudes. The higher levels of genetic variation in northern, bisexual populations suggest southern and largely unisexual populations originated from southwardly expanding parthenogenetic lineages. Bayesian skyline analysis also provides support for a recent population size increase consistent with a large increase in geographic distribution in the late Pleistocene. These results exemplify the utility of integrating ENM and phylogeographic analysis in testing hypotheses on the origin of geographic parthenogenesis and effects of Pleistocene environmental change on biodiversity.     

Postglacial recolonization of North America by spadefoot toads: integrating niche and corridor modeling to study species’ range dynamics over geologic time

Understanding the factors that shape species’ distributions is a key topic in biogeography. As climates change, species can either cope with these changes through evolution, plasticity or by shifting their ranges to track the optimal climatic conditions. Ecological niche modeling (ENM) is a widespread technique in biogeography that estimates the niche of the organism by using occurrences and environmental data to estimate species’ potential distributions. ENMs are often criticized for failing to take species’ dispersal abilities into consideration. Here, we attempt to fill this gap by combining ENMs with dispersal and corridor modeling to study the range dynamics of North American spadefoot toads (Scaphiopodidae) over the Holocene. We first estimated the current and past distributions of spadefoot toads and then estimated their past distributions from the Last Glacial Maximum (LGM) to the present day. Then, we estimated how each taxon recolonized North American by using dispersal and corridor modeling. By combining these two modeling approaches we were able to 1) estimate the LGM refugia used by the North American spadefoot toads, 2) further refine these projections by estimating which of the putative LGM refugia contributed to the recolonization of North America via dispersal, and 3) estimate the relative influence of each LGM refugium to the current species’ distributions. The models were tested using previously published phylogeographic data, revealing a high degree of congruence between our models and the genetic data. These results suggest that combining ENMs and dispersal modeling over time is a promising approach to investigate both historical and future species’ range dynamics.     

Post-glacial range formation of temperate forest understorey herbs – Insights from a spatio-temporally explicit modelling approach

Aim

Our knowledge of Pleistocene refugia and post-glacial recolonization routes of forest understorey plants is still very limited. The geographical ranges of these species are often rather narrow and show highly idiosyncratic, often fragmented patterns indicating either narrow and species-specific ecological tolerances or strong dispersal limitations. However, the relative roles of these factors are inherently difficult to disentangle.

Location

Central and south-eastern Europe.

Time period

17,100 BP – present.

Major taxa studied

Five understorey herbs of European beech forests: Aposeris foetida, Cardamine trifolia, Euphorbia carniolica, Hacquetia epipactis and Helleborus niger.

Methods

We used spatio-temporally explicit modelling to reconstruct the post-glacial range dynamics of the five forest understorey herbs. We varied niche requirements, demographic rates and dispersal abilities across plausible ranges and simulated the spread of species from potential Pleistocene refugia identified by phylogeographical analyses. Then we identified the parameter settings allowing for the most accurate reconstruction of their current geographical ranges.

Results

We found a largely homogenous pattern of optimal parameter settings among species. Broad ecological niches had to be combined with very low but non-zero rates of long-distance dispersal via chance events and low rates of seed dispersal over moderate distances by standard dispersal vectors. However, long-distance dispersal events, although rare, led to high variation among replicated simulation runs.

Main conclusions

Small and fragmented ranges of many forest understorey species are best explained by a combination of broad ecological niches and rare medium- and long-distance dispersal events. Stochasticity is thus an important determinant of current species ranges, explaining the idiosyncratic distribution patterns of the study species despite strong similarities in refugia, ecological tolerances and dispersal abilities.     

Ancient forest fragmentation or recent radiation? Testing refugial speciation models in chameleons within an African biodiversity hotspot

Aim East Africa is one of the most biologically diverse regions, especially in terms of endemism and species richness. Hypotheses put forward to explain this high diversity invoke a role for forest refugia through: (1) accumulation of new species due to radiation within refugial habitats, or (2) retention of older palaeoendemic species in stable refugia. We tested these alternative hypotheses using data for a diverse genus of East African forest chameleons, Kinyongia. Location East Africa. Methods We constructed a dated phylogeny for Kinyongia using one nuclear and two mitochondrial markers. We identified areas of high phylogenetic diversity (PD) and evolutionary diversity (ED), and mapped ancestral areas to ascertain whether lineage diversification could best be explained by vicariance or dispersal. Results Vicariance best explains the present biogeographic patterns, with divergence between three major Kinyongia clades (Albertine Rift, southern Eastern Arc, northern Eastern Arc) in the early Miocene/Oligocene (> 20 Ma). Lineage diversification within these clades pre‐dates the Pliocene (> 6 Ma). These dates are much older than the Plio‐Pleistocene climatic shifts associated with cladogenesis in other East African taxa (e.g. birds), and instead point to a scenario whereby palaeoendemics are retained in refugia, rather than more recent radiations within refugia. Estimates of PD show that diversity was highest in the Uluguru, Nguru and East Usambara Mountains and several lineages (from Mount Kenya, South Pare and the Uluguru Mountains) stand out as being evolutionarily distinct as a result of isolation in forest refugia. PD was lower than expected by chance, suggesting that the phylogenetic signal is influenced by an unusually low number of extant lineages with long branch lengths, which is probably due to the retention of palaeoendemic lineages. Main conclusions The biogeographic patterns associated with Kinyongia are the result of long evolutionary histories in isolation. The phylogeny is dominated by ancient lineages whose origins date back to the early Miocene/Oligocene as a result of continental wide forest fragmentation and contraction due to long term climatic changes in Africa. The maintenance of palaeoendemic lineages in refugia has contributed substantially to the remarkably high biodiversity of East Africa.     

Historical biogeography and phenotype‐phylogeny of Chroodiscus (lichenized Ascomycota: Ostropales: Graphidaceae)

Aim To analyse the historical biogeography of the lichen genus Chroodiscus using a phenotype‐based phylogeny in the context of continental drift and evolution of tropical rain forest vegetation. Location All tropical regions (Central and South America, Africa, India, Southeast Asia, north‐east Australia). Methods We performed a phenotype‐based phylogenetic analysis and ancestral character state reconstruction of 14 species of the lichen genus Chroodiscus, using paup * and mesquite ; dispersal–vicariance analysis (DIVA) and dispersal–extinction–cladogenesis (DEC) modelling to trace the geographical origin of individual clades; and ordination and clustering by means of pc‐ord , based on a novel similarity index, to visualize the biogeographical relationships of floristic regions in which Chroodiscus occurs. Results The 14 species of Chroodiscus show distinctive distribution patterns, with one pantropical and one amphi‐Pacific taxon and 12 species each restricted to a single continent. The genus comprises four clades. DIVA and DEC modelling suggest a South American origin of Chroodiscus in the mid to late Cretaceous (120–100 Ma), with subsequent expansion through a South American–African–Indian–Southeast Asian–Australian dispersal route and late diversification of the argillaceus clade in Southeast Asia. Based on the abundance of extant taxa, the probability of speciation events in Chroodiscus is shown to be extremely low. Slow dispersal of foliicolous rain forest understorey lichens is consistent with estimated phylogenetic ages of individual species and with average lengths of biological species intervals in fungi (10–20 Myr). Main conclusions The present‐day distribution of Chroodiscus can be explained by vicariance and mid‐distance dispersal through the interconnection or proximity of continental shelves, without the need for recent, trans‐oceanic long‐distance dispersal. Phylogenetic reconstruction and age estimation for Chroodiscus are consistent with the ‘biotic ferry’ hypothesis: a South American origin and subsequent eastward expansion through Africa towards Southeast Asia and north‐eastern Australia via the Indian subcontinent. The present‐day pantropical distributions of many clades and species of foliicolous lichens might thus be explained by eastward expansion through continental drift, along with the evolution of modern rain forests starting 120 Ma, rather than by the existence of a hypothetical continuous area of pre‐modern rain forest spanning South America, Africa and Southeast Asia during the mid and late Cretaceous.     

The distribution of forest passerine birds and Quaternary climatic change in tropical Africa

A cluster analysis of the distribution patterns of forest passerine birds in tropical Africa showed that they could be divided into three main distribution types: those of restricted distribution (144 species), those of disjunct distributions across West, Central and rarely to East Africa (65 species) and those of very wide distribution (81 species). Centres of species richness, endemism and disjunction coincide spatially, and are identified as forest refugia, where forest persisted throughout Quaternary climatic vicissitudes. These distribution patterns agree with modern interpretations of Quaternary palaeoclimatic changes, which show that glacial periods were arid and interglacials humid. Glacial periods were therefore too dry for montane forest to have spread into areas at present occupied by lowland forest, as advocated by Moreau and other supporters of ihe "pluvial" theory; montane forest is probably as extensive now as at any time during the Quaternary and there is no evidence of past connections between currently isolated montane forests. Bird distribution can be explained largely by slow dispersal outwards from refugia as climatic conditions allowed forest to spread, combined with a relaxing of the distinction between "montane" and "lowland" species under conditions of reduced interspecific competition.     

Identification of glacial refugia in south-eastern North America by phylogeographical analyses of a forest understorey plant, Trillium cuneatum

Aim We examine several hypotheses emerging from biogeographical and fossil records regarding glacial refugia of a southern thermophilic plant species. Specifically, we investigated the glacial history and post‐glacial colonization of a forest understorey species, Trillium cuneatum. We focused on the following questions: (1) Did T. cuneatum survive the Last Glacial Maximum (LGM) in multiple refugia, and (if so) where were they located, and is the modern genetic structure congruent with the fossil record‐based reconstruction of refugia for mesic deciduous forests? (2) What are the post‐glacial colonization patterns in the present geographical range? Location South‐eastern North America. Methods We sampled 45 populations of T. cuneatum throughout its current range. We conducted phylogeographical analyses based on maternally inherited chloroplast DNA (cpDNA haplotypes) and used TCS software to reconstruct intraspecific phylogeny. Results We detected six cpDNA haplotypes, geographically highly structured into non‐overlapping areas. With one exception, none of the populations had mixed haplotype composition. TCS analysis resulted in two intraspecific cpDNA lineages, with one clade subdivided further by shallower diversification. Main conclusions Our investigation revealed that T. cuneatum survived the LGM in multiple refugia, belonging to two (western, eastern) genealogical lineages geographically structured across south‐eastern North America. The western clade is confined to the south‐western corner of T. cuneatum’s modern range along the Lower Mississippi Valley, where fossil records document a major refugium of mesic deciduous forest. For the eastern clade, modern patterns of cpDNA haplotype distribution suggest cryptic vicariance, in the form of forest contractions and subsequent expansions associated with Pleistocene glacial cycles, rather than simple southern survival and subsequent northward colonization. The north–south partitioning of cpDNA haplotypes was unexpected, suggesting that populations of this rather southern thermophilic species may have survived in more northern locations than initially expected based on LGM climate reconstruction, and that the Appalachian Mountains functioned as a barrier to the dispersal of propagules originating in more southern refugia. Furthermore, our results reveal south‐west to north‐east directionality in historical migration through the Valley and Ridge region of north‐west Georgia.     

Ancient islands acted as refugia and pumps for conifer diversity

Island species are thought to be extinction‐prone because of small population sizes, restricted geographical distribution and limited dispersal ability. However, the topographical and environmental heterogeneity, geographical isolation and stability of islands over long timescales could create refugia for taxa whose source area is threatened by environmental changes. We address this possibility by inferring the evolution of the New Caledonia (NC) and New Zealand (NZ) conifer diversity, which represents over 10% of the world's diversity for this group. We estimate speciation and extinction rates in relation to the presence/absence on these islands, and dispersal rates between the islands and surrounding areas. We also test the Eocene submersion of NC and the Oligocene drowning of NZ by comparing the fit of biogeographical scenarios using ancestral area estimations. We find that extinction rates were significantly lower for island species, and dispersal “out of islands” was higher. A model including a diversification shift when NC emerged better explains the diversification dynamics. Biogeographical analyses corroborate that conifers experienced high continental extinctions, but survived on islands. NC and NZ have thus contributed to the world's conifer diversity as “island refugia”, by maintaining early‐diverging lineages from continents during environmental changes on continents. These ancient islands also acted as “species pumps”, providing species into adjacent areas. Our study highlights the important but neglected role of islands in promoting the evolution and conservation of biodiversity.     

Local or global? Biogeography of some primitive Lepidoptera in New Zealand

Abstract

Distribution maps are presented for 30 species of small forest moths representing two genera of Micropterigidae and one genus of Mnesarchaeidae. Evolutionary patterns are interpreted by means of the panbiogeographic method in order to seek a relationship between dispersal patterns within New Zealand and the overseas links of the taxa in question. It is established that different evolutionary lines indeed exhibit different geographic dispersal patterns within New Zealand. Moreover, certain characteristics of these patterns can be attributed to the overseas affinities of the groups, i.e., groups of organisms with New Caledonian or Australian affinities have a western disjunct pattern, whereas those with circum-Pacific affinities have a different pattern in which the taxa occupying a basal position in the phylogeny are to be found in the eastern South Island. However, both groups have overlapping centres of diversity, particularly in NW Nelson.     

Broad‐scale geographic patterns in body size and hind wing development of western Palaearctic carabid beetles(Coleoptera: Carabidae)

Research into large‐scale ecological rules has a long tradition but has received increasing attention over the last two decades. Whereas environmental, especially climatic, influences on the geographic distribution of species traits such as body size are well understood in mammals and birds, our knowledge of the determinants and mechanisms which shape spatial patterns in invertebrate traits is still limited. This study analyzes macroecological patterns in two traits of the highly diverse invertebrate taxon of carabid beetles: body size and hind wing development, the latter being directly linked to species’ dispersal abilities. We tested for potential impacts of environmental variables (spatial, areal, topographic and climate‐related) representing both contemporary conditions and historical processes on large‐scale patterns in the two traits. Regression models revealed hump‐shaped relationships with latitude for both traits in the categories 1) all species, 2) widespread and 3) endemic (restricted‐range) species: body size and the proportion of flightless species increased from northern towards southern Europe and then decreased towards North Africa. The shared and independent influence of environmental factors was analyzed by variation partitioning. While contemporary environmental productivity and stability (represented by measures of ambient energy and water energy balance) had strong positive relationships with carabid body size, patterns in hind wing development were most notably influenced by topography (elevation range). Regions with high elevation range and low historical climate variability (since the last ice age), which likely offer long‐term stable habitats (i.e. glacial refugia), coincide with regions with high proportions of flightless species. Thus geographic patterns in carabid traits tend to be formed not only by recent climate but also by dispersal and historical climate and processes (i.e. glaciations and postglacial colonization).     

The phylogenetic pattern of speciation and wing pattern change in neotropical Ithomia butterflies (Lepidoptera: nymphalidae)

Species level phylogenetic hypotheses can be used to explore patterns of divergence and speciation. In the tropics, speciation is commonly attributed to either vicariance, perhaps within climate-induced forest refugia, or ecological speciation caused by niche adaptation. Mimetic butterflies have been used to identify forest refugia as well as in studies of ecological speciation, so they are ideal for discriminating between these two models. The genus Ithomia contains 24 species of warningly colored mimetic butterflies found in South and Central America, and here we use a phylogenetic hypothesis based on seven genes for 23 species to investigate speciation in this group. The history of wing color pattern evolution in the genus was reconstructed using both parsimony and likelihood. The ancestral pattern for the group was almost certainly a transparent butterfly, and there is strong evidence for convergent evolution due to mimicry. A punctuationist model of pattern evolution was a significantly better fit to the data than a gradualist model, demonstrating that pattern changes above the species level were associated with cladogenesis and supporting a model of ecological speciation driven by mimicry adaptation. However, there was only one case of sister species unambiguously differing in pattern, suggesting that some recent speciation events have occurred without pattern shifts. The pattern of geographic overlap between clades over time shows that closely related species are mostly sympatric or, in one case, parapatric. This is consistent with modes of speciation with ongoing gene flow, although rapid range changes following allopatric speciation could give a similar pattern. Patterns of lineage accumulation through time differed significantly from that expected at random, and show that most of the extant species were present by the beginning of the Pleistocene at the latest. Hence Pleistocene refugia are unlikely to have played a major role in Ithomia diversification.     

Comparative phylogeography reveals a shared impact of pleistocene environmental change in shaping genetic diversity within nine Anopheles mosquito species across the Indo-Burma biodiversity hotspot

South-East Asia is one of the world's richest regions in terms of biodiversity. An understanding of the distribution of diversity and the factors shaping it is lacking, yet essential for identifying conservation priorities for the region's highly threatened biodiversity. Here, we take a large-scale comparative approach, combining data from nine forest-associated Anopheles mosquito species and using statistical phylogeographical methods to disentangle the effects of environmental history, species-specific ecology and random coalescent effects. Spatially explicit modelling of Pleistocene demographic history supports a common influence of environmental events in shaping the genetic diversity of all species examined, despite differences in species' mtDNA gene trees. Populations were periodically restricted to allopatric northeastern and northwestern refugia, most likely due to Pleistocene forest fragmentation. Subsequent southwards post-glacial recolonization is supported by a north-south gradient of decreasing genetic diversity. Repeated allopatric fragmentation and recolonization have led to the formation of deeply divergent geographical lineages within four species and a suture zone where these intraspecific lineages meet along the Thai-Myanmar border. A common environmental influence for this divergence was further indicated by strong support for simultaneous divergence within the same four species, dating to approximately 900 thousand years ago (kya). Differences in the geographical structuring of genetic diversity between species are probably the result of varying species' biology. The findings have important implications for conservation planning; if the refugial regions and suture zone identified here are shared by other forest taxa, the unique and high levels of genetic diversity they house will make these areas conservation priorities.     

Avian host composition,local speciation and dispersal drive the regional assembly of avian malaria parasites in South American birds

Identifying the ecological factors that shape parasite distributions remains a central goal in disease ecology. These factors include dispersal capability, environmental filters and geographic distance. Using 520 haemosporidian parasite genetic lineages recovered from 7,534 birds sampled across tropical and temperate South America, we tested (a) the latitudinal diversity gradient hypothesis and (b) the distance–decay relationship (decreasing proportion of shared species between communities with increasing geographic distance) for this host–parasite system. We then inferred the biogeographic processes influencing the diversity and distributions of this cosmopolitan group of parasites across South America. We found support for a latitudinal gradient in diversity for avian haemosporidian parasites, potentially mediated through higher avian host diversity towards the equator. Parasite similarity was correlated with climate similarity, geographic distance and host composition. Local diversification in Amazonian lineages followed by dispersal was the most frequent biogeographic events reconstructed for haemosporidian parasites. Combining macroecological patterns and biogeographic processes, our study reveals that haemosporidian parasites are capable of circumventing geographic barriers and dispersing across biomes, although constrained by environmental filtering. The contemporary diversity and distributions of haemosporidian parasites are mainly driven by historical (speciation) and ecological (dispersal) processes, whereas the parasite community assembly is largely governed by host composition and to a lesser extent by environmental conditions.