Tracing the origin of disjunct distributions: a case of biogeographical convergence in Pyrgus butterflies

Aim To study the biogeographical factors responsible for the current disjunct distributions of two closely related species of butterflies (Pyrgus cinarae and Pyrgus sidae, Lepidoptera: Hesperioidea). Both species have small populations in the Iberian Peninsula that are isolated by more than 1000 km from their nearest conspecifics. Because these species possess similar ecological preferences and geographical distributions, they are excellent candidates for congruent biogeographical histories. Location The Palaearctic region, with a special focus on the Mediterranean peninsulas as glacial refugia. Methods We integrated phylogeography and population genetic analyses with ecological niche modelling. The mitochondrial gene cytochrome c oxidase subunit 1 (COI) and the non‐coding nuclear marker internal transcribed spacer 2 (ITS2) were analysed for 62 specimens of P. cinarae and for 80 of P. sidae to infer phylogeography and to date the origin of disjunct distributions. Current and ancestral [Last Glacial Maximum using MIROC (Model for Interdisciplinary Research on Climate) and CCSM (Community Climate System Model) circulation models] distribution models were calculated with Maxent . Using present climatic conditions, we delimited the ecological space for each species. Results The genetic structure and potential ancestral distribution of the two species were markedly different. While the Iberian population of P. cinarae had an old origin (c. 1 Ma), that of P. sidae was closely related to French and Italian lineages (which jointly diverged from eastern populations c. 0.27 Ma). Ecological niche modelling showed that minor differences in the ecological preferences of the two species seem to account for their drastically different distributional response to the last glacial to post‐glacial environmental conditions. Although the potential distribution of P. cinarae was largely unaffected by climate change, suitable habitat for P. sidae strongly shifted in both elevation and latitude. This result might explain the early origin of the disjunct distribution of P. cinarae, in contrast to the more recent disjunction of P. sidae. Main conclusions We show that convergent biogeographical patterns can be analysed with a combination of genetic and ecological niche modelling data. The results demonstrate that species with similar distributional patterns and ecology may still have different biogeographical histories, highlighting the importance of including the temporal dimension when studying biogeographical patterns.     

A revision of the Neotropical species of Bolitogyrus Chevrolat,a geographically disjunct lineage of Staphylinini (Coleoptera,Staphylinidae)

The Neotropical species of the rarely collected genus Bolitogyrus (Coleoptera: Staphylinidae: Staphylininae: Staphylinini) are revised. The genus exhibits an uncommon, disjunct distribution between the Neotropical and Oriental Regions and is of unknown phylogenetic position within Staphylinini. Morphological evolution remarkable for Staphylinini was discovered within Bolitogyrus, including sexually dimorphic modifications of the pronotum that may be involved in male competition for females. rSEM interactive animations were used to establish morphological species boundaries between two highly variable species and are provided to illustrate diagnostic characters of the genitalia in unconventional views. The genus is redescribed based on the world fauna and twenty-eight Neotropical species are considered valid. Of these, nineteen are described as new to science: Bolitogyrus ashei sp. n.; B. apicofasciatus sp. n.; B. brevistellus sp. n.; B. bufo sp. n.; B. cheungi sp. n.; B. cornutus sp. n.; B. divisus sp. n.; B. falini sp. n.; B. gracilis sp. n.; B. inexspectatus sp. n.; B. longistellus sp. n.; B. marquezi sp. n.; B. newtoni sp. n.; B. pseudotortifolius sp. n.; B. pulchrus sp. n.; B. silex sp. n.; B. thomasi sp. n.; B. tortifolius sp. n.; and B. viridescens sp. n. Bolitogyrus sallei (Kraatz), stat. r. is removed from synonymy with B. buphthalmus (Erichson) and the following new synonyms are proposed: Cyrtothorax cyanescens Sharp, 1884, syn. n. = Quedius buphthalmus Erichson, 1840; C. nevermanni Scheerpeltz, 1974, syn. n. = C. costaricensis Wendeler, 1927. A summary of all available bionomic and distributional data, as well as an illustrated identification key to and diagnoses of all Neotropical species are provided.     

Molecular phylogeny of the squeak beetles, a family with disjunct Palearctic-Australian range

Many higher groups of plants and animals show distributional patterns which have been shown or have at some point in time been suggested to be correlated with plate tectonics and the ancient supercontinents Laurasia and Gondwana. Here, we study the family of squeak beetles (Coleoptera: Adephaga: Hygrobiidae) and its enigmatic distribution pattern, with one species in the Western Palearctic, one in China and four in Australia. We present a molecular phylogeny including five of the six extant species, showing the monophyly of the Australian radiation. We use a molecular clock approach, which indicates that Hygrobiidae is an ancient group dating back to the breakup of Pangea and discuss the possibility of vicariance as explanation for its current distribution.     

The complex history of the olive tree: from Late Quaternary diversification of Mediterranean lineages to primary domestication in the northern Levant

The location and timing of domestication of the olive tree, a key crop in Early Mediterranean societies, remain hotly debated. Here, we unravel the history of wild olives (oleasters), and then infer the primary origins of the domesticated olive. Phylogeography and Bayesian molecular dating analyses based on plastid genome profiling of 1263 oleasters and 534 cultivated genotypes reveal three main lineages of pre-Quaternary origin. Regional hotspots of plastid diversity, species distribution modelling and macrofossils support the existence of three long-term refugia; namely the Near East (including Cyprus), the Aegean area and the Strait of Gibraltar. These ancestral wild gene pools have provided the essential foundations for cultivated olive breeding. Comparison of the geographical pattern of plastid diversity between wild and cultivated olives indicates the cradle of first domestication in the northern Levant followed by dispersals across the Mediterranean basin in parallel with the expansion of civilizations and human exchanges in this part of the world.     

First fossil Molinaranea Mello‐Leitão, 1940 (Araneae: Araneidae), from middle Miocene Dominican amber,with a phylogenetic and palaeobiogeographical analysis of the genus

The first fossil Molinaranea is described, from middle Miocene Dominican amber. This record extends the known range of the genus back 16 million years; it also extends the geographical range of the genus through time, with extant species known only from Chile, Argentina, the Falkland Islands, and Juan Fernandez Island. A parsimony‐based phylogenetic analysis was performed, which indicates that the fossil species, Molinaranea mitnickii sp. nov. , is nested with Molinaranea magellanica Walckenaer, 1847 and Molinaranea clymene Nicolet, 1849 . A modified Brooks parsimony analysis was conducted in order to examine the biogeography and origins of the fossil species in the Dominican Republic; the analysis suggests that M. mitnickii sp. nov. arrived in Hispaniola from South America as a result of a chance dispersal event. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 711–725.     

Diversification of Lupinus (Leguminosae) in the western New World: derived evolution of perennial life history and colonization of montane habitats

Previous phylogenetic studies of Lupinus (Leguminosae) based on nuclear DNA have shown that the western New World taxa form a monophyletic group representing the majority of species in the genus, with evidence for high rates of recent diversification in South America following final uplift of the Andes 2–4 million years ago (Mya). For this study, three regions of rapidly evolving non-coding chloroplast DNA (trnL intron, trnS–trnG, and trnT–trnL) were examined to estimate the timing and rates of diversification in the western New World, and to infer ancestral states for geographic range, life history, and maximum elevation. The western New World species (5.0–9.3 Mya, 0.6–1.1 spp./My) comprise a basally branching assemblage of annual plants endemic to the lower elevations of western North America, from which two species-rich clades are recently derived: (i) the western North American perennials from the Rocky Mountains, Great Basin, and Pacific Slope (0.7–2.1 Mya, 2.0–5.9 spp./My) and (ii) the predominantly perennial species from the Andes Mountains of South America and highlands of Mexico (0.8–3.4 Mya, 1.4–5.7 spp./My). Bayesian posterior predictive tests for association between life history and maximum elevation demonstrate that perennials are positively correlated with higher elevations. These results are consistent with a series of one or more recent radiations in the western New World, and indicate that rapid diversification of Lupinus coincides with the derived evolution of perennial life history, colonization of montane habitats, and range expansion from North America to South America.     

A coupled phylogeographical and species distribution modelling approach recovers the demographical history of a Neotropical seasonally dry forest tree species

We investigated here the demographical history of Tabebuia impetiginosa (Bignoniaceae) to understand the dynamics of the disjunct geographical distribution of South American seasonally dry forests (SDFs), based on coupling an ensemble approach encompassing hindcasting species distribution modelling and statistical phylogeographical analysis. We sampled 17 populations (280 individuals) in central Brazil and analysed the polymorphisms at chloroplast (trnS‐trnG, psbA‐trnH, and ycf6‐trnC intergenic spacers) and nuclear (ITS nrDNA) genomes. Phylogenetic analyses based on median‐joining network showed no haplotype sharing among population but strong evidence of incomplete lineage sorting. Coalescent analyses showed historical constant populations size, negligible gene flow among populations, and an ancient time to most recent common ancestor dated from ~4.7 ± 1.1 Myr BP. Most divergences dated from the Lower Pleistocene, and no signal of important population size reduction was found in coalescent tree and tests of demographical expansion. Demographical scenarios were built based on past geographical range dynamic models, using two a priori biogeographical hypotheses (‘Pleistocene Arc’ and ‘Amazonian SDF expansion’) and on two additional hypotheses suggested by the palaeodistribution modelling built with several algorithms for distribution modelling and palaeoclimatic data. The simulation of these demographical scenarios showed that the pattern of diversity found so far for T. impetiginosa is in consonance with a palaeodistribution expansion during the last glacial maximum (LGM, 21 kyr BP), strongly suggesting that the current disjunct distribution of T. impetiginosa in SDFs may represent a climatic relict of a once more wide distribution.     

Modelling the distribution of a threatened habitat: the California sage scrub

Aim Using predictive species distribution and ecological niche modelling our objectives are: (1) to identify important climatic drivers of distribution at regional scales of a locally complex and dynamic system – California sage scrub; (2) to map suitable sage scrub habitat in California; and (3) to distinguish between bioclimatic niches of floristic groups within sage scrub to assess the conservation significance of analysing such species groups. Location Coastal mediterranean‐type shrublands of southern and central California. Methods Using point localities from georeferenced herbarium records, we modelled the potential distribution and bioclimatic envelopes of 14 characteristic sage scrub species and three floristic groups (south‐coastal, coastal–interior disjunct and broadly distributed species) based upon current climate conditions. Maxent was used to map climatically suitable habitat, while principal components analysis followed by canonical discriminant analysis were used to distinguish between floristic groups and visualize species and group distributions in multivariate ecological space. Results Geographical distribution patterns of individual species were mirrored in the habitat suitability maps of floristic groups, notably the disjunct distribution of the coastal–interior species. Overlap in the distributions of floristic groups was evident in both geographical and multivariate niche space; however, discriminant analysis confirmed the separability of floristic groups based on bioclimatic variables. Higher performance of floristic group models compared with sage scrub as a whole suggests that groups have differing climate requirements for habitat suitability at regional scales and that breaking sage scrub into floristic groups improves the discrimination between climatically suitable and unsuitable habitat. Main conclusions The finding that presence‐only data and climatic variables can produce useful information on habitat suitability of California sage scrub species and floristic groups at a regional scale has important implications for ongoing efforts of habitat restoration for sage scrub. In addition, modelling at a group level provides important information about the differences in climatic niches within California sage scrub. Finally, the high performance of our floristic group models highlights the potential a community‐level modelling approach holds for investigating plant distribution patterns.     

The application of environmental ethics in biological conservation: a case study from the southernmost tip of the Americas

Biological conservation is not only about facts and technical measures concerning ecology, rather it must also consider values. This pertains to both the balancing of various human interests and also to the ethical evaluation of human actions towards nature. Here we discuss how environmental ethics can be incorporated into conservation decisions, and what implications the inclusion of ethical valuation has for the practice of conservation biology. While this is done mostly on a rather abstract level, we illustrate this here by applying ethical theory to a case study: the options for management of the introduced North American beaver (Castor canadensis) in the very south of Chile (Navarino Island). The beaver is an exotic species to the area and has substantially altered the ecological systems of the region. We discuss different options for dealing with the beaver (eradicate, control, tolerate, promote) from the viewpoint of anthropocentric environmental ethics and biocentric ethics. The results of our analysis demonstrate the value of ethical discussions in clarifying and underpinning arguments for and against specific actions. At the same time, they also show that ethical arguments do not decrease the need for sound scientific data but, on the contrary, may even increase this demand. We also highlight that the conclusions regarding adequate actions to be taken vary depending on the specific ethical theory embraced.

Colonization history of Mallorca Island by the European rabbit,Oryctolagus cuniculus,and the Iberian hare,Lepus granatensis (Lagomorpha: Leporidae)

The Mediterranean islands have a long history of human‐mediated introductions resulting in frequent replacements of their fauna and flora. Although these histories are sometimes well documented or may be inferred from paleontological studies, the use of phylogenetic and population genetic reconstruction methods provides a complementary perspective for answering questions related to the history of insular species. In the present study, we infer the colonization history of Mallorca (Balearic Islands) by the European rabbit (Oryctolagus cuniculus) and the Iberian hare (Lepus granatensis) using sequence variation of the mitochondrial DNA control region from continental and insular specimens (total of 489 sequences). Additionally, the taxonomic identity of Mallorcan L. granatensis was confirmed using a diagnostic nuclear marker. For both Mallorcan rabbits and hares, genetic diversity was comparable to the continental populations, suggesting the introduction of multiple lineages. Two Mallorcan haplogroups were found in hares, which likely correspond to two introduction events. Rabbits from Mallorca were identified as belonging to the subspecies Oryctolagus cuniculus cuniculus, and may have been originated both from Iberian and French populations. The molecular estimates of the timing of the colonization events of the Mallorcan lagomorphs are consistent with human‐mediated introductions by early settlers on the islands. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 748–760.     

Sea warming and fish distribution: the case of the small pelagic fish, Sardinella aurita, in the western Mediterranean

This study analyses the temporal and spatial changes in abundance and distribution of the warm water species round sardinella (Sardinella aurita) in the western Mediterranean over the last decades in relation to sea water temperature. In the western Mediterranean basin (1950–2003), a significant positive relationship was found between round sardinella landings and temperature anomalies. Along a latitudinal gradient off the Mediterranean Iberian coast (1989–2004), a gradual increase in species abundance was observed from south to north, with a certain time lag going northwards, associated with the increase in sea water temperature. The abundance of round sardinella in the two warmest and southernmost areas was positively and significantly correlated with sea surface temperature registered during the start of gonad maturation the previous year. In addition, the positive relationship established between water temperature and abundance of round sardinella in the coldest and northernmost study area demonstrates that there is a temperature limit for the distribution of this species in the western Mediterranean. In addition, this study analyses round sardinella larvae distribution and abundance in the summers of 2003 and 2004, and conducts a comparison with the situation 20 years ago (summer 1983). Results show a marked increase in larval abundance during the last decades and the present appearance of larvae in the northernmost study areas, where they did not occur 20 years ago. This indicates the successful reproduction of round sardinella in the northern part of the Mediterranean, where the species has expanded, confirming its establishment in the area.     

Phylogeography,historical demography and habitat suitability modelling of freshwater fishes inhabiting seasonally fluctuating Mediterranean river systems: a case study using the Iberian cyprinid Squalius valentinus

The Mediterranean freshwater fish fauna has evolved under constraints imposed by the seasonal weather/hydrological patterns that define the Mediterranean climate. These conditions have influenced the genetic and demographic structure of aquatic communities since their origins in the Mid‐Pliocene. Freshwater species in Mediterranean‐type climates will likely constitute genetically well‐differentiated populations, to varying extents depending on basin size, as a consequence of fragmentation resulting from drought/flood cycles. We developed an integrative framework to study the spatial patterns in genetic diversity, demographic trends, habitat suitability modelling and landscape genetics, to evaluate the evolutionary response of Mediterranean‐type freshwater fish to seasonal fluctuations in weather. To test this evolutionary response, the model species used was Squalius valentinus, an endemic cyprinid of the Spanish Levantine area, where seasonal weather fluctuations are extreme, although our findings may be extrapolated to other Mediterranean‐type species. Our results underscore the significant role of the Mediterranean climate, along with Pleistocene glaciations, in diversification of S. valentinus. We found higher nuclear diversity in larger drainage basins, but higher mitochondrial diversity correlated to habitat suitability rather than basin size. We also found strong correlation between genetic structure and climatic factors associated with Mediterranean seasonality. Demographic and migration analyses suggested population expansion during glacial periods that also contributed to the current genetic structure of S. valentinus populations. The inferred models support the significant contribution of precipitation and temperature to S. valentinus habitat suitability and allow recognizing areas of habitat stability. We highlight the importance of stable habitat conditions, fostered by typical karstic springs found on the Mediterranean littoral coasts, for the preservation of freshwater species inhabiting seasonally fluctuating river systems.