Modelling the distribution of the invasive Roesel’s bushcricket (Metrioptera roeselii) in a fragmented landscape

The development of conservation strategies to mitigate the impact of invasive species requires knowledge of the species ecology and distribution. This is, however, often lacking as collecting biological data may be both time-consuming and resource intensive. Species distribution models can offer a solution to this dilemma by analysing the species-environment relationship with help of Geographic information systems (GIS). In this study, we model the distribution of the non-native bush-cricket Metrioptera roeselii in the agricultural landscape in mid-Sweden where the species has been rapidly expanding in its range since the 1990s. We extract ecologically relevant landscape variables from Swedish CORINE land-cover maps and use species presence-absence data from large-scale surveys to construct a species distribution model (SDM). The aim of the study is to increase the knowledge of the species range expansion pattern by examining how its distribution is affected by landscape composition and structure, and to evaluate SDM performance at two different spatial scales. We found that models including data on a scale of 1 × 1 km were able to explain more of the variation in species distribution than those on the local scale (10 m buffer on each side of surveyed road). The amount of grassland in the landscape, estimated from the area of arable land, pasture and rural settlements, was a good predictor of the presence of the species on both scales. The measurements of landscape structure – linear elements and fragmentation - gave ambivalent results which differed from previous small scaled studies on species dispersal behaviour and occupancy patterns. The models had good predictive ability and showed that areas dominated by agricultural fields and their associated grassland edges have a high probability being colonised by the species. Our study identified important landscape variables that explain the distribution of M. roeselii in Mid-Sweden that may also be important to other range expanding orthopteran species. This work will serve as a foundation for future analyses of species spread and ecological processes during range expansion

A clue to invasion success: genetic diversity quickly rebounds after introduction bottlenecks

One of the fundamental questions in invasion biology is to understand the genetic mechanisms behind success or failure during the establishment of a species. However, major limitations to understanding are usually a lack of spatiotemporal population data and information on the populations' colonisation history. In a large-scale, detailed study on the bush-cricket Metrioptera roeselii 70 groups of founders were introduced in areas outside the species' distribution range. We examined how (1) the number of founders (2-32 individuals), (2) the time since establishment (7 or 15 years after introduction) and (3) possible gene flow affected establishment success and temporal genetic changes of the introduced populations. We found higher establishment success in introductions with larger propagule sizes but genetic diversity indices were only partly correlated to propagule size. As expected, introduced populations were more similar to their founder population the larger the propagule size was. However, even if apparent at first, most of the differentiation in the small propagule introductions disappeared over time. Surprisingly, genetic variability was regained to a level comparable to the large and outbreeding founder population only 15 generations after severe demographic bottlenecks. We suggest that the establishment of these populations could be a result of several mechanisms acting in synergy. Here, rapid increase in genetic diversity of few introductions could potentially be attributed to limited gene flow from adjacent populations, behavioural adaptations and/or even increased mutation rate. We present unique insights into genetic processes that point towards traits that are important for understanding species' invasiveness

Are peripheral populations special? Congruent patterns in two butterfly species

Populations at range margins may be genetically different from more central ones for a number of mutually non-exclusive reasons. Specific selection pressures may operate in environments that are more marginal for the species. Genetic drift may also have a strong effect in these populations if they are small, isolated and/or have experienced significant bottlenecks during the colonisation phase. The question if peripheral populations are special, and if yes then how and why, is of obvious relevance for speciation theory, as well as for conservation biology. To evaluate the uniqueness of populations at range margins and the influence of gene flow and selection, we performed a morphometric study of two grassland butterfly species: Coenonympha arcania and C. hero (Lepidoptera: Nymphalidae). The samples were collected from Swedish populations that are peripheral and isolated from the main area of the species distributions and from populations in the Baltic states that are peripheral but connected to the main area of the species distributions. These samples were compared to those from central parts of the species distributions. The isolated populations in both species differed consistently from both peripheral and central populations in their wing size and shape. We interpret this as a result of selection caused by differences in population structure in these isolated locations, presumably favoring different dispersal propensity of these butterflies. Alternative explanations based on colonisation history, latitudinal effects, inbreeding or phenotypic plasticity appear less plausible. As a contrast, the much weaker and seemingly random among-region differences in wing patterns are more likely to be ascribed to weaker selection pressures allowing genetic drift to be influential. In conclusion, both morphological data and results from neutral genetic markers in earlier studies of the same system provide congruent evidence of both adaptation and genetic drift in the isolated Swedish populations of both species

Data from: Evaluating range-expansion models for calculating nonnative species’ expansion rate

1. Species range shifts associated with environmental change or biological invasions are increasingly important study areas. However, quantifying range expansion rates may be heavily influenced by methodology and/or sampling bias. 2. We compared expansion rate estimates of Roesel’s bush-cricket (Metrioptera roeselii, Hagenbach 1822), a non-native species currently expanding its range in south-central Sweden, from range statistic models based on distance measures (mean, median, 95th gamma quantile, marginal mean, maximum and conditional maximum) and an area-based method (grid occupancy). We used sampling simulations to determine the sensitivity of the different methods to incomplete sampling across the species’ range. 3. For periods when we had comprehensive survey data, range expansion estimates clustered into two groups: (i) those calculated from range margin statistics (gamma, marginal mean, maximum and conditional maximum: ~3 km/yr), and (ii) those calculated from the central tendency (mean and median) and the area-based method of grid occupancy (~1.5 km/yr). 4. Range statistic measures differed greatly in their sensitivity to sampling effort; the proportion of sampling required to achieve an estimate within 10% of the true value ranged from 0.17-0.9. Grid occupancy and median were most sensitive to sampling effort, and the maximum and gamma quantile the least. 5. If periods with incomplete sampling were included in the range expansion calculations, this generally lowered the estimates (range 16-72%), with exception of the gamma quantile that was slightly higher (6%). 6. Care should be taken when interpreting rate expansion estimates from data sampled from only a fraction of the full distribution. Methods based on the central tendency will give rates approximately half that of methods based on the range margin. The gamma quantile method appears to be the most robust to incomplete sampling bias and should be considered as the method of choice when sampling the entire distribution is not possible

Gene flow relates to evolutionary divergence among populations at the range margin

Background: Morphological differentiation between populations resulting from local adaptations to environmental conditions is likely to be more pronounced in populations with increasing genetic isolation. In a previous study a positive clinal variation in body size was observed in isolated Roesel's bush-cricket, Metrioptera roeselii, populations, but were absent from populations within a continuous distribution at the same latitudinal range. This observational study inferred that there was a phenotypic effect of gene flow on climate-induced selection in this species.Methods: To disentangle genetic versus environmental drivers of population differences in morphology, we measured the size of four different body traits in wild-caught individuals from the two most distinct latitudinally-matched pairs of populations occurring at about 60 degrees N latitude in northern Europe, characterised by either restricted or continuous gene flow, and corresponding individuals raised under laboratory conditions.Results: Individuals that originated from the genetically isolated populations were always bigger (femur, pronotum and genital appendages) when compared to individuals from latitudinally-matched areas characterised by continuous gene flow between populations. The magnitude of this effect was similar for wild-caught and laboratory-reared individuals. We found that previously observed size cline variation in both male and female crickets was likely to be the result of local genetic adaptation rather than phenotypic plasticity.Conclusions: This strongly suggests that restricted gene flow is of major importance for frequencies of alleles that participate in climate-induced selection acting to favour larger phenotypes in isolated populations towards colder latitudes

Data from: Limited emigration from an outbreak of a forest pest insect

Population density and individual dispersal behaviour affect species´ distribution dynamics. Population densities vary over time, and some species occasionally increase to very high numbers, for example during outbreaks. In such situations populations are expected to expand into new areas due to density-dependent dispersal and sometimes even result in range expansion. A local population of the northern pine processionary moth Thaumetopoea pinivora has recently reached outbreak densities at the edge of its northern range at the southern tip of Gotland Island in the Baltic Sea. We first investigated if the outbreak had resulted in establishment of populations in suitable habitats on Gotland Island outside of the outbreak area. Six small populations were found that could potentially have originated from the outbreak area. However, data from 12 microsatellite markers strongly suggest that these populations did not originate from the recent outbreak. Genetic variability was not reduced in these small, isolated populations and there were several unique alleles, indicating instead a different population history and that there has been no recent range expansion. In addition, there was apparent genetic isolation by geographic distance, implying that despite that the outbreak population reached a high enough density to cause density-dependent dispersal, significant gene flow has not occurred

Roeselii_invasion_data

The .xls file contains the years and relative positions that Roesel's bush-crickets were observed in southern-central Sweden from 1981-2010 during their ongoing invasion of this area. The distance measures are in metres relative to the original observation location (see Preuss et al. 2014, Ecology and Evolution for more details). The observation locations were summarised as the centre of 5x5km landscape squares

Gene flow relates to evolutionary divergence among populations at the range margin

Background: Morphological differentiation between populations resulting from local adaptations to environmental conditions is likely to be more pronounced in populations with increasing genetic isolation. In a previous study a positive clinal variation in body size was observed in isolated Roesel's bush-cricket, Metrioptera roeselii, populations, but were absent from populations within a continuous distribution at the same latitudinal range. This observational study inferred that there was a phenotypic effect of gene flow on climate-induced selection in this species. Methods: To disentangle genetic versus environmental drivers of population differences in morphology, we measured the size of four different body traits in wild-caught individuals from the two most distinct latitudinally-matched pairs of populations occurring at about 60 degrees N latitude in northern Europe, characterised by either restricted or continuous gene flow, and corresponding individuals raised under laboratory conditions. Results: Individuals that originated from the genetically isolated populations were always bigger (femur, pronotum and genital appendages) when compared to individuals from latitudinally-matched areas characterised by continuous gene flow between populations. The magnitude of this effect was similar for wild-caught and laboratory-reared individuals. We found that previously observed size cline variation in both male and female crickets was likely to be the result of local genetic adaptation rather than phenotypic plasticity. Conclusions: This strongly suggests that restricted gene flow is of major importance for frequencies of alleles that participate in climate-induced selection acting to favour larger phenotypes in isolated populations towards colder latitudes

Wing morphology of the butterfly Coenonympha arcania in Europe : Traces of both historical isolation in glacial refugia and current adaptation

In this study, we examined the evolutionary outcome of and interplay between historic isolation and current selection pressures on traits more or less closely connected to fitness in the Pearly Heath butterfly (Coenonympha arcania) across its range in Europe. We hypothesized that a trait mean is more related to historic events if it has low connection to fitness, while a trait more closely connected with fitness is expected to have a mean that relates more to current selection pressures. In order to test this, we collected 322 butterflies from across the species range in Europe and measured five wing traits relating to size and color patterns. To infer a phylogeographic history for each individual, we sequenced a 594 bp fragment of the COI gene. The morphological data were then analyzed in relation to selected climatic variables and the history of individuals to disentangle which factors best correlated with morphological variation. The results supported our hypothesis in that wing sizes correlated with summer precipitation but not with its inferred location during the last glaciation. Eyespot position, on the other hand, correlated with the history of individuals but not with the analyzed climatic indicators. The sizes of the black spot and the white band, two traits that were expected to have intermediate selection pressure, were associated with both history and current conditions. Thus, this study illustrates the fascinating interplay between events and processes that lead to a specific evolutionary outcome