Dispersal as a key element of community structure: the case of ground beetles on lake islands

In order to study the influence of dispersal ability on community structure of ground beetles (carabidae), 15 lake islands and 2 mainland sites of the lake Mamry archipelago, northern Poland, were sampled by using pitfall traps. Of the 71 ground beetle species detected, 47 were macropterous, 16 wing-dimorphic and 8 brachypterous. Macropterous species had lower site abundances and occupied fewer sites than dimorphic and brachypterous species. There were trends from macropterous to brachypterous species towards a nested distribution across the sites and towards over-dispersed (aggregated) species co-occurrences. Canonical correspondence analysis pointed to site isolation and area as main factors influencing site abundance and spatial distribution. Our results imply that further studies on species co-occurrences and community assembly have to consider dispersal ability as a key element influencing ecological distributions at the regional scale.     

The impact of hedge-forest connectivity and microhabitat conditions on spider and carabid beetle assemblages in agricultural landscapes

Agricultural intensification in terms of decreasing landscape complexity and connectivity has negatively affected biodiversity. Linear landscape elements composed of woody vegetation like hedges may counteract this negative trend by providing habitats and enhancing habitat connectivity for different organisms. Here, we tested the impacts of habitat type (forest edges vs. hedges) and hedges’ isolation (connected vs. isolated hedges) from forests as well as microhabitat conditions (percentage of bare ground and width) on trait-specific occurrence of ground-dwelling arthropods, namely spiders and carabids. Arthropods were grouped by habitat specialisation (forest vs. open-habitat species vs. generalists), hunting strategy (web-building or hunting spiders) and dispersal ability (wing morphology of carabids). Spider and carabid assemblage composition was strongly influenced by habitat type and isolation, but not by microhabitat conditions. Activity density of forest species and brachypterous carabids was higher in forest edges compared to hedges, whereas open-habitat species and macropterous carabids showed reverse patterns, with no effects of isolation. Occurrence of generalist carabids, but not spiders, was higher in hedges compared to forest edges. Habitat type and isolation did not affect spiders with different hunting strategy. Microhabitat conditions were less important for spider and carabid occurrence. Our study concludes that on a landscape scale, type of linear woody habitat is more important for arthropod occurrence than isolation effects and microhabitat conditions, depending on traits. Hedges provide refuges for species specialised to open habitats and species with high dispersal ability, such as macropterous carabids. Forest edges enhance persistence of species specialised to forests and species with low dispersal ability, such as brachypterous carabids.     

The impact of high-altitude ski pistes on ground-dwelling arthropods in the Alps

The development of winter sport resorts above the timberline may affect every ecosystem component. We analyzed the effect of ski-pistes on the abundance and species richness of arthropods (namely carabids, spiders, opilionids, and grasshoppers) trapped in grasslands adjacent to the ski-run, on ski-pistes and at the edge between these two habitat types. Our results showed that diversity of brachypterous carabids, spiders, and grasshoppers decreased significantly from natural grasslands to ski-pistes. This was not true for the macropterous carabid guild, which included species with contrasting ecological requirements. The analysis of indicator species (IndVal) showed that most of the species (some of them precinctive to restricted areas in the north-western Alps) had clear preferences for natural grassland and few taxa were limited to ski-pistes. Generalized linear models suggested that the local extent of grass and rock cover can significantly affect assemblages: the low grass cover of ski-pistes, in particular, was a serious hindrance to colonization by spider, grasshopper, brachypterous, and some macropterous carabid species. The results obtained, support concerns over the possible disruption of local ecosystem functionality and over the conservation of arthropod species which are endemic to restricted alpine areas. In order to retain arthropod ground-dwelling fauna we suggest that: (i) new, environmentally friendly ways of constructing pistes should be developed to preserve as much soil and grass cover as possible; (ii) existing ski-pistes should be restored through management to promote the recovery of local vegetation.     

The impact of forest ski-pistes on diversity of ground-dwelling arthropods and small mammals in the Alps

Forest clearing for winter sport activities is the major force driving loss and fragmentation of the alpine forests. The establishment of ski-pistes involves impacts on every ecosystem component. To assess the extent of this threat we studied ground-dwelling arthropods (namely ground beetles and spiders) and small mammals (shrews and voles) at two ski resorts in north-western Italian Alps by pitfall trapping. Diversity parameters (mean abundance, species richness and Shannon index) of spiders and macropterous carabids increased from forest interior to open habitats (i.e., ski-piste or pasture), whereas parameters of brachypterous carabids significantly decreased from forest interior to open habitats. Diversity parameters of macropterous ground beetles were higher on pastures than on ski-pistes. Small mammals were virtually absent from ski-pistes. Observed frequencies in the three adjacent habitats were significantly different from expected ones for the bank vole Myodes glareolus and the pygmy shrew Sorex minutus. Generalized linear models showed that abundance, species richness and diversity of spiders and macropterous carabids of ski-pistes were best modelled by combination of factors, including grass cover and width of the ski-piste. Indicator Species Analysis showed that species that significantly preferred ski-pistes were less than those preferring pastures, and species which were exclusive of ski-pistes were very few. To retain arthropod ground-dwelling fauna of open habitats environmentally friendly ways of constructing pistes should be developed. After tree clearing, only the roughest ground surfaces should be levelled, in order to preserve as much natural vegetation as possible. Where necessary, ski-pistes should be restored through the recovery of local vegetation.     

Dispersal and reproductive properties of sugarcane bugs,Cavelerius saccharivorus

The distribution of sugarcane bugs, Cavelerius saccharivorus is remarkably patchy at time of the peak emergence of the first generation adult in early summer. In population near carrying capacity 70% of adults become macropterous, and they move to sparse populations or vacuum areas. The experiment on the reproductive difference between females from field populations with different densities, showed that the overcrowding restricted their oviposition. On the further experiment it was shown that reproductive ability of macropterous adults or dispersers was not much inferior to that of brachypterous from sparse populations. Overcrowding at time of the peak emergence is reduced by the dispersal. Hence the intraspecific competition is a transient condition. The oviposition is at first restricted in residents of dense populations as compared with brachypterous adults of sparse populations, but the difference becomes small as the former density decreases by the emigration. The last section considers the relation of the reproductive rate and dispersal risk to the rate of dispersal with the simple model, which explains the characteristic dispersal of this species.     

Habitat type mediates equilibrium with climatic conditions in the distribution of Iberian diving beetles

Aim The contrasting habitat permanence over geological time‐scales of lotic and lentic habitats may impose different constraints on the dispersal ability of their macroinvertebrate populations, and ultimately on the degree of equilibrium with current climate. We aim to test for differences between species typical of either habitat type in their potential versus realized distributions as a surrogate measure of degree of climate equilibrium, both in refuges and more recently deglaciated areas. Location Western Europe. Methods We focus on 99 Iberian diving beetles (family Dytiscidae). A multidimensional envelope procedure was used to estimate their potential distributions, which were projected for different spatial scales. At the continental scale we calculated the percentage of countries with climatically suitable conditions for each species over those actually occupied (range filling). At the regional scale, we estimated realized distributions using: (1) convex hull polygons for Sweden and the Iberian Peninsula; and (2) generalized linear models for the Iberian Peninsula. Results In the Iberian Peninsula, differences in the degree of equilibrium with climatic conditions between lotic and lentic species were few, if any. However, at the continental scale we found significant differences, with lentic species closer to equilibrium than lotic species. In the recently deglaciated area (Sweden) the subset of species with ranges wide enough to encompass Iberia and Scandinavia were mostly lentic, and all were closer to climatic equilibrium without significant differences between habitat types. Main conclusions Our results show that, at continental scales, climate equilibrium is not concordant between the habitat types across western Europe. We hypothesize that: (1) the differences between refuge areas in dispersal ability are erased probably due to long‐term climatic stability, allowing enough time to reach equilibrium, and (2) the species with wide geographical ranges able to recolonize recently deglaciated areas should have the highest dispersal abilities, and are closer to climatic equilibrium.     

From climate change predictions to actions – conserving vulnerable animal groups in hotspots at a regional scale

Current climate change is a major threat to biodiversity. Species unable to adapt or move will face local or global extinction and this is more likely to happen to species with narrow climatic and habitat requirements and limited dispersal abilities, such as amphibians and reptiles. Biodiversity losses are likely to be greatest in global biodiversity hotspots where climate change is fast, such as the Iberian Peninsula. Here we assess the impact of climate change on 37 endemic and nearly endemic herptiles of the Iberian Peninsula by predicting species distributions for three different times into the future (2020, 2050 and 2080) using an ensemble of bioclimatic models and different combinations of species dispersal ability, emission levels and global circulation models. Our results show that species with Atlantic affinities that occur mainly in the North‐western Iberian Peninsula have severely reduced future distributions. Up to 13 species may lose their entire potential distribution by 2080. Furthermore, our analysis indicates that the most critical period for the majority of these species will be the next decade. While there is considerable variability between the scenarios, we believe that our results provide a robust relative evaluation of climate change impacts among different species. Future evaluation of the vulnerability of individual species to climate change should account for their adaptive capacity to climate change, including factors such as physiological climate tolerance, geographical range size, local abundance, life cycle, behavioural and phenological adaptability, evolutionary potential and dispersal ability.     

Wing morph-related physiological differences in adults of temperate population of Pyrrhocoris apterus (L.) (Heteroptera: Pyrrhocoridae)

The reproductive and diapausing adult females of brachypterous morph and macropterous females with reproductive arrest of non-diapause type, originating from the laboratory cultures of Pyrrhocoris apterus, were studied for their feeding and drinking behaviour, digestive enzyme activities, and carbohydrate and lipid contents. The highest feeding and drinking activities were observed in reproductive brachypters, the lowest in macropters. Macropters also differed from brachypters by lower activities of gut lipase, peptidase and protease, lower concentration of haemolymph sugars, and lower weight of fat body, which probably reflects their low feeding activity. The total content of fat body lipids was also lower in macropters (0.6 mg) than in reproductive and diapausing brachypters (4.6 and 7.5 mg, respectively) on day 14. A very high amount of glycogen was found in the fat body of diapausing brachypters, 363 micrograms on day 14, as opposed to 15 and 80 micrograms in macropterous and reproductive brachypterous females, respectively. The obtained data indicate that the most important difference between macropterous and brachypterous females with different types of reproductive arrest consists of an enhanced mobilization of lipids for dispersal in macropters and accumulation of energetic reserves for hibernation in brachypters.     

Habitat structure and neighbor linear features influence more carabid functional diversity in olive groves than the farming system

The effects of land-use management and environmental features at different scales on carabid beetle diversity and trait structure were assessed across olive groves in northeastern Portugal. We selected organic and integrated olive groves that were distinct in terms of specific management practices, local linear features and landscape configurations. Besides the management intensification levels, differences in carabid diversity and community traits were mainly due to local habitat and ecological linear structures at a finer spatial scale. Carabid community traits related to disturbance, namely traits of body size and species dispersal ability, responded to land-use intensity and particular olive grove features were influencing diversity patterns. Within the olive grove patches, larger and brachypterous species were associated to plots with more dense vegetation cover while macropterous and small-sized species were more associated to open areas. Also, larger carabid species benefitted from higher patch size heterogeneity within the landscape mosaics. Our findings indicate that the effects of farming system is contingent on the specific management practices, local and linear features present in agroecosystems such as olive groves. Particularly, the influence of local features on carabid diversity patterns and community traits linked to dispersal and movement may be crucial in maintaining pest control at a landscape scale.     

Spatial and temporal pattern of colonization of Nabidae (Heteroptera) in alfalfa (Medicago sativa)

Abstract: Six species of Nabidae (Heteroptera) were collected by standardized sweep net sampling in alfalfa fields in Thuringia, Germany, from 1993 to 1995: Nabis pseudoferus , N. ferus , N. brevis , N. major , Nabicula flavomarginata and Aptus mirmicoides . Colonization of a newly cultivated field was studied over a 3-year period. The density of all the studied nabid species was low (less than five individuals per 100 sweeps) and not related to time since colonization started, or to the distance from the margin of the field. Macropterous species were able to colonize the whole field within one season. The density of one macropterous species, N. pseudoferus , varied between the years of study and was mainly affected by the harvest regime. The brachypterous species reached the margin within one season but for density it took three seasons to reach satiated values also in the centre of the field. The abundance of the brachypterous N. brevis was significantly different both between years and sampling sites. This indicates the importance of the surroundings on the succession of this species. Nabis major , a fully winged species, showed a migration pattern intermediate to macropterous and brachypterous nabids. These results suggest that the total abundance of nabid predators cannot be predicted by time or distance from the expansion source (shelter belts). The abundance of brachypterous nabid individuals can be predicted from time since colonization but is best analysed at the species level.     

Wing morph-related physiological differences in adults of temperate population of Pyrrhocoris apterus (L.) (Heteroptera: Pyrrhocoridae)

The reproductive and diapausing adult females of brachypterous morph and macropterous females with reproductive arrest of non-diapause type, originating from the laboratory cultures of Pyrrhocoris apterus, were studied for their feeding and drinking behaviour, digestive enzyme activities, and carbohydrate and lipid contents. The highest feeding and drinking activities were observed in reproductive brachypters, the lowest in macropters. Macropters also differed from brachypters by lower activities of gut lipase, peptidase and protease, lower concentration of haemolymph sugars, and lower weight of fat body, which probably reflects their low feeding activity. The total content of fat body lipids was also lower in macropters (0.6 mg) than in reproductive and diapausing brachypters (4.6 and 7.5 mg, respectively) on day 14. A very high amount of glycogen was found in the fat body of diapausing brachypters, 363 μg on day 14, as opposed to 15 and 80 μg in macropterous and reproductive brachypterous females, respectively. The obtained data indicate that the most important difference between macropterous and brachypterous females with different types of reproductive arrest consists of an enhanced mobilization of lipids for dispersal in macropters and accumulation of energetic reserves for hibernation in brachypters.     

The occurrence and distribution of carabid beetles (Carabidae) on islands in the Baltic Sea: a review

The distribution and occurrence of carabid beetles have been studied extensively in a number of archipelagoes in the Baltic Sea, a ca. 377,000 km², fairly shallow and young sea in northern Europe. This work has revealed some surprising results related to colonisation success and maintenance of populations. Dispersal from the mainland and between islands appears to be relatively easy because inter-island distances are small (tens or hundreds of metres) and the salinity of the Baltic is low, which allows survival during drift. This is reflected in the higher than expected proportion of brachypterous species on small islands. A lack of energetically expensive wings, flight muscles and flight fuel may give brachypterous species an advantage - in terms of resource allocation for survival and fecundity - over macropterous species in the colonisation of an island. Once an island is reached, successful colonisation mainly depends on the presence of suitable habitat. The most abundantly collected carabids on these islands are habitat generalists, but some degree of habitat association exists. Yet, the composition of carabid assemblages differs substantially among islands, possibly a result of chance effects. There is evidence that species on smaller islands occur in a wider variety of vegetation types than species on larger islands. Both habitat diversity and area per se are implicated in the positive species-area relationships found, but the small island effect has been identified as being significant in most of the archipelagoes investigated. Pterostichus melanarius, a eurytopic, wing-polymorphic species abundantly collected on the mainland is, unexpectedly, absent from small islands. Possible reasons for this include; (1) interspecific predation and competition for food by the closely related but slightly larger P. niger, which is dominant on small islands, (2) the apparent inability of P. melanarius to reproduce on small islands, (3) its reduced dispersal power as compared to P. niger and (4) the possible absence of some essential resources from small islands. In conclusion, the conservation value of highly threatened cultural landscapes on Baltic Sea islands, such as species rich wooded meadows, is discussed as well as the possible negative effects of climate change on carabid beetles.     

THE EVOLUTION OF WING DIMORPHISM IN INSECTS

Wing-dimorphic insects are excellent subjects for a study of the evolution of dispersal since the nondispersing brachypterous morph is easily recognized. The purpose of this paper is to develop a framework within which the evolution of wing dimorphism can be understood. A review of the literature indicates that the presence or absence of wings may be controlled by a single locus, two-allele genetic system or a polygenic system. Both types of inheritance can be subsumed within a general threshold model. An increase in the frequency of a brachypterous morph in a population may result from an increased relative fitness of this morph or the emigration of the macropterous type. The abundance of wing-polymorphic species argues for an increased fitness of the brachypterous form. An analysis of the life-history characteristics of 22 species of insects indicates that the brachypterous morph is both more fecund and reproduces earlier that the macropterous morph. Unfortunately, data on males are generally lacking. It is suggested that suppression of wing production results when some hormone, perhaps juvenile hormone, exceeds a threshold value during a critical stage of development. Further, it is known that in the monomorphically winged species Oncopeltus fasciatus both flight and oviposition are regulated by the titer of juvenile hormone. These observations are used to construct a possible pathway for the evolution of wing dimorphism. This suggests that evolution to a dimorphic species requires both an increase in the rate of production of the wing suppressing hormone and a change in the threshold level at which wing and wing-muscle production are suppressed. The stage in this evolutionary sequence that an organism will reach depends on the stability of the habitat.     

基于形态及分子特征的鳖扁蜡蝉长翅型研究

研究了鳖扁蜡蝉属Cixiopsis Matsumura长翅型雌虫的形态和分子特征.通过和该属的模式种,即鳖扁蜡蝉Cixiopsis punctatus Matsumura,1900短翅型的形态比较,发现该长翅型雌虫在形态学方面存在某些差异.为了明确该长翅型雌虫的归属,还测定了该长翅型雌虫和鳖扁蜡蝉短翅型的COI序列,结...     

Adaptive significance of wing dimorphism in males ofNilaparvata lugens

The trade-offs associated with macroptery in males were investigated in a wing-dimorphic planthopper, Nilaparvata lugens (Stål) (Delphacidae, Hemiptera). Developmental time of brachypters was significantly shorter than that of macropters across a wide range of population densities, while no within-population differences between wing morphs in body weight were found. In mating experiments, brachypterous males 12 to 24 h old were almost three times more successful in competition for females than macropterous males of the same age, while in five days old males, macropters were twice as successful as brachypters. In competition for females involving direct body contact between males, brachypterous and macropterous males were equally successful. It is suggested that macroptery and delayed peak in sexual activity in males may both be influenced by the titre of juvenile hormone, as is known from females. The combination of prolonged development and delayed maximum competitive ability for mates, associated with macroptery, is likely to be disad vantageous, representing probably a penalty associated with macroptery in males in undisturbed habitats.     

Biogeographical tests of the vicariance model in Mediterranean land snails

Aim To investigate the importance of vicariance in shaping land snail faunas. Location Three data sets of Mediterranean land snails were analysed: Helicoidea of the Iberian Peninsula and the complete land snail faunas of the central and eastern Aegean Islands and of Israel and Palestine. Methods The vicariance model predicts a clustering of species ranges. We tested for clustering of species ranges with a Monte Carlo simulation. For this simulation we used a null model that generates range data sets in such a way that their range size distribution, their species richness distribution and the spatial autocorrelation of the ranges approximate the parameters in the real data set. Biotic elements (clusters of species ranges) were delimited with model‐based Gaussian clustering. A second prediction of the vicariance model is that closely related species belong to different biotic elements. This was tested with a chi‐squared test. Results The distribution areas of the Iberian Helicoidea and the Israeli/Palestinian land snails are significantly clustered. The same is true for Israeli/Palestinian land snail species belonging to groups with geographically restricted species. However, the clustering is not significant in the complete central and eastern Aegean land snail data set, the Iberian Helicoidea species, and the central and eastern Aegean land snails belonging to groups with geographically restricted species. Contrary to the prediction of the vicariance model, closely related Iberian Helicoidea species and Israeli/Palestinian land snails belong significantly more often to the same biotic element than expected by chance. The null hypothesis that closely related species are homogeneously distributed across biotic elements cannot be rejected only for the data set that includes the Israeli/Palestinian land snails belonging to groups with geographically restricted species. Main conclusions The patterns found in the central and eastern Aegean land snail and the Iberian Helicoidea data sets do not correspond with the predictions of the vicariance model. This indicates that speciation modes other than vicariance were frequent, or that the distribution areas of many species in these faunas were largely modified by extensive post‐speciation dispersal and/or regional extinction. Hardly any possible vicariance events that might have contributed to the origin of the observed biotic elements in the Israeli/Palestinian land snail fauna could be identified. On the contrary, the biotic elements of the Israeli/Palestinian land snail fauna correspond with ecological factors. Vicariance has had only a limited influence on current biogeographical patterns in Mediterranean land snails.     

Rare plants are common where you find them

Broad patterns in distribution and abundance can elucidate processes of evolution. A positive association between local abundance and the size of the geographic range has been demonstrated for closely related species across many taxa. This pattern is usually explained by assuming that species with smaller ranges are ecologically inferior (e.g., poor competitors or dispersers). Many areas of high endemism support local species that have evolved recently. The distribution of these neoendemics may reflect historical processes not accounted for by ecological, equilibrium hypotheses. We asked whether such traditional macroecological hypotheses also applied to the local abundance of seven narrowly endemic species and ecologically similar widespread congeners in the northern Rocky Mountains. For each of the 14 species, we estimated abundance of five randomly chosen populations by counting plants in 10 randomly located plots. The association between range size and local abundance was not positive. Instead, all seven narrow endemics were more abundant than their widespread congeners. Ecological specialization or differences in dispersal ability are not likely explanations for our results. We believe the local abundance of narrowly endemic species may be a sign of recent speciation. Most or all of our narrowly distributed species have probably not yet had time to spread to their full potential. Furthermore, theory predicts that speciation is more likely to occur in locally abundant populations. Our results suggest that strictly ecological mechanisms cannot explain abundance and distribution in regions with high neoendemism.     

稻飞虱翅型纯系后代个体的翅型分化对光周期变化不敏感

【目的】为了明确光周期和遗传因子在稻飞虱翅型分化中的作用, 研究了3种稻飞虱（褐飞虱Nilaparvata lugens、白背飞虱Sogatella furcifera和灰飞虱Laodelphax striatellus）翅型纯系或近纯系在不同光照时数下的翅型分化比率。【方法】以经过5~45代连续翅型筛选后的褐飞虱、白背飞虱和灰飞虱的长翅型和短翅型纯系或近纯系为材料, 在室内分别测定了其在长光照（16和20 h）、短光照(4~12 h)和正常光照(14 h) 3类光周期条件下饲养后, 雌、雄成虫中长翅和短翅个体出现的比率及存活率。【结果】白背飞虱和灰飞虱的长翅型纯系M♂×M♀或短翅型纯系B♂×B♀在不同光周期下的翅型比率均无显著差异（P>0.05）。褐飞虱短翅型近纯系B♂×B♀的雌虫短翅率和成虫总短翅率在不同光周期下也无显著差异（P>0.05）, 但雄虫短翅率在正常光照14 h和短光照4 h下显著高于长光照20 h下的（P<0.05）。当褐飞虱短翅型达到纯系后, 其后代翅型在6~16 h光照条件下无显著差异。褐飞虱长翅型近纯系M♂×M♀的后代虽有短翅个体出现, 但是雌虫和雄虫的各自短翅率在不同光周期下无显著差异（P>0.05), 仅总体短翅率在12 h光照条件下的显著高于16 h下的（P<0.05）。褐飞虱长、短翅型杂交筛选品系M♂×B♀的雌虫短翅率随光照时数的延长而升高; 灰飞虱杂交筛选品系M♂×B♀的短翅雄虫随光照时数的缩短而增多（P<0.05）, 但当筛选代次达到45代时, 这种趋势不再显著。3种稻飞虱长翅型和短翅型纯系或近纯系若虫的存活率会稍低于长、短翅型杂交后代的存活率, 但长、短翅型品系的存活率在6~16 h光照条件下差异不显著(P>0.05)。【结论】稻飞虱翅型分化对光周期的反应受飞虱本身遗传背景的影响, 翅型纯系后代个体的翅型分化对光周期变化不敏感。     

The relationship between geographic range size and life history traits: is biogeographic history uncovered? A test using the Iberian butterflies

The geographic range of a species is influenced by past phylogenetic and biogeographic patterns. However, other historical interactions, including the interplay between life history and geography, are also likely involved. Therefore, the range size of a species can be explained on the basis of niche‐breadth or dispersal related hypotheses, and previous work on European butterflies suggests that both, under the respective guise of ecological specialisation and colonising ability may apply. In the present study, data from 205 species of butterflies from the Iberian peninsula were processed through multiple regression analyses to test for correlations between geographic range size, life history traits and geographic features of the species distribution types. In addition, the percentage of variance explained by the subsets of variables analyzed in the study, with and without control for phylogenetic effects was tested. Despite a complex pattern of bivariate correlations, we found that larval polyphagy was the single best correlate of range size, followed by dispersal. Models that combined both life history traits and geographic characteristics performed better than models generated independently. The combined variables explained at least 39% of the variance. Bivariate correlations between range size and body size, migratory habits or egg size primarily reflected taxonomic patterning and reciprocal correlations with larval diet breadth and adult phenology. Therefore, aspects of niche breadth i.e. potential larval diet breadth emerged as the most influential determinants of range size. However, the relationships between these types of ecological traits and biogeographic history must still be considered when associations between life history and range size are of interest.