Phylogeography of Parnassius apollo: hints on taxonomy and conservation of a vulnerable glacial butterfly invader

Parnassius apollo (Linnaeus, 1758) is probably the most renowned Eurasian montane butterfly. Its specialized ecology makes it very sensitive to habitat and climate changes, so that it is now experiencing range contraction and local extinction across most of its range. We sequenced 869 bp of the mitochondrial DNA (mtDNA) cytochrome oxidase I gene in 78 P. apollo populations (201 individuals) in order to: (1) assess the phylogeographic pattern of the species; (2) shed light on the historical biogeographic processes that shaped the distribution of the species; and (3) identify geographic population units of special value for the conservation of the species' genetic diversity. Our analyses revealed a very strong phylogeographic structure in P. apollo, which displays a number of distinctive mtDNA lineages populating geographically distinct areas. Overall sequence divergence is relatively shallow, and is consistent with a recent (late Pleistocene) colonization of most of the range. We propose that P. apollo is best viewed as an atypical glacial invader in southern and western Europe, the isolated, montane populations of which, threatened by climate warming, retain a large fraction of the species evolutionary heritage. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 169–183.     

Species traits help explain butterfly habitat use in a tropical forest

1. When species can access all parts of the landscape, species-sorting metacommunity theory predicts that community composition depends on habitat choice and interactions with other species and the environment. These filtering processes can also depend on species' traits.
2. The authors investigated how traits mediate a species-sorting process in determining butterfly community composition in a naturally patchy landscape in the tropical Western Ghats, India. The authors asked, do traits mediate access to certain habitats and does seasonality affect these patterns? The authors surveyed 56 habitat patches in three habitat types: laterite plateau grasslands, ridge grassland, and moist-deciduous forest, in a 65-km² landscape.
3. Non-palatable butterflies showed similar occurrences across seasons and habitats, but palatable butterflies were less commonly encountered in open habitats in the dry season. Polyphagous butterflies occurred infrequently in the dry season in laterite habitats, potentially indicating emigration or diapause patterns are linked to diet breadth.
4. All species were present in all habitats, implying dispersal does not limit access to different habitat patches, consistent with the species-sorting metacommunity concept. Nevertheless, butterfly occurrence was strongly influenced by the interaction of mobility and habitat type with sedentary species occurring less often in low-resource open laterite patches than mobile species.
5. Species sorting is typically regarded as occurring directly through environmental filters, but here the authors suggest that the environmental filter acts through movement limitations. Studies integrating landscape heterogeneity and species characteristics will help us better understand metacommunities and species distributions in nature.

     

Prediction of butterfly diversity hotspots in Belgium: a comparison of statistically focused and land use-focused models

Aim We evaluate differences between and the applicability of three linear predictive models to determine butterfly hotspots in Belgium for nature conservation purposes. Location The study is carried out in Belgium for records located to Universal Transverse Mercator (UTM) grid cells of 5 × 5 km. Methods We first determine the relationship between factors correlated to butterfly diversity by means of modified t‐tests and principal components analysis; subsequently, we predict hotspots using linear models based on land use, climate and topographical variables of well‐surveyed UTM grid cells (n = 197). The well‐surveyed squares are divided into a training set and an evaluation set to test the model predictions. We apply three different models: (1) a ‘statistically focused’ model where variables are entered in descending order of statistical significance, (2) a ‘land use‐focused’ model where land use variables known to be related to butterfly diversity are forced into the model and (3) a ‘hybrid’ model where the variables of the ‘land use‐focused model’ are entered first and subsequently complemented by the remaining variables entered in descending order of statistical significance. Results A principal components analyses reveals that climate, and to a large extent, land use are locked into topography, and that topography and climate are the variables most strongly correlated with butterfly diversity in Belgium. In the statistically focused model, biogeographical region alone explains 65% of the variability; other variables entering the statistically focused model are the area of coniferous and deciduous woodland, elevation and the number of frost days; the statistically focused model explains 77% of the variability in the training set and 66% in the evaluation set. In the land use‐focused model, biogeographical region, deciduous and mixed woodland, natural grassland, heathland and bog, woodland edge, urban and agricultural area and biotope diversity are forced into the model; the land use‐focused model explains 68% of the variability in the training set and 57% in the evaluation set. In the hybrid model, all variables from the land use‐focused model are entered first and the covariates elevation, number of frost days and natural grassland area are added on statistical grounds; the hybrid model explains 78% of the variability in the training set and 67% in the evaluation set. Applying the different models to determine butterfly diversity hotspots resulted in the delimitation of spatially different areas. Main conclusions The best predictions of butterfly diversity in Belgium are obtained by the hybrid model in which land use variables relevant to butterfly richness are entered first after which climatic and topographic variables were added on strictly statistical grounds. The land use‐focused model does not predict butterfly diversity in a satisfactory manner. When using predictive models to determine butterfly diversity, conservation biologists need to be aware of the consequences of applying such models. Although, in conservation biology, land use‐focused models are preferable to statistically focused models, one should always check whether the applied model makes sense on the ground. Predictive models can target mapping efforts towards potentially species‐rich sites and permits the incorporation of un‐surveyed sites into nature conservancy policies. Species richness distribution maps produced by predictive modelling should therefore be used as pro‐active conservation tools.     

广州市不同城市化发展区域蝶类多样性

2005～2006年,对广州地区4个不同城市化发展区域(森林区、农田区、城市区、沿海湿地区)的蝶类进行6次调查,共统计到10科46属73种.森林区科、属、物种和个体数目都最多,沿海湿地区各项数量指标都最小,农田区的个体数量大于城市区,但科、属和物种数却与城市区差异不大.多样性分析结果表明,森林区的种类丰度、多样性指数最高,沿海湿地区的种类丰度和多样性指数最低;均匀度大小依次为沿海湿地区﹥农田区﹥森林区﹥城市区;优势度指数大小依次为沿海湿地区﹥城市区﹥农田区﹥森林区.相似性分析结果表明,森林区和农田区具有的相同物种数最多,相似性系数最高;沿海湿地区和森林区、农田区、城市区具有的相同物种数都较少,相似性系数也较低.     

The case of the monarch butterfly: a verdict is returned

A publication reporting the harmful effects on the monarch butterfly of maize genetically modified to express insecticidal delta-endotoxins from the soil bacterium Bacillus thuringiensis (Bt) caused much public interest. A series of ecologically based studies were subsequently carried out to evaluate rigorously the impact of pollen from such crops and to quantify the risks. The results demonstrated that the commercial large-scale cultivation of current Bt-maize hybrids did not pose a significant risk to the monarch population. Further studies also demonstrated that Bt-expressing crops posed little risk to other nontarget insects, including beneficial insects such as pollinators and natural enemies.     

Environmental controls on butterfly occurrence and species richness in Israel: The importance of temperature over rainfall

Butterflies are considered important indicators representing the state of biodiversity and key ecosystem functions, but their use as bioindicators requires a better understanding of how their observed response is linked to environmental factors. Moreover, better understanding how butterfly faunas vary with climate and land cover may be useful to estimate the potential impacts of various drivers, including climate change, botanical succession, grazing, and afforestation. It is particularly important to establish which species of butterflies are sensitive to each environmental driver.The study took place in Israel, including the West Bank and Golan Heights.To develop a robust and systematic approach for identifying how butterfly faunas vary with the environment, we analyzed the occurrence of 73 species and the abundance of 24 species from Israeli Butterfly Monitoring Scheme (BMS‐IL) data. We used regional generalized additive models to quantify butterfly abundance, and generalized linear latent variable models and generalized linear models to quantify the impact of temperature, rainfall, soil type, and habitat on individual species and on the species community.Species richness was higher for cooler transects, and also for hilly and mountainous transects in the Mediterranean region (rendzina and Terra rossa soils) compared with the coastal plain (Hamra soil) and semiarid northern Jordan Vale (loessial sierozem soil). Species occurrence was better explained by temperature (negative correlation) than precipitation, while for abundance the opposite pattern was found. Soil type and habitat were insignificant drivers of occurrence and abundance.Butterfly faunas responded very strongly to temperature, even when accounting for other environmental factors. We expect that some butterfly species will disappear from marginal sites with global warming, and a large proportion will become rarer as the region becomes increasingly arid.     

Conserving a wetland butterfly: quantifying early lifestage survival through seasonal flooding, adult nectar, and habitat preference

Conservation of once thought extinct populations of Lycaena xanthoides in western Oregon will require specific information of how the butterfly interacts with its now rare wetland habitat. Three experiments were conducted to yield information directly applicable to wetland restoration work: (1) to quantify the survival of L. xanthoides eggs laid on inundated plants, (2) to quantify adult nectar preferences, (3) to investigate the role of adult resources and how butterflies assess habitat quality. Survival of eggs laid on inundated plants was nearly seven times lower than the survival of eggs laid on uninundated plants, indicating that eggs laid on plants that are seasonally flooded are a population sink. Adult L. xanthoides preferred an endemic native nectar plant, Grindelia integrifolia × nana, as an adult resource and used it approximately 88% of the time while other butterfly species preferred to nectar on the non-native Mentha pulegium. Adult L. xanthoides had a significantly greater short-term recapture rate in two restored study sites that had a high relative amount of Grindelia integrifolia × nana compared to a degraded site that lacked the preferred nectar source. Based on the results from the three small experiments, restoration of wetlands for L. xanthoides should concentrate plantings of host plant in non-flooded areas and propagate conspicuous patches of the preferred nectar plant.     

Additive partitioning of butterfly diversity in a fragmented landscape: importance of scale and implications for conservation

Aim Most of the Atlantic Forest in Brazil occurs in fragments of various sizes. Previous studies indicate that forest fragmentation affects fruit‐feeding butterflies. Conservation strategies that seek to preserve organisms that are distributed in high‐fragmented biomes need to understand the spatial distribution of these organisms across the landscape. In view of the importance of understanding the fauna of these forest remnants, the objective of the present work is to investigate the extent to which the diversity of this group varies across spatial scales ranging from within‐forest patches to between landscapes. Location South America, south‐eastern Brazil, São Paulo State. Methods We used bait traps to sample fruit feeding butterflies at 50 points in 10 fragments in two different landscapes during a period of 12 months. Total species richness and Shannon index were partitioned additively in diversity at trap level, and beta diversity was calculated among traps, among forest patches, and between landscapes. We used permutation tests to compare these values to the expected ones under the null hypothesis that beta diversity is only a random sampling effect. Results There was significant beta diversity at the smallest scale examined; however, the significance at higher scales depends on the diversity measurement used. Beta diversity with Shannon index was smaller than expected by chance among fragments, whereas species richness was not. Among landscapes, only beta diversity in richness was higher than expected by chance. Main conclusions The results observed occur because there is great variability in species composition among forest patches in the same landscape, changing this diversity even though the communities are formed from the same pool of species. At the largest scale evaluated (between landscapes), these pattern changes and differences in beta diversity in richness were detectable. This difference is probably caused by the presence of rare species. Thus, a conservation strategy that seeks to preserve as many species as possible per unit of area in high‐fragmented biomes should give priority to protecting fragments in different landscapes, rather than more fragments in the same landscape.     

Patterns of butterfly distribution in the Andaman islands: implications for conservation

Twenty-five islands of different sizes were rapidly surveyed in the Andaman islands for patterns of butterfly distribution and abundance. The surveys were conducted in the dry seasons of 1992 in the South Andaman islands, 1994 in the North Andaman islands and on both these years on the Little Andaman Island. Different habitat types were identified on each island and butterflies were sampled by the line transect method in each habitat type. Sixty-five species of butterflies were recorded from six families. Fifty-one species were less common and contributed to 25 % of the total count. Six species were very common. The overall distribution patterns of the species were nested. This suggests that small islands share their species with the larger islands but not vice versa. Many uncommon species were found exclusively on large islands. The presence of evergreen forest on islands significantly influenced the species encountered. Small and medium sized islands with evergreen forests had significantly more species than those without evergreen forests. Loss of primary forests due to logging and encroachment will result in the loss of many butterfly species. It is recommended that the large patches of primary evergreen forests be protected on a priority basis on large islands.     

Preservation of biodiversity in small rainforest patches: rapid evaluations using butterfly trapping

Determining the capacity of small forest remnants to support biodiversity is of critical importance, especially in the tropics where high rates of land conversion coincide with extraordinarily high species richness and endemism. Using fruit-baited traps, we conducted rapid evaluations in 1993 and 1994 of the forest butterfly diversity of seven small patches (3–30 ha) and a single remaining large patch (227 ha) of Costa Rican mid-elevation moist forest. Our results suggest that even recently isolated 20–30 ha fragments of primary forest retain surprisingly depauperate butterfly faunas relative to that supported by the 227 ha patch only 0.5–1.0 km away. If forest butterflies are an index of the diversity of small-bodied organisms in general, preservation of the latter may require unexpectedly large patches. In 1994 we also surveyed a 16 ha botanical garden, situated between and contiguous to both the 227 ha patch and an exceptionally species-rich 25 ha patch. In the garden, we discovered adults of many butterfly species associated with forest interior, suggesting that even heavily managed systems of largely exotic plants (such as agricultural systems) could be designed to serve as corridors for butterflies and perhaps some other groups of organisms. We discuss some implications for a planned restoration of biotic connections between lowland and montane forests in southern Costa Rica.     

Lack of gene flow: Narrow and dispersed differentiation islands in a triplet of Leptidea butterfly species

Genome scans in recently separated species can inform on molecular mechanisms and evolutionary processes driving divergence. Large‐scale polymorphism data from multiple species pairs are also key to investigate the repeatability of divergence—whether radiations tend to show parallel responses to similar selection pressures and/or underlying molecular forces. Here, we used whole‐genome resequencing data from six wood white (Leptidea sp.) butterfly populations, representing three closely related species with karyomorph variation, to infer the species' demographic history and characterize patterns of genomic diversity and differentiation. The analyses supported previously established species relationships, and there was no evidence for postdivergence gene flow. We identified significant intraspecific genetic structure, in particular between karyomorph extremes in the wood white (L. sinapis)—a species with a remarkable chromosome number cline across the distribution range. The genomic landscapes of differentiation were erratic, and outlier regions were narrow and dispersed. Highly differentiated (F_ST) regions generally had low genetic diversity (θ_π), but increased absolute divergence (D_XY) and excess of rare frequency variants (low Tajima's D). A minority of differentiation peaks were shared across species and population comparisons. However, highly differentiated regions contained genes with overrepresented functions related to metabolism, response to stimulus and cellular processes, indicating recurrent directional selection on a specific set of traits in all comparisons. In contrast to the majority of genome scans in recently diverged lineages, our data suggest that divergence landscapes in Leptidea have been shaped by directional selection and genetic drift rather than stable recombination landscapes and/or introgression.     

基于地形因子的三峡库区腹地耕地演变——以草堂溪流域为例

以1990、2000、2004和2007年4期遥感影像数据和1∶5万的DEM数据为数据源,利用ArcGIS 9.3空间分析功能和Fragstas 3.3景观格局分析功能对库区腹地耕地演变进行了研究.首先建立草堂溪流域耕地数据库以及高程、坡度和地形位指数图,然后将高程、坡度和地形位梯度分别与耕地数据进行图层运算,提取坡耕地的面积,探讨不同地形下耕地的分布特征.结果表明:耕地在各地形因素上的分布表现为15° ～35°坡度带、500～1000 m高程级别、东南坡及南坡以及中高地形上.1990-2007年,研究区耕地在面积和空间格局上均发生很大的变化,随地形梯度的增大,4个时期耕地分布面积随地形位指数的增大表现为先升后降;聚集度减少特别显著的主要集中在第4(1.2～1.5)和5(1.5 ～2.0)两个地形梯度带,即中高地形位的耕地之间聚集程度降低,破碎程度加剧,生态环境好转.     

长江三峡库区蝶类群落的物种多样性

报道了多样性指数、丰富度和均匀度.各小生境物种丰富度的变化在0～28之间,其下限值的数理分布范围主要在0与5之间,而上限值的分布主要在9、11和12.物种多样性指数的变化范围是：样本间0～4.9285,小生境间0～2.1143,生境类型间0～1.7091和植被型0.9740～1.3143;而物种多样性指数最高的样本（4.9285）在1500～1500m针阔混交林的小生境中,物种多样性指数最高的小生境（2.1143）是1000～1500m阔叶林,物种多样性指数最高的生境类型是阔叶林灌丛（1.4373）,物种多样性指数最高的植被型是灌丛（1.3143）;而灌丛各生境类型的物种多样性指数最高,从1.2773到1.4373;草地居第二位1.0588～1.2402,森林最低,0.8088～0.9618（仅分布在一个海拔梯度的生境类型除外）;就整体有而言,物种多样性指数居前5位的生境类型都是灌丛.可见,三峡库区灌丛是最适宜于蝴蝶繁衍的植被型;完全成片的森林与纯粹的农田均不太适合蝴蝶生存.因环境的复杂性,物种多样性指数的最高值（4.9285）和最低值（0.0）均出现森林植被型,样本间物种多样性指数变异系数最大的也是森林植被型,表明它给蝴蝶生存的影响,也将是最复杂的.海拔高度对物种多样性指数的影响明显,最高的物种多样性指数出现在海拔1000～1500m（1.2363）,最低是2000m以上（0.2536）,但没有显著的规律性.灌丛和森林各个小生境的蝴蝶均匀度与物种多样性的变化趋势基本是一致的,表明这两种植被型内各小生境的物种多样性指数主要受物种均匀度变化的影响;农田和草地两种植被型的物种多样性指数的变化除受均匀度影响外,还较大程度地受物种丰富度的影响.在生境类型的水平上,蝴蝶均匀度与物种多样性指数、丰富度间有着比较复杂的关系.这些结果反映了三峡库区蝶类生存环境的多样性和破碎化,从而影响了蝴蝶分布的丰富度、多样性和均匀性.     

Rapid assessment of butterfly diversity in a montane landscape

We present the results of a rapid assessment of butterfly diversity in the 754 ha Beaver Meadows study area in Rocky Mountain National Park, Larimer County, Colorado. We measured butterfly species richness and relative abundance as part of a landscape-scale investigation of diversity patterns involving several groups of organisms. A stratified random sampling design was used to include replication in both rare and common vegetation types. We recorded 49 butterfly species from the twenty-four 0.1 ha plots that were sampled four times during June, July, and August 1996. Butterfly species richness, diversity, and uniqueness were highest in quaking aspen (Populus tremuloides Michaux) groves and wet meadows, which occupy only a small proportion of the studied landscape. This result supports the suggestion that aspen areas represent hotspots of biological diversity in this montane landscape. Patterns of butterfly species richness were positively correlated with total vascular plant species richness (r = 0.69; P < 0.001), and native plant species richness (r = 0.64; P < 0.001). However, exotic plant species richness (r = 0.70; P < 0.001) and the cover of exotic plant species (r = 0.70; P < 0.001) were the best predictors of butterfly species richness.     

Quantifying the ecological stability of a phytoplankton community: The Lake Kinneret case study

The widely used term “stability” has multiple meanings and is rarely quantified in limnological studies. The main objective of this study was to develop an approach for quantifying the stability of a phytoplankton community using Lake Kinneret as a case study. It is a first attempt of calculating an index of stability for each of the five main taxonomic groups of the Kinneret phytoplankton (Bacillariophyta, Chlorophyta, Cryptophyta, Cyanophyta and Dinophyta), and for the entire community. A simple statistical approach to calculate the stability index was devised, using phytoplankton wet-weight biomass as the parameter being manipulated. The period 1970–1979 was selected as a reference period. The following stability indices were established and applied (each at three time scales): (1) a stability index for each of five main taxonomic groups; (2) a combined index of the stability, aggregating the stabilities of the individual taxonomic groups and (3) a stability index of entire community based on total phytoplankton biomass. The dynamics of these indices during 1969–2011 were examined. Destabilization of the community structure was triggered by an increase in the variability of Bacillariophyta biomass shortly after the reference period, in 1981–1983. Only 10 years later, the community destabilization become associated with progressively increasing biomass of Cyanobacteria. Dinophyta were the last to destabilize in the mid 1990s. Despite notable changes in the community structure, the total phytoplankton biomass remained relatively stable. Therefore, in 1969–2011 the stability index based on total phytoplankton biomass was higher than the combined index based on the stabilities of the individual taxonomic groups. Only weak relationships were found between the stability index values and potential driving forces (lake water level fluctuations and nutrient loads). While this approach was applied to Lake Kinneret, the concept presented is not lake specific and could be applied to other lakes.