An experimental examination of the effects of habitat quality on the dispersal and local abundance of the butterfly Parnassius smintheus

Abstract 1. Nectar flower abundance was manipulated through flower removal, and sex ratio was manipulated by moving individual butterflies within a series of nine alpine meadows. The movement and abundance of the butterfly Parnassius smintheus in the meadows were monitored using mark–release–recapture methods.
2. A total of 937 butterflies, 698 males and 239 females, was captured. There were 223 observed between-meadow movements. Fifty-two per cent of males and 35% of females moved among meadows.
3. The immigration of male butterflies was related positively to nectar flowers, host plant abundance, and female butterflies. Male emigration was not affected by any of the treatments. The number of males captured was related positively to nectar flowers and host plants but not affected by sex ratio. The number of resident male butterflies was greater in meadows containing flowers and was related positively to host plant abundance, but unaffected by sex ratio.
4. Flower removal, sex ratio, and abundance of Sedum had no significant effect on the abundance, movement, or residence time for female butterflies, in part due to small sample size.
5. The fact that males immigrate to higher quality meadows suggests that male butterflies are assessing meadow quality, either by sampling meadows or potentially from a distance using olfactory cues.     

Inclusion of trophic interactions increases the vulnerability of an alpine butterfly species to climate change

Climate change is expected to have significant and complex impacts on ecological communities. In addition to direct effects of climate on species, there can also be indirect effects through an intermediary species, such as in host–plant interactions. Indirect effects are expected to be more pronounced in alpine environments because these ecosystems are sensitive to temperature changes and there are limited areas for migration of both species (i.e. closed systems), and because of simpler trophic interactions. We tested the hypothesis that climate change will reduce the range of an alpine butterfly (Parnassius smintheus) because of indirect effects through its host plant (Sedum sp.). To test for direct and indirect effects, we used the simulations of climate change to assess the distribution of P. smintheus with and without Sedum sp. We also compared the projected ranges of P. smintheus to four other butterfly species that are found in the alpine, but that are generalists feeding on many plant genera. We found that P. smintheus gained distributional area in climate‐only models, but these gains were significantly reduced with the inclusion of Sedum sp. and in dry‐climate scenarios which resulted in a reduction in net area. When compared to the more generalist butterfly species, P. smintheus exhibited the largest loss in suitable habitat. Our findings support the importance of including indirect effects in modelling species distributions in response to climate change. We highlight the potentially large and still neglected impacts climate change can have on the trophic structure of communities, which can lead to significant losses of biodiversity. In the future, communities will continue to favour species that are generalists as climate change induces asynchronies in the migration of species.     

Genetic structure of French populations of the mountain butterfly Parnassius mnemosyne L. (Lepidoptera: Papilionidae)

Geographical variation in allozyme frequencies among 24 populations of the butterfly Parnassius mnemosyne in southern France was studied by enzyme electrophoresis. Clustering procedures were used to define the geographical organization of genetic variation and an analysis of genetic differentiation was performed to test the degree of potential gene flow between the colonies using the methodologies derived from F-statistics and the private alleles model. Further, an approach using reduced space ordinations (factorial analysis of correspondences and discriminant analysis), was used to allow the visualization of genetically intermediate populations and to determine the rôles of individual alleles at various levels of geographical structure. In P. mnemosyne , the effect of both isolation by distance and geographical barriers upon gene flow is complex. Some barriers appear to be very efficient, but others, at least as strong geographically, do not act in such a way. Multivariate analyses underlined the progressive transition between some groups while in other instances, a sharp cut was observed without obvious topographical barriers. Ecogeographical data and historical events (such as colonization processes) should probably be taken into account in order to explain the geographical variation of genetic structure.     

Simulating effects of climate change under direct and diapause development in a butterfly

Temperature is one of the most important ecological factors affecting species survival and distributions. Therefore, global climate change, involving increases in mean surface temperature and the occurrence of extreme weather events, may pose a substantial challenge to biodiversity. Whereas tropical ectotherms are believed to be very sensitive to climate change, temperate‐zone species may actually benefit from higher temperatures. However, as in temperate zones large parts of the year are unsuitable for growth and reproduction, seasonal time constraints may complicate matters. Against this background we here investigate the impact of simulated climate change, involving increased mean temperatures and heat waves, across developmental pathways of the butterfly Lycaena tityrus (Poda) (Lepidoptera: Lycaenidae). Increased temperatures speeded up development but decreased pupal mass as expected. However, we found no evidence for detrimental effects of increased temperatures or even simulated heat waves. Furthermore, patterns did not differ between indirectly and directly developing individuals, which are assumed to be more time constrained. Our findings support the notion that not all species will be detrimentally affected by climate change, and suggest that species attributes may be more important than potential time constraints imposed by different developmental pathways.     

Projecting demographic responses to climate change: adult and juvenile survival respond differently to direct and indirect effects of weather in a passerine population

Few studies have quantitatively projected changes in demography in response to climate change, yet doing so can provide important insights into the processes that may lead to population declines and changes in species distributions. Using a long‐term mark‐recapture data set, we examined the influence of multiple direct and indirect effects of weather on adult and juvenile survival for a population of Song Sparrows (Melospiza melodia) in California. We found evidence for a positive, direct effect of winter temperature on adult survival, and a positive, indirect effect of prior rainy season precipitation on juvenile survival, which was consistent with an effect of precipitation on food availability during the breeding season. We used these relationships, and climate projections of significantly warmer and slightly drier winter weather by the year 2100, to project a significant increase in mean adult survival (12–17%) and a slight decrease in mean juvenile survival (4–6%) under the B1 and A2 climate change scenarios. Together with results from previous studies on seasonal fecundity and postfledging survival in this population, we integrated these results in a population model and projected increases in the population growth rate under both climate change scenarios. Our results underscore the importance of considering multiple, direct, and indirect effects of weather throughout the annual cycle, as well as differences in the responses of each life stage to climate change. Projecting demographic responses to climate change can identify not only how populations will be affected by climate change but also indicate the demographic process(es) and specific mechanisms that may be responsible. This information can, in turn, inform climate change adaptation plans, help prioritize future research, and identify where limited conservation resources will be most effectively and efficiently spent.     

Cumulative effects of climate and landscape change drive spatial distribution of Rocky Mountain wolverine (Gulo gulo L.)

Contemporary landscapes are subject to a multitude of human‐derived stressors. Effects of such stressors are increasingly realized by population declines and large‐scale extirpation of taxa worldwide. Most notably, cumulative effects of climate and landscape change can limit species’ local adaptation and dispersal capabilities, thereby reducing realized niche space and range extent. Resolving the cumulative effects of multiple stressors on species persistence is a pressing challenge in ecology, especially for declining species. For example, wolverines (Gulo gulo L.) persist on only 40% of their historic North American range. While climate change has been shown to be a mechanism of range retractions, anthropogenic landscape disturbance has been recently implicated. We hypothesized these two interact to effect declines. We surveyed wolverine occurrence using camera trapping and genetic tagging at 104 sites at the wolverine range edge, spanning a 15,000 km² gradient of climate, topographic, anthropogenic, and biotic variables. We used occupancy and generalized linear models to disentangle the factors explaining wolverine distribution. Persistent spring snow pack—expected to decrease with climate change—was a significant predictor, but so was anthropogenic landscape change. Canid mesocarnivores, which we hypothesize are competitors supported by anthropogenic landscape change, had comparatively weaker effect. Wolverine population declines and range shifts likely result from climate change and landscape change operating in tandem. We contend that similar results are likely for many species and that research that simultaneously examines climate change, landscape change, and the biotic landscape is warranted. Ecology research and species conservation plans that address these interactions are more likely to meet their objectives.     

Avoiding verisimilitude when modelling ecological responses to climate change: the influence of weather conditions on trapping efficiency in European badgers (Meles meles)

The signal for climate change effects can be abstruse; consequently, interpretations of evidence must avoid verisimilitude, or else misattribution of causality could compromise policy decisions. Examining climatic effects on wild animal population dynamics requires ability to trap, observe or photograph and to recapture study individuals consistently. In this regard, we use 19 years of data (1994–2012), detailing the life histories on 1179 individual European badgers over 3288 (re‐) trapping events, to test whether trapping efficiency was associated with season, weather variables (both contemporaneous and time lagged), body‐condition index (BCI) and trapping efficiency (TE). PCA factor loadings demonstrated that TE was affected significantly by temperature and precipitation, as well as time lags in these variables. From multi‐model inference, BCI was the principal driver of TE, where badgers in good condition were less likely to be trapped. Our analyses exposed that this was enacted mechanistically via weather variables driving BCI, affecting TE. Notably, the very conditions that militated for poor trapping success have been associated with actual survival and population abundance benefits in badgers. Using these findings to parameterize simulations, projecting best‐/worst‐case scenario weather conditions and BCI resulted in 8.6% ± 4.9 SD difference in seasonal TE, leading to a potential 55.0% population abundance under‐estimation under the worst‐case scenario; 38.6% over‐estimation under the best case. Interestingly, simulations revealed that while any single trapping session might prove misrepresentative of the true population abundance, due to weather effects, prolonging capture–mark–recapture studies under sub‐optimal conditions decreased the accuracy of population estimates significantly. We also use these projection scenarios to explore how weather could impact government‐led trapping of badgers in the UK, in relation to TB management. We conclude that population monitoring must be calibrated against the likelihood that weather conditions could be altering trap success directly, and therefore biasing model design.     

长白山红松不同树高处径向生长特征及其对气候的响应

利用长白山红松不同树高(0.3、1.3、4、10、15、20、25 m)处的径向生长资料,分析各树高处径向生长特征,建立红松生长与气候因子的相关关系,以期完善红松种群对气候变化的响应机制。结果表明:(1)红松不同树高处年径向生长量变化趋势基本一致,除在1980年前后,20 m处径向生长量出现异常增加外,其他各高度径向生长均出现下降趋势,红松基部和顶端(0.3、1.3 m和20 m)处径向生长年际变化更明显。随着树高增加,各处年径向生长率有所降低,0.3m处生长速率最大,且与10 m和15 m处径向生长差异显著(P < 0.05)。(2)不同树高处径向生长对气候因子的响应存在明显差异,10 m树高是红松径向生长对温度和降水响应差异的分界线。10 m以下红松径向生长主要受到生长季温度的负作用,尤其是4 m处,与当年生长季初期(4月和5月)温度显著负相关(P < 0.05)。0.3 m和1.3 m处径向生长分别与上年9月平均温度显著正相关(P < 0.05),当年6月平均和最高温度显著负相关(P < 0.05)。随着树高上升,降水对径向生长的促进作用增强,而温度对径向生长的作用也发生改变。10 m(含)以上则受到温度和降水的共同作用。10 m处径向生长对气候因子响应最敏感,受到当年生长季高温的抑制作用,还与上年和当年生长季末(9月)降水显著正相关(P < 0.05)。15 m处径向生长与上年9月最低温度和降水显著正相关(P < 0.05),而与当年5月月平均温度显著负相关(P < 0.05)。20 m处径向生长与当年3月月平均、最低和最高温度,当年7月月平均温度以及当年5月降水显著正相关(P < 0.05),而与当年1月降水显著负相关(P < 0.05)。     

Population growth potential of Tuta absoluta (Meyrick) (Lepidoptera: Gelechiidae) on tomato,potato, and eggplant

The influence of host plant on population dynamics of an invasive pest, Tuta absoluta was studied on three economically important solanaceous crops. Experiments were conducted in laboratory (29 ± 0.5°C, 75 ± 5% RH and a photoperiod of 14:10 hr [L:D]) using tomato (Solanum lycopersicum L.), potato (Solanum tuberosum L.) and eggplant (Solanum melongena L.). Results indicated that intrinsic rate of increase (r), finite rate of increase (λ) and net reproductive rate (R₀) were higher, and mean generation time (T) was the shortest on tomato. Results suggested that T. absoluta developed on all the three plants, and tomato plant was most preferred one. Results suggested that T. absoluta has a potential to become a serious pest on potato and even on eggplant under favourable conditions. We used the life tables of 0.025th and 0.975th percentiles of bootstraps to project the uncertainty of population growth, a new concept.     

Spatial heterogeneity in climate change effects decouples the long‐term dynamics of wild reindeer populations in the high Arctic

The ‘Moran effect’ predicts that dynamics of populations of a species are synchronized over similar distances as their environmental drivers. Strong population synchrony reduces species viability, but spatial heterogeneity in density dependence, the environment, or its ecological responses may decouple dynamics in space, preventing extinctions. How such heterogeneity buffers impacts of global change on large‐scale population dynamics is not well studied. Here, we show that spatially autocorrelated fluctuations in annual winter weather synchronize wild reindeer dynamics across high‐Arctic Svalbard, while, paradoxically, spatial variation in winter climate trends contribute to diverging local population trajectories. Warmer summers have improved the carrying capacity and apparently led to increased total reindeer abundance. However, fluctuations in population size seem mainly driven by negative effects of stochastic winter rain‐on‐snow (ROS) events causing icing, with strongest effects at high densities. Count data for 10 reindeer populations 8–324 km apart suggested that density‐dependent ROS effects contributed to synchrony in population dynamics, mainly through spatially autocorrelated mortality. By comparing one coastal and one ‘continental’ reindeer population over four decades, we show that locally contrasting abundance trends can arise from spatial differences in climate change and responses to weather. The coastal population experienced a larger increase in ROS, and a stronger density‐dependent ROS effect on population growth rates, than the continental population. In contrast, the latter experienced stronger summer warming and showed the strongest positive response to summer temperatures. Accordingly, contrasting net effects of a recent climate regime shift—with increased ROS and harsher winters, yet higher summer temperatures and improved carrying capacity—led to negative and positive abundance trends in the coastal and continental population respectively. Thus, synchronized population fluctuations by climatic drivers can be buffered by spatial heterogeneity in the same drivers, as well as in the ecological responses, averaging out climate change effects at larger spatial scales.     

寄主作物对B型烟粉虱生长发育和种群增殖的影响

在恒温28℃、湿度80％的条件下研究了5种寄主植物对B型烟粉虱(Bemisia tubaci)生长、发育和繁殖的影响。结果表明：取食棉花、烟草、花生、大豆和玉米的烟粉虱。其形态、发育历期、存活率、成虫繁殖力和寿命等生命参数均有明显差异。棉花、大豆和玉米不利于烟粉虱卵和若虫的生长发育。表现为虫体较小、卵期和生命周期延长、存活率降低。烟粉虱在棉花、大豆、花生、烟草和玉米上的生命周期分别为32．03d、32．11d、25．69d、24．43d和20．68d。其存活率分别为49．86％、54．41％、86．86％、69．93％和29．38％。与烟草和玉米相比。棉花、大豆和花生明显有利于烟粉虱的种群繁殖．在棉花、大豆、花生、烟草和玉米上生长发育的成虫寿命和单雌产卵量分别是27．8d和235．O粒、23．2d和191．1粒、22．Od和131．1粒、6．25d和28．O粒、2．42d和5．1粒。在花生、大豆、棉花、烟草和玉米上的内禀增长率(rm)分别为O．1590、O．1364、O．1236、O．0841和-O．0285．其种群趋势指数(I)分别为113．85、117．38、103．98、19．58和O．4274。     

Pleistocene evolutionary history of the Clouded Apollo (Parnassius mnemosyne): genetic signatures of climate cycles and a 'time-dependent' mitochondrial substitution rate

Genetic data are currently providing a large amount of new information on past distribution of species and are contributing to a new vision of Pleistocene ice ages. Nonetheless, an increasing number of studies on the 'time dependency' of mutation rates suggest that date assessments for evolutionary events of the Pleistocene might be overestimated. We analysed mitochondrial (mt) DNA (COI) sequence variation in 225 Parnassius mnemosyne individuals sampled across central and eastern Europe in order to assess (i) the existence of genetic signatures of Pleistocene climate shifts; and (ii) the timescale of demographic and evolutionary events. Our analyses reveal a phylogeographical pattern markedly influenced by the Pleistocene/Holocene climate shifts. Eastern Alpine and Balkan populations display comparatively high mtDNA diversity, suggesting multiple glacial refugia. On the other hand, three widely distributed and spatially segregated lineages occupy most of northern and eastern Europe, indicating postglacial recolonization from different refugial areas. We show that a conventional 'phylogenetic' substitution rate cannot account for the present distribution of genetic variation in this species, and we combine phylogeographical pattern and palaeoecological information in order to determine a suitable intraspecific rate through a Bayesian coalescent approach. We argue that our calibrated 'time-dependent' rate (0.096 substitutions/ million years), offers the most convincing time frame for the evolutionary events inferred from sequence data. When scaled by the new rate, estimates of divergence between Balkan and Alpine lineages point to c. 19 000 years before present (last glacial maximum), and parameters of demographic expansion for northern lineages are consistent with postglacial warming (5-11 000 years before present).     

Synergistic effects of the invasive Chinese tallow (Triadica sebifera) and climate change on aquatic amphibian survival

Changes in climate and the introduction of invasive species are two major stressors to amphibians, although little is known about the interaction between these two factors with regard to impacts on amphibians. We focused our study on an invasive tree species, the Chinese tallow (Triadica sebifera), that annually sheds its leaves and produces leaf litter that is known to negatively impact aquatic amphibian survival. The purpose of our research was to determine whether the timing of leaf fall from Chinese tallow and the timing of amphibian breeding (determined by weather) influence survival of amphibian larvae. We simulated a range of winter weather scenarios, ranging from cold to warm, by altering the relative timing of when leaf litter and amphibian larvae were introduced into aquatic mesocosms. Our results indicate that amphibian larvae survival was greatly affected by the length of time Chinese tallow leaf litter decomposes in water prior to the introduction of the larvae. Larvae in treatments simulating warm winters (early amphibian breeding) were introduced to the mesocosms early in the aquatic decomposition process of the leaf litter and had significantly lower survival compared with cold winters (late amphibian breeding), likely due to significantly lower dissolved oxygen levels. Shifts to earlier breeding phenology, linked to warming climate, have already been observed in many amphibian taxa, and with most climate models predicting a significant warming trend over the next century, the trend toward earlier breeding should continue if not increase. Our results strongly suggest that a warming climate can interact with the effects of invasive plant species, in ways we have not previously considered, to reduce the survival of an already declining group of organisms.     

Land‐use and climate change effects on population size and extinction risk of Andean plants

Andean plant species are predicted to shift their distributions, or ‘migrate,’ upslope in response to future warming. The impacts of these shifts on species' population sizes and their abilities to persist in the face of climate change will depend on many factors including the distribution of individuals within species' ranges, the ability of species to migrate and remain at equilibrium with climate, and patterns of human land‐use. Human land‐use may be especially important in the Andes where anthropogenic activities above tree line may create a hard barrier to upward migrations, imperiling high‐elevation Andean biodiversity. In order to better understand how climate change may impact the Andean biodiversity hotspot, we predict the distributional responses of hundreds of plant species to changes in temperature incorporating population density distributions, migration rates, and patterns of human land‐use. We show that plant species from high Andean forests may increase their population sizes if able to migrate onto the expansive land areas above current tree line. However, if the pace of climate change exceeds species' abilities to migrate, all species will experience large population losses and consequently may face high risk of extinction. Using intermediate migration rates consistent with those observed for the region, most species are still predicted to experience population declines. Under a business‐as‐usual land‐use scenario, we find that all species will experience large population losses regardless of migration rate. The effect of human land‐use is most pronounced for high‐elevation species that switch from predicted increases in population sizes to predicted decreases. The overriding influence of land‐use on the predicted responses of Andean species to climate change can be viewed as encouraging since there is still time to initiate conservation programs that limit disturbances and/or facilitate the upward migration and persistence of Andean plant species.     

Effects of climate change on different geographical populations of the cotton bollworm Helicoverpa armigera (Lepidoptera,Noctuidae)

The effects of climate change on pest phenology and population size are highly variable. Understanding the impacts of localized climate change on pest distribution and phenology is helpful for improving integrated pest management strategies. Here, the population dynamics of cotton bollworms (Helicoverpa armigera) from Maigaiti County, south Xinjiang, and Shawan County, north Xinjiang, China, were analyzed using a 29‐year dataset at lower latitudes and a 23‐year dataset at higher latitudes to determine the effects of climate change on the population dynamics of H. armigera. The results showed that all generations of H. armigera at both sites showed increasing trends in population size with climate warming. Abrupt changes in phenology and population number occurred after abrupt temperature changes. Climate change had a greater effect on the phenology of H. armigera at higher latitudes than at lower latitudes and led to a greater increase in population size at lower latitudes than at higher latitudes; the temperature increase at higher latitudes will cause a greater increase in the adult moth population size in the future compared to that at lower latitudes; and abrupt changes in the phenology, temperature increase, and population size at lower latitudes occurred earlier than those at higher latitudes. Thus, it is necessary to develop sustainable management strategies for Helicoverpa armigera at an early stage.     

Heritability of and strong single gene (Pgi) effects on life‐history traits in the Glanville fritillary butterfly

We estimated broad‐sense heritabilities (H²) of 13 female and seven male life‐history traits of the Glanville fritillary butterfly (Melitaea cinxia) under semi‐natural conditions in a large outdoor population cage. The analysis was based on full‐sib families collected as young larvae in the field and reared under common garden conditions. We found significant genetic variance in female lifespan, fecundity, number of matings and host‐plant preference as well as in male body mass and mobility. Apart from host‐plant preference, female traits that were more strongly correlated with lifetime reproductive success (LRS; measured as total number of eggs laid) had higher H². LRS itself exhibited significant heritability. Host‐plant preference had very high H², consistent with a previously reported genetically determined geographical cline in host‐plant preference in the study area. Lifespan and egg hatching rate were significantly associated with a SNP in the coding region of the Pgi gene, for which there is previous evidence for balancing selection. Selection on Pgi, which furthermore shows spatial and temporal variation, may maintain genetic variance in fitness‐related life‐history traits. In contrast, we found no strong evidence for life‐history trade‐offs.     

Genetic architecture for normal and novel host-plant use in two local populations of the herbivorous ladybird beetle, Epilachna pustulosa

Trade-offs in host-plant use are thought to promote the evolution of host specificity. However, usually either positive or no genetic correlations have been found. Whereas factors enhancing variation in overall viability have been claimed to mask negative genetic correlations, alternative hypotheses emphasize the sequential changes in genetic correlation in the course of host-range evolution. In this study, the genetic architectures of performances on different hosts were compared in two populations of the herbivorous ladybird beetle, Epilachna pustulosa, using three host plants, one being normal for both, one novel for only one population, and the other novel for both populations. The genetic correlations between larval periods on normal hosts were significantly positive whereas those between normal and novel hosts were not different from zero. There was no evidence for reduced genetic variation on the normal host-plants. These results suggest that the host-range is not restricted by the antagonistic genetic associations among exploitation abilities on different plant species, but rather that selection of different host-plants may improve the coordination between genes responsible for the use of different plants.     

Genetic diversity analysis and conservation implications for the Iberian threatened populations of the irano-turanian relict Krascheninnikovia ceratoides (Chenopodiaceae)

Krascheninnikovia ceratoides (L.) Gueldenst. (Chenopodiaceae) is an irano‐turanian steppe plant with a striking geographical vicariance between the western Mediterranean and the central Asian regions. Its westernmost populations are located in the north‐east part of the Iberian Peninsula, in the narrow middle Ebro and Alfambra valleys, where they are threatened and have been catalogued as ‘vulnerable’. Genetic diversity and structure was studied in 150 individuals from five populations of these two valleys using inter‐simple sequence repeats (ISSR). A total of 121 informative bands allowed the identification of 150 genotypes. Nei's genetic diversity found in all natural populations (h = 0.448) was higher than that of other geographical restricted species. Analysis of molecular variance and spatial correlations analyses showed a strong genetic differentiation among populations (35.88%) and among valleys (22.33%). The unweighted pair‐group method of arithmetical averages and Neighbour‐joining clusterings further demonstrated a substructure of two population cores within the Ebro valley. Our genetic data suggest that the high levels of genetic diversity and the strong genetic structure found among the northern Iberian populations and ranges of K. ceratoides might result from the summed effects of its tetraploidy, its outbreeding nature, and its success in colonizing newly altered areas. However, the ISSR data might also reflect the existence of an ancient wider distribution range of the plant in the Iberian Peninsula coupled with a more recent history of habitat fragmentation. Different management conservation guidelines are recommended for this plant after our genetic analysis: a microreserve of approximately 2 km² is proposed for the highly threatened population of Osera (Ebro valley), whereas seed collection strategies and occasional population reinforcements are suggested for the remaining populations. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 92 , 419–429.