Insect responses to heat: physiological mechanisms,evolution and ecological implications in a warming world

Surviving changing climate conditions is particularly difficult for organisms such as insects that depend on environmental temperature to regulate their physiological functions. Insects are extremely threatened by global warming, since many do not have enough physiological tolerance even to survive continuous exposure to the current maximum temperatures experienced in their habitats. Here, we review literature on the physiological mechanisms that regulate responses to heat and provide heat tolerance in insects: (i) neuronal mechanisms to detect and respond to heat; (ii) metabolic responses to heat; (iii) thermoregulation; (iv) stress responses to tolerate heat; and (v) hormones that coordinate developmental and behavioural responses at warm temperatures. Our review shows that, apart from the stress response mediated by heat shock proteins, the physiological mechanisms of heat tolerance in insects remain poorly studied. Based on life‐history theory, we discuss the costs of heat tolerance and the potential evolutionary mechanisms driving insect adaptations to high temperatures. Some insects may deal with ongoing global warming by the joint action of phenotypic plasticity and genetic adaptation. Plastic responses are limited and may not be by themselves enough to withstand ongoing warming trends. Although the evidence is still scarce and deserves further research in different insect taxa, genetic adaptation to high temperatures may result from rapid evolution. Finally, we emphasize the importance of incorporating physiological information for modelling species distributions and ecological interactions under global warming scenarios. This review identifies several open questions to improve our understanding of how insects respond physiologically to heat and the evolutionary and ecological consequences of those responses. Further lines of research are suggested at the species, order and class levels, with experimental and analytical approaches such as artificial selection, quantitative genetics and comparative analyses.     

Detecting climate change induced range shifts: Where and how should we be looking?

Abstract: Global climate warming is expected to cause systematic shifts in the distribution of species and consequently increase extinction risk. Conservation managers must be able to detect, measure and accurately predict range shifts in order to mitigate impacts on biodiversity. However, important responses to climate change may go unnoticed or be dismissed if we fail to collect sufficient baseline data and apply the most sensitive analytical tests. Here we use randomizations of a contemporary data set on rainforest birds of north‐eastern Australia to quantify the sensitivity of three measures for assessing range shifts along altitudinal gradients. We find that smaller range shifts are detectable by analysing change in the mean altitude of presence records rather than upper or lower range boundaries. For a moderate survey effort of 96 surveys, measurements of change in the mean altitude of 34 species have the capacity to provide strong inference for a mean altitudinal range shift as small as 40 m across the species assemblage. We also show that range shifts measured at range boundaries can be potentially misleading when differences in sampling effort between contemporary and historical data sets are not taken into account.     

Potential impacts of climate change on stable flies, investigated along an altitudinal gradient

Adult populations of stable flies were sampled along an altitudinal transect in Reunion Island to determine whether higher temperatures were associated with: (a) higher numbers of flies; (b) a longer season of infestation, and/or (c) different responses to warming in the cosmopolitan Stomoxys calcitrans (L) and the tropical Stomoxys niger niger Macquart (Diptera: Muscidae). Flies of both species were trapped at seven farms situated at four altitudes (100-1600 m a.s.l.) over a 90-week period. For both species, there were no relationships between the maximum or mean fly abundance and altitude. Only minimum abundance in winter was significantly higher at lower altitudes. Maximum and mean abundances differed significantly between nearby farms under similar climatic conditions. Seasonal fluctuations in fly abundance changed along the gradient. At lower altitudes, population growth started earlier after the winter but abundance declined earlier in summer, which resulted in a shift of the season of infestation. Seasonal fluctuations of both species were strongly related to climate variables at high altitude, mainly temperature. However, climate variables explained a decreasing proportion of the variations in abundance at lower altitudes. Stomoxys calcitrans was the most abundant species overall, but the proportion of S. n. niger increased significantly at lower altitudes and this species became predominant at 100 m a.s.l. It is concluded that stable fly infestations are unlikely to worsen in response to global warming. Maximum abundance is limited by local factors, possibly larval resources, which suggests that adequate husbandry practices could override the impact of climate change. Because S. n. niger tends to be the predominant pest at elevated temperatures, it is recommended that this species should not be introduced in areas where climate is changing.     

全球气候变暖对“植物-害虫-天敌”互作系统的影响

全球气候变化是近来人类关注的焦点问题,其最显著的特征是气候变暖。因为昆虫具有生活周期短、繁殖率高等特点,所以,气候变暖对昆虫的发育、繁殖和存活会产生强烈的直接影响。气候变暖促使一些昆虫提前春天的物候现象,向高纬度或高海拔地区迁移。然而,昆虫在自然界并非孤立地存在,它们与寄主植物和自然天敌相互联系、相互作用,并在长期的进化过程中逐渐适应特定区域的气候条件。因此,全球气候变暖对"植物-害虫-天敌"的种间关系必然产生直接或间接的影响,导致不同昆虫之间以及昆虫与其相关营养层的物种之间的相互关系在气候变化下呈现出时间上的异步性和空间上的错位,从而影响植物的适应性和抗虫性、害虫的发生规律和危害程度以及天敌的种群消长和控害效能。昆虫除了可以通过休眠或滞育的方式在时间上避开高温的影响外,还可以通过迁飞或移动的方式在空间上避开高温的影响,在这种迁移和扩散不同步的情况下可能使害虫食性和取食植物的害虫及其天敌的种类发生变化,从而改变生物群落的组成与结构,影响生态系统的服务和功能。     

Plants,Birds and Butterflies: Short-Term Responses of Species Communities to Climate Warming Vary by Taxon and with Altitude

As a consequence of climate warming, species usually shift their distribution towards higher latitudes or altitudes. Yet, it is unclear how different taxonomic groups may respond to climate warming over larger altitudinal ranges. Here, we used data from the national biodiversity monitoring program of Switzerland, collected over an altitudinal range of 2500 m. Within the short period of eight years (2003–2010), we found significant shifts in communities of vascular plants, butterflies and birds. At low altitudes, communities of all species groups changed towards warm-dwelling species, corresponding to an average uphill shift of 8 m, 38 m and 42 m in plant, butterfly and bird communities, respectively. However, rates of community changes decreased with altitude in plants and butterflies, while bird communities changed towards warm-dwelling species at all altitudes. We found no decrease in community variation with respect to temperature niches of species, suggesting that climate warming has not led to more homogenous communities. The different community changes depending on altitude could not be explained by different changes of air temperatures, since during the 16 years between 1995 and 2010, summer temperatures in Switzerland rose by about 0.07°C per year at all altitudes. We discuss that land-use changes or increased disturbances may have prevented alpine plant and butterfly communities from changing towards warm-dwelling species. However, the findings are also consistent with the hypothesis that unlike birds, many alpine plant species in a warming climate could find suitable habitats within just a few metres, due to the highly varied surface of alpine landscapes. Our results may thus support the idea that for plants and butterflies and on a short temporal scale, alpine landscapes are safer places than lowlands in a warming world.     

Predicting potential responses to future climate in an alpine ungulate: interspecific interactions exceed climate effects

The altitudinal shifts of many montane populations are lagging behind climate change. Understanding habitual, daily behavioural rhythms, and their climatic and environmental influences, could shed light on the constraints on long‐term upslope range‐shifts. In addition, behavioural rhythms can be affected by interspecific interactions, which can ameliorate or exacerbate climate‐driven effects on ecology. Here, we investigate the relative influences of ambient temperature and an interaction with domestic sheep (Ovis aries) on the altitude use and activity budgets of a mountain ungulate, the Alpine chamois (Rupicapra rupicapra). Chamois moved upslope when it was hotter but this effect was modest compared to that of the presence of sheep, to which they reacted by moving 89–103 m upslope, into an entirely novel altitudinal range. Across the European Alps, a range‐shift of this magnitude corresponds to a 46% decrease in the availability of suitable foraging habitat. This highlights the importance of understanding how factors such as competition and disturbance shape a given species’ realised niche when predicting potential future responses to change. Furthermore, it exposes the potential for manipulations of species interactions to ameliorate the impacts of climate change, in this case by the careful management of livestock. Such manipulations could be particularly appropriate for species where competition or disturbance already strongly restricts their available niche. Our results also reveal the potential role of behavioural flexibility in responses to climate change. Chamois reduced their activity when it was warmer, which could explain their modest altitudinal migrations. Considering this behavioural flexibility, our model predicts a small 15–30 m upslope shift by 2100 in response to climate change, less than 4% of the altitudinal shift that would be predicted using a traditional species distribution model‐type approach (SDM), which assumes that species’ behaviour remains unchanged as climate changes. Behavioural modifications could strongly affect how species respond to a changing climate.     

气候变暖对昆虫影响研究进展

"全球气候变化"已成为国内外最受关注的环境问题。气候变化中以温度升高为特征的气候变暖对变温动物昆虫自身及其所在的生物群落产生直接或间接影响。从研究内容与研究方法2个方面综述了气候变暖对昆虫影响研究的国内外进展。气候变暖导致昆虫发生期提前,地理分布向更高纬度和海拔地区扩散,低温适生种种群密度下降,高温适生种种群密度增加。气候变暖改变寄主植物—害虫—天敌的物候同步性和昆虫原有种间互作关系,影响植食性昆虫的寄主植物范围和取食为害程度。长期的气候变暖带来的强烈的选择性压力引起某些昆虫种群的基因组发生变异。以日均温升高、日最高气温升高和昼夜温差变化等为主要特征气候变暖对昆虫发育、繁殖及存活等生态学指标产生重要影响。研究方法上主要是利用野外直接观察法、回归预测模型、有效积温模型、CLIMEX和GIS等生态风险评估软件、生物化石比较技术、人工气候下生态试验、检测标记基因频率变化等方法来研究气候变暖对昆虫的影响。最后简要评述了已有研究的不足并指出未来的研究方向:(1)气候变暖情景下开展昆虫种间互作研究并拓展研究对象;(2)高温下昆虫适应性研究;(3)建立完善人工模拟气候下的实验方法;(4)构建昆虫有效生态机理模型。     

Water loss in insects: an environmental change perspective

In the context of global environmental change much of the focus has been on changing temperatures. However, patterns of rainfall and water availability have also been changing and are expected to continue doing so. In consequence, understanding the responses of insects to water availability is important, especially because it has a pronounced influence on insect activity, distribution patterns, and species richness. Here we therefore provide a critical review of key questions that either are being or need to be addressed in this field. First, an overview of insect behavioural responses to changing humidity conditions and the mechanisms underlying sensing of humidity variation is provided. The primary sensors in insects belong to the temperature receptor protein superfamily of cation channels. Temperature-activated transient receptor potential ion channels, or thermoTRPs, respond to a diverse range of stimuli and may be a primary integrator of sensory information, such as environmental temperature and moisture. Next we touch briefly on the components of water loss, drawing attention to a new, universal model of the water costs of gas exchange and its implications for responses to a warming, and in places drying, world. We also provide an overview of new understanding of the role of the sub-elytral chamber for water conservation, and developments in understanding of the role of cuticular hydrocarbons in preventing water loss. Because of an increasing focus on the molecular basis of responses to dehydration stress we touch briefly on this area, drawing attention to the role of sugars, heat shock proteins, aquaporins, and LEA proteins. Next we consider phenotypic plasticity or acclimation responses in insect water balance after initial exposures to altered humidity, temperature or nutrition. Although beneficial acclimation has been demonstrated in several instances, this is not always the case. Laboratory studies show that responses to selection for enhanced ability to survive water stress do evolve and that genetic variation for traits underlying such responses does exist in many species. However, in others, especially tropical, typically narrowly distributed species, this appears not to be the case. Using the above information we then demonstrate that habitat alteration, climate change, biological invasions, pollution and overexploitation are likely to be having considerable effects on insect populations mediated through physiological responses (or the lack thereof) to water stress, and that these effects may often be non-intuitive.     

Responses of insects to the current climate changes: from physiology and behavior to range shifts

Climate change (first of all the rise in temperature) is currently considered one of the most serious global challenges facing mankind. Here we review the diversity of insect responses to the current climate warming, with particular focus on true bugs (Heteroptera). Insects as ectotherms are bound to respond to the temperature change, and different species respond differently depending on their specific physiological and ecological traits, seasonal cycle, trophic relations, etc. Insect responses to climate warming can be divided into six categories: changes in (1) ranges, (2) abundance, (3) phenology, (4) voltinism, (5) morphology, physiology, and behavior, and (6) relationships with other species and in the structure of communities. Changes in ranges and phenology are easier to notice and record than other responses. Range shifts have been reported more often in Lepidoptera and Odonata than in other insect orders. We briefly outline the history and eco-physiological background of the recent range limit changes in the Southern green stink bug Nezara viridula (Heteroptera, Pentatomidae) in central Japan. Range expansion in individual species can lead to enrichment of local faunas, especially at high latitudes. Phenological changes include not only advances in development in spring but also shifts in phenology later in the season. The phenophases related to the end of activity usually shift to later dates, thus prolonging the period of active development. This may have both positive and negative consequences for the species and populations. As with any other response, the tendencies in phenological changes may vary among species and climatic zones. The proven cases of change in voltinism are rare, but such examples do exist. Application of models based on thermal parameters of development suggests that a rise in temperature by 2°C will result in an increased number of annual generations in many species from different arthropod taxa (up to three or four additional generations in Thysanoptera, Aphidoidea, and Acarina). The warming-mediated changes in physiology, morphology, or behavior are difficult to detect and prove, first of all because of the absence of reliable comparative data. Nevertheless, there are examples of changes in photoperiodic responses of diapause induction and behavioral responses related to search of shelters for summer diapause (aestivation). Since (1) individual species do not exist in isolation and (2) the direction and magnitude of responses even to the same environmental changes vary between species, it may be expected that in many cases the current stable relationships between species will be affected. Thus, unequal range shifts in insects and their host plants may disrupt their trophic interactions near the species?? range boundaries. Studies of responses to climate warming in more than one interrelated species or in entire communities are extremely rare. The loss of synchronism in seasonal development of community members may indicate inability of the higher trophic levels to adapt fully to climate warming or an attempt of the lower trophic level to escape from the pressure of the higher trophic levels. It is generally supposed that many insect species in the Temperate Climate Zone will benefit in some way from the current climate warming. However, there is some experimental evidence of an opposite or at least much more complex response; the influence of warming might be deleterious for some species or populations. It is suggested that species or populations from the cold or temperate climate have sufficient phenotypic plasticity to survive under the conditions of climate warming, whereas species and populations which already suffer from stress under extreme seasonal temperatures in warmer regions may have a limited ??maneuver space?? since the current temperatures are close to their upper thermal limits. Without genetic changes, even moderate warming will put these species or populations under serious physiological stress. The accumulated data suggest that responses of insects and the entire biota to climate warming will be complex and will vary depending on the rate of warming and ecological peculiarities of species and regions. Physiological responses will vary in their nature, direction, and magnitude even within one species or population, and especially between seasons. The responses will also differ in different seasons. For example, warming may negatively affect nymphal development during the hot season but at the same time accelerate growth and development during the cold season and/or ensure milder and more favorable overwintering conditions for adults. All these factors will affect population dynamics of particular species and relationships among the members of ecosystems. We should keep in mind that (1) not only selected insect species but almost all the species will be affected, (2) temperature is not the only component of the climatic system that is changing, and (3) responses will be different in different seasons. Host plants, phytophagous insects, their competitors, symbionts, predators, parasites, and pathogens will not only respond separately to climate changes; individual responses will further affect the responses of other species, thus making reliable prediction extremely complicated. Responses are expected to (1) be species- or population-specific, (2) concern basically all the aspects of organism/ species biology and ecology (individual physiology, population structure, abundance, local adaptations, phenology, voltinism, and distribution), and (3) occur at scales ranging from an undetectable cellular level to major distribution range shifts or regional extinctions. The scale of insect responses will depend on the extent and rate of climate warming. Slight to moderate warming may cause responses only in a limited number of species with more flexible life cycles, whereas a substantial increase in temperature may affect a greater number of different species and ecological groups.     

卧龙竹类非结构性碳水化合物与叶氮含量对海拔的响应

山地由于海拔变化导致的环境因子显著差异,成为研究植物环境适应性及其对全球气候变化响应的理想区域。以卧龙自然保护区内油竹子(Fargesia angustissima Yi)与华西箭竹(Fargesia nitida(Mitford)Keng f.ex Yi)为对象,沿海拔梯度研究了两种竹子在各自海拔分布区间内组织非结构性碳水化合物(NSC)含量、比叶面积(SLA)、以及基于单位叶面积和单位叶质量的叶氮含量(Narea,Nmass)。除油竹子叶NSC,Narea和华西箭竹Nmass随海拔升高不发生变化外,两种竹子其它调查因子对海拔的响应均是非线性的,反映了环境因子随海拔的非线性变化。所有调查因子对海拔的响应均表现出明显的种间差别,这一结果除了种间生理生态特性差别的原因外,可能意味着两竹种对温度的敏感性不同。高海拔种(华西箭竹)比(中)低海拔种(油竹子)对全球气候暖化可能更加敏感。     

Mutualism meltdown in insects: bacteria constrain thermal adaptation

Predicting whether and how organisms will successfully cope with climate change presents critical questions for biologists and environmental scientists. Models require knowing how organisms interact with their abiotic environment, as well understanding biotic interactions that include a network of symbioses in which all species are embedded. Bacterial symbionts of insects offer valuable models to examine how microbes can facilitate and constrain adaptation to a changing environment. While some symbionts confer plasticity that accelerates adaptation, long-term bacterial mutualists of insects are characterized by tight lifestyle constraints, genome deterioration, and vulnerability to thermal stress. These essential bacterial partners are eliminated at high temperatures, analogous to the loss of zooanthellae during coral bleaching. Recent field-based studies suggest that thermal sensitivity of bacterial mutualists constrains insect responses. In this sense, highly dependent mutualisms may be the Achilles' heel of thermal responses in insects.     

An elevational shift of cryophilous bryophytes in the last century – an effect of climate warming?

Aim To investigate altitudinal range shifts of bryophytes in Switzerland by comparing recent altitudinal distributions with historical distributions derived from herbarium specimens. Location Switzerland, covering 41,285 km² in Central Europe. Methods We used a dataset of 8520 herbarium specimens of 61 bryophyte species and compared altitudinal data between the two periods 1880–1920 and 1980–2005. The records we used were not specifically sampled for climatological analyses, but originate from non‐systematic fieldwork by various collectors. Historical and recent records were distributed all over Switzerland with occurrences in all major biogeographical areas. To account for different sampling efforts in the two time periods, different subsampling procedures were applied. Results Overall, we found a significant mean increase in altitude of 89 ± 29 m which was mainly driven by the cryophilous species (+222 ± 50 m). The mean increase in altitude of cryophilous species corresponds to a decadal upward shift of 24 m. The upper range limit of cryophilous species also increased by 189 ± 55 m, but there was no effect on the lower range limit. For intermediate and thermophilous species neither mean, nor upper or lower range limits changed. However, the proportion of records of thermophilous to cryophilous species increased considerably at lower altitudes, but levelled off above approximately 1800 m. Main conclusions We conclude that cryophilous bryophytes are expanding their range to higher elevations in Switzerland and that at lower elevations, a slow extinction process is going on, probably as a result of climate warming trends. The observed decadal upward shifts of cryophilous species closely match those reported from vascular plants in Europe and those expected, given recent estimates of climate warming trends. We emphasize that herbaria provide valuable data that can be used to detect ongoing changes in the distribution of species.     

Small shifts in microsite occupation could mitigate climate change consequences for mountain top endemics: a test analyzing saxicolous lichen distribution patterns

The extent to which small shifts among local topographic microsites could mitigate the effects of larger-scale climate change in arctic–alpine systems including mountain top organisms is largely unknown. This study is among the first to evaluate the relative contribution of microsite and altitude as a proxy for climate change on saxicolous lichen communities. We registered 107 lichen species in 54 boulders ranging from 900 to 2700 m.a.s.l. and in a large array of microsites in central Argentina. Communities ordinated along NMS multivariate analysis axes 1, 2 and 3 presented a cumulative R² of 80%. The three axes were explained by altitude with axis 1 only being explained by altitude. Axis 2 was also explained by slope and aspect whereas axis 3 was explained by the interaction of altitude with aspect indicating that aspect was important only at lower altitudes but not at the mountain top. Lichen cover and richness were similar throughout the altitudinal gradient. We interpret that under a climate warming scenario, lower altitude species occupying pole ward facing slopes will have to migrate upwards while at the mountain top—for most communities—there still is scope for microsite segregation to compensate climate change.     

Early effects of climate change: do they include changes in vector-borne disease?

The world's climate appears now to be changing at an unprecedented rate. Shifts in the distribution and behaviour of insect and bird species indicate that biological systems are already responding to this change. It is well established that climate is an important determinant of the spatial and temporal distribution of vectors and pathogens. In theory, a change in climate would be expected to cause changes in the geographical range, seasonality (intra-annual variability), and in the incidence rate (with or without changes in geographical or seasonal patterns). The detection and then attribution of such changes to climate change is an emerging task for scientists. We discuss the evidence required to attribute changes in disease and vectors to the early effects of anthropogenic climate change. The literature to date indicates that there is a lack of strong evidence of the impact of climate change on vector-borne diseases (i.e. malaria, dengue, leishmaniasis, tick-borne diseases). New approaches to monitoring, such as frequent and long-term sampling along transects to monitor the full latitudinal and altitudinal range of specific vector species, are necessary in order to provide convincing direct evidence of climate change effects. There is a need to reassess the appropriate levels of evidence, including dealing with the uncertainties attached to detecting the health impacts of global change.     

寒温带森林白桦径向生长的海拔差异及其气候响应——以奥克里堆山为例

气候变化深刻影响森林生态系统的结构和功能。在全球升温背景下,不同环境中不同树种的生长模式及其气候响应决定着生态系统的发展和稳定。本研究基于大兴安岭地区奥克里堆山的白桦年轮宽度数据,采用树木年代学方法,分析(兴安)落叶松森林中先锋树种白桦的生长-气候响应与升温和海拔变化的关联。结果表明: 气候变暖使具有明显海拔环境差异的白桦产生生长分异。较低海拔(1050 m)区域的白桦生长显著增加,而在相对偏高海拔(1250 m)区域的生长变化不大;在具有明显快速升温变化(1980年)之前,各海拔区域白桦生长的冬季(上年10月至当年2月)低温胁迫均达到显著水平;在快速升温阶段(1981—2010年),白桦生长的冬季低温胁迫降低,生长季(5—7月)温度成为偏高海拔处白桦生长的主要限定性因子;在水热条件较为调和的低海拔区域,白桦生长加快。研究区内白桦的分布总体上随着气候的持续变暖逐渐向高海拔区域扩散。     

昆虫分布南界的形成原因及影响因素

生物都有一定的适生区,低温导致其分布北界的形成是不争的事实,然而一些种类还存在分布上的南界。昆虫的生长发育与环境的关系非常密切,研究其分布南界的形成原因对预测昆虫种群的扩散和暴发具有重要意义。从环境、寄主和人类活动等方面总结了北半球昆虫分布南界形成的机制及影响因素。低纬度地区夏季高温引起昆虫死亡率上升、生殖率下降、昆虫体内共生菌解体和冬季低温过高导致昆虫滞育节律被破坏等是昆虫分布南界形成的主要原因。此外,寄主的分布和人类活动也对昆虫分布南界存在一定影响。气候变暖对不同种类昆虫的分布南界影响不同,大部分昆虫的分布范围会扩大,但一些受环境限制的昆虫分布南界往往北移。建议今后对全球气候变暖背景下昆虫分布南界的变化、检疫性害虫在其原产地和入侵地分布南界的差异及其原因进行研究,以期为分析昆虫分布的变化提供依据,也为害虫防治提供新的可能途径。     

Multi‐scale responses to warming in an experimental insect metacommunity

In metacommunities, diversity is the product of species interactions at the local scale and dispersal between habitat patches at the regional scale. Although warming can alter both species interactions and dispersal, the combined effects of warming on these two processes remains uncertain. To determine the independent and interactive effects of warming‐induced changes to local species interactions and dispersal, we constructed experimental metacommunities consisting of enclosed milkweed patches seeded with five herbivorous milkweed specialist insect species. We treated metacommunities with two levels of warming (unwarmed and warmed) and three levels of connectivity (isolated, low connectivity, high connectivity). Based on metabolic theory, we predicted that if plant resources were limited, warming would accelerate resource drawdown, causing local insect declines and increasing both insect dispersal and the importance of connectivity to neighboring patches for insect persistence. Conversely, given abundant resources, warming could have positive local effects on insects, and the risk of traversing a corridor to reach a neighboring patch could outweigh the benefits of additional resources. We found support for the latter scenario. Neither resource drawdown nor the weak insect‐insect associations in our system were affected by warming, and most insect species did better locally in warmed conditions and had dispersal responses that were unchanged or indirectly affected by warming. Dispersal across the matrix posed a species‐specific risk that led to declines in two species in connected metacommunities. Combined, this scaled up to cause an interactive effect of warming and connectivity on diversity, with unwarmed metacommunities with low connectivity incurring the most rapid declines in diversity. Overall, this study demonstrates the importance of integrating the complex outcomes of species interactions and spatial structure in understanding community response to climate change.     

Response of insect parasitism to elevation depends on host and parasitoid life-history strategies

How global warming will affect insect parasitoids and their role as natural enemies of insect pests is difficult to assess within a short period of time. Considering that elevation gradients can be used as analogues for global warming, we carried out meta-analyses of 27 correlations between parasitoid richness and elevation and 140 correlations between parasitism rate and elevation in natural and semi-natural environments. We also explored various covariates that may explain the observed responses. Both parasitism rates and parasitoid species richness significantly decreased with increasing elevation. The decrease was greater for ectoparasitoids and parasitoids of ectophagous insects than for endoparasitoids and parasitoids of endophagous hosts, possibly because these latter are better protected from adverse and extreme climatic conditions occurring at higher elevations. Although our results suggest an increase of parasitism with increasing temperature, other factors regulating herbivorous insects have to be considered before concluding that climate warming will lead to a decrease in pest density.     

Alpine bird distributions along elevation gradients: the consistency of climate and habitat effects across geographic regions

Many species have shown recent shifts in their distributions in response to climate change. Patterns in species occurrence or abundance along altitudinal gradients often serve as the basis for detecting such changes and assessing future sensitivity. Quantifying the distribution of species along altitudinal gradients acts as a fundamental basis for future studies on environmental change impacts, but in order for models of altitudinal distribution to have wide applicability, it is necessary to know the extent to which altitudinal trends in occurrence are consistent across geographically separated areas. This was assessed by fitting models of bird species occurrence across altitudinal gradients in relation to habitat and climate variables in two geographically separated alpine regions, Piedmont and Trentino. The ten species studied showed non-random altitudinal distributions which in most cases were consistent across regions in terms of pattern. Trends in relation to altitude and differences between regions could be explained mostly by habitat or a combination of habitat and climate variables. Variation partitioning showed that most variation explained by the models was attributable to habitat, or habitat and climate together, rather than climate alone or geographic region. The shape and position of the altitudinal distribution curve is important as it can be related to vulnerability where the available space is limited, i.e. where mountains are not of sufficient altitude for expansion. This study therefore suggests that incorporating habitat and climate variables should be sufficient to construct models with high transferability for many alpine species.     

中国地卷属地衣海拔分布分析

海拔是地衣多样性的重要影响因素。了解地衣的海拔分布格局是地衣多样性保护的基础。研究表明中国地卷属地衣的物种丰富度和个体丰富度沿海拔梯度呈单峰曲线,它们倾向于分布在受全球变暖影响较高的高海拔地区[（2,022±995）m],且具有较窄的海拔分布幅（约68%的物种分布于海拔为1,694－2,954m的地带）,因而对其进行保护生物学的研究已十分迫切。地卷属11种地衣具有不同的海拔分布规律,这些分布规律与各自分布的海拔区间有关。基于11物种的海拔分析支持Rapoport法则。全球暖化对4种地衣的潜在威胁尤其强烈,即：大陆地卷Peltigera continentalis、长孢地卷P. dolichospora、克氏地卷P. kristinssonii和小地卷P. venosa。