Combined effects of climate and biotic interactions on the elevational range of a phytophagous insect

1. The ranges of many species have expanded in cool regions but contracted at warm margins in response to recent climate warming, but the mechanisms behind such changes remain unclear. Particular debate concerns the roles of direct climatic limitation vs. the effects of interacting species in explaining the location of low latitude or low elevation range margins. 2. The mountains of the Sierra de Guadarrama (central Spain) include both cool and warm range margins for the black-veined white butterfly, Aporia crataegi, which has disappeared from low elevations since the 1970s without colonizing the highest elevations. 3. We found that the current upper elevation limit to A. crataegi's distribution coincided closely with that of its host plants, but that the species was absent from elevations below 900 m, even where host plants were present. The density of A. crataegi per host plant increased with elevation, but overall abundance of the species declined at high elevations where host plants were rare. 4. The flight period of A. crataegi was later at higher elevations, meaning that butterflies in higher populations flew at hotter times of year; nevertheless, daytime temperatures for the month of peak flight decreased by 6.2 degrees C per 1 km increase in elevation. 5. At higher elevations A. crataegi eggs were laid on the south side of host plants (expected to correspond to hotter microclimates), whereas at lower sites the (cooler) north side of plants was selected. Field transplant experiments showed that egg survival increased with elevation. 6. Climatic limitation is the most likely explanation for the low elevation range margin of A. crataegi, whereas the absence of host plants from high elevations sets the upper limit. This contrasts with the frequent assumption that biotic interactions typically determine warm range margins, and thermal limitation cool margins. 7. Studies that have modelled distribution changes in response to climate change may have underestimated declines for many specialist species, because range contractions will be exacerbated by mismatch between the future distribution of suitable climate space and the availability of resources such as host plants.     

Fish introductions reveal the temperature dependence of species interactions

A major area of current research is to understand how climate change will impact species interactions and ultimately biodiversity. A variety of environmental conditions are rapidly changing owing to climate warming, and these conditions often affect both the strength and outcome of species interactions. We used fish distributions and replicated fish introductions to investigate environmental conditions influencing the coexistence of two fishes in Swedish lakes: brown trout (Salmo trutta) and pike (Esox lucius). A logistic regression model of brown trout and pike coexistence showed that these species coexist in large lakes (more than 4.5 km²), but not in small, warm lakes (annual air temperature more than 0.9–1.5°C). We then explored how climate change will alter coexistence by substituting climate scenarios for 2091–2100 into our model. The model predicts that brown trout will be extirpated from approximately half of the lakes where they presently coexist with pike and from nearly all 9100 lakes where pike are predicted to invade. Context dependency was critical for understanding pike–brown trout interactions, and, given the widespread occurrence of context-dependent species interactions, this aspect will probably be critical for accurately predicting climate impacts on biodiversity.     

The ability of climate envelope models to predict the effect of climate change on species distributions

Climate envelope models (CEMs) have been used to predict the distribution of species under current, past, and future climatic conditions by inferring a species' environmental requirements from localities where it is currently known to occur. CEMs can be evaluated for their ability to predict current species distributions but it is unclear whether models that are successful in predicting current distributions are equally successful in predicting distributions under different climates (i.e. different regions or time periods). We evaluated the ability of CEMs to predict species distributions under different climates by comparing their predictions with those obtained with a mechanistic model (MM). In an MM the distribution of a species is modeled based on knowledge of a species' physiology. The potential distributions of 100 plant species were modeled with an MM for current conditions, a past climate reconstruction (21 000 years before present) and a future climate projection (double preindustrial CO₂ conditions). Point localities extracted from the currently suitable area according to the MM were used to predict current, future, and past distributions with four CEMs covering a broad range of statistical approaches: Bioclim (percentile distributions), Domain (distance metric), GAM (general additive modeling), and Maxent (maximum entropy). Domain performed very poorly, strongly underestimating range sizes for past or future conditions. Maxent and GAM performed as well under current climates as under past and future climates. Bioclim slightly underestimated range sizes but the predicted ranges overlapped more with the ranges predicted with the MM than those predicted with GAM did. Ranges predicted with Maxent overlapped most with those produced with the MMs, but compared with the ranges predicted with GAM they were more variable and sometimes much too large. Our results suggest that some CEMs can indeed be used to predict species distributions under climate change, but individual modeling approaches should be validated for this purpose, and model choice could be made dependent on the purpose of a particular study.     

Predicting species distribution and abundance responses to climate change: why it is essential to include biotic interactions across trophic levels

Current predictions on species responses to climate change strongly rely on projecting altered environmental conditions on species distributions. However, it is increasingly acknowledged that climate change also influences species interactions. We review and synthesize literature information on biotic interactions and use it to argue that the abundance of species and the direction of selection during climate change vary depending on how their trophic interactions become disrupted. Plant abundance can be controlled by aboveground and belowground multitrophic level interactions with herbivores, pathogens, symbionts and their enemies. We discuss how these interactions may alter during climate change and the resulting species range shifts. We suggest conceptual analogies between species responses to climate warming and exotic species introduced in new ranges. There are also important differences: the herbivores, pathogens and mutualistic symbionts of range-expanding species and their enemies may co-migrate, and the continuous gene flow under climate warming can make adaptation in the expansion zone of range expanders different from that of cross-continental exotic species. We conclude that under climate change, results of altered species interactions may vary, ranging from species becoming rare to disproportionately abundant. Taking these possibilities into account will provide a new perspective on predicting species distribution under climate change.     

Modelling species' range shifts in a changing climate: the impacts of biotic interactions, dispersal distance and the rate of climate change

There is an urgent need for accurate prediction of climate change impacts on species ranges. Current reliance on bioclimatic envelope approaches ignores important biological processes such as interactions and dispersal. Although much debated, it is unclear how such processes might influence range shifting. Using individual-based modelling we show that interspecific interactions and dispersal ability interact with the rate of climate change to determine range-shifting dynamics in a simulated community with two growth forms--mutualists and competitors. Interactions determine spatial arrangements of species prior to the onset of rapid climate change. These lead to space-occupancy effects that limit the rate of expansion of the fast-growing competitors but which can be overcome by increased long-distance dispersal. As the rate of climate change increases, lower levels of long-distance dispersal can drive the mutualists to extinction, demonstrating the potential for subtle process balances, non-linear dynamics and abrupt changes from species coexistence to species loss during climate change.     

Predicting the impacts of climate change on the distribution of species: are bioclimate envelope models useful?

Modelling strategies for predicting the potential impacts of climate change on the natural distribution of species have often focused on the characterization of a species’ bioclimate envelope. A number of recent critiques have questioned the validity of this approach by pointing to the many factors other than climate that play an important part in determining species distributions and the dynamics of distribution changes. Such factors include biotic interactions, evolutionary change and dispersal ability. This paper reviews and evaluates criticisms of bioclimate envelope models and discusses the implications of these criticisms for the different modelling strategies employed. It is proposed that, although the complexity of the natural system presents fundamental limits to predictive modelling, the bioclimate envelope approach can provide a useful first approximation as to the potentially dramatic impact of climate change on biodiversity. However, it is stressed that the spatial scale at which these models are applied is of fundamental importance, and that model results should not be interpreted without due consideration of the limitations involved. A hierarchical modelling framework is proposed through which some of these limitations can be addressed within a broader, scale‐dependent context.     

The legacy of past climate and landscape change on species' current experienced climate and elevation ranges across latitude: a multispecies study utilizing mammals in western North America

Aim Elevation and climate ranges across latitude experienced by 21 wide‐ranging mammal species in western North America were summarized to examine two questions: (1) do populations in the northern and southern portions of a species’ range experience different climates or are environments selected to remain similar to climates at the core of ranges; and (2) how do species’ elevational ranges, experienced temperature seasonality and temperature ranges change across latitude? Given the larger effects of climate oscillations in the north vs. the south, a predicted outcome is for species to conserve climate niches across latitude and to show reduced climate and elevation ranges in the north. An alternative outcome is latitudinal niche diversification and increased climate variation in the north. Location Western North America. Methods The questions above were examined using a combination of species occurrence data bases, climate data bases, simple summaries of means and standard deviations and by testing summaries against random distributions across latitude for 21 mammal species from a variety of orders. Results The results showed that: (i) most species conserve their niche strongly or weakly given overall temperature gradients from north to south; (ii) seasonality experienced by species is relatively static until the highest latitudes despite directional trends across the region; and (iii) the elevation range and temperature variation that species experience decreases from south to north. Main conclusions Populations at range edges appear to partition environments to remain closer to temperature values similar to those at the core of the range. In addition, seasonality is not a likely explanatory factor of genetic diversity in latitudinal gradients. The data are instead more consistent with predictions that a combination of higher gene‐flow, increasing environmental instability and decreasing elevation gradients in the north compared to the south may lead to negative correlations between latitude and species’ climate variation. The results corroborate risks faced by northern mammal populations to global climate changes.     

Changes in species interactions across a 2.5 km elevation gradient: effects on plant migration in response to climate change

Predicted climate change in the Andes will require plant species to migrate upslope to avoid extinction. Central to predictions of species responses to climate change is an understanding of species distributions along environmental gradients. Environmental gradients are frequently modelled as abiotic, but biotic interactions can play important roles in setting species distributions, abundances, and life history traits. Biotic interactions also have the potential to influence species responses to climate change, yet they remain mostly unquantified. An important interaction long studied in tropical forests is postdispersal seed predation which has been shown to affect the population dynamics, community structure, and diversity of plant species in time and space. This paper presents a comparative seed predation study of 24 species of tropical trees across a 2.5 km elevation gradient in the Peruvian Andes and quantifies seed predation variation across the elevational gradient. We then use demographic modelling to assess effects of the observed variation in seed predation on population growth rates in response to observed increasing temperatures in the area. We found marked variation among species in total seed predation depending on the major seed predator of the species and consistent changes in seed predation across the gradient. There was a significant increase in seed survival with increasing elevation, a trend that appears to be driven by regulation of seed predators via top–down forces in the lowlands giving way to bottom–up (productivity) regulation at mid‐ to high elevations, resulting in a ninefold increase in effective fecundity for trees at high elevations. This potential increase in seed crop size strongly affects modelled plant population growth and seed dispersal distances, increasing population migration potential in the face of climate change. These results also indicate that species interactions can have effects on par with climate in species responses to global change.     

The data void in modeling current and future distributions of tropical species

Conserving biodiversity in the face of climate change requires a predictive ecology of species distributions. Nowhere is this need more acute than in the tropics, which harbor the majority of Earth's species and face rapid and large climate and land‐use changes. However, the study of species distributions and their responses to climate change in high diversity tropical regions is potentially crippled by a lack of basic data. We analyzed a database representing more than 800 000 unique geo‐referenced natural history collections to determine what fraction of tropical plant species has sufficient numbers of available collections for use in the habitat or niche models commonly used to predict species responses to climate change. We found that more than nine out of 10 species from the three principle tropical realms are so poorly collected (n < 20 records) that they are essentially invisible to modern modeling and conservation tools. In order to predict the impact of climate change on tropical species, efforts must be made to increase the amount of data available from tropical countries through a combination of collecting new specimens and digitizing existing records.     

Disentangling the relative importance of climate,size and competition on tree growth in Iberian forests: implications for forest management under global change

Most large‐scale multispecies studies of tree growth have been conducted in tropical and cool temperate forests, whereas Mediterranean water‐limited ecosystems have received much less attention. This limits our understanding of how growth of coexisting tree species varies along environmental gradients in these forests, and the implications for species interactions and community assembly under current and future climatic conditions. Here, we quantify the absolute effect and relative importance of climate, tree size and competition as determinants of tree growth patterns in Iberian forests, and explore interspecific differences in the two components of competitive ability (competitive response and effect) along climatic and size gradients. Spatially explicit neighborhood models were developed to predict tree growth for the 15 most abundant Iberian tree species using permanent‐plot data from the Spanish Second and Third National Forest Inventory (IFN). Our neighborhood analyses showed a climatic and size effect on tree growth, but also revealed that competition from neighbors has a comparatively much larger impact on growth in Iberian forests. Moreover, the sensitivity to competition (i.e. competitive response) of target trees varied markedly along climatic gradients causing significant rank reversals in species performance, particularly under xeric conditions. We also found compelling evidence for strong species‐specific competitive effects in these forests. Altogether, these results constitute critical new information which not only furthers our understanding of important theoretical questions about the assembly of Mediterranean forests, but will also be of help in developing new guidelines for adapting forests in this climatic boundary to global change. If we consider the climatic gradients of this study as a surrogate for future climatic conditions, then we should expect absolute growth rates to decrease and sensitivity to competition to increase in most forests of the Iberian Peninsula (in all but the northern Atlantic forests), making these management considerations even more important in the future.