Expressed sequence tag‐derived microsatellites for the cool‐water marine copepod Calanus finmarchicus

The copepod Calanus finmarchicus is the major contributor to zooplankton biomass in the North Atlantic and Norwegian Sea, but recent studies have shown a 70% decrease in abundance as well as a northward shift in the species’ range. Insights into dispersal capabilities gained from population genetic studies will be crucial in predicting the response of C. finmarchicus communities to climate change and, consequently, we have developed a set of expressed sequence tag‐derived microsatellite markers to allow fine‐scale elucidation of population structuring and dispersal. Ten polymorphic markers displayed between two and 19 alleles, with levels of expected heterozygosity ranging from 0.044 to 0.924.     

Post-bloom feeding of Calanus finmarchicus copepodites: Selection for autotrophic versus heterotrophic prey

Feeding and faecal pellet production of late copepodite stages of Calanus finmarchicus were measured in mixtures of cultured autotrophic and heterotrophic food, as well as in a natural post-bloom plankton assemblage, in order to evaluate food selection and its potential effect on sedimentation of organic matter. Calanus finmarchicus consistently selected for diatoms, both in mixtures with the heterotrophic dinoflagellate Oxyrrhis marina, and in natural seston containing dinoflagellates, ciliates and flagellates. Similarly, the filtration, ingestion and faecal pellet production rates were significantly higher feeding on diatoms than when feeding on other food species. Calanus finmarchicus selection appeared relatively inflexible, so that changes in seston composition induced large changes in diet quantity and composition. Our results support the traditional view of C. finmarchicus as a major grazer of diatoms, and suggest potentially high post-bloom faecal pellet production rates.     

Effects of climate‐driven primary production change on marine food webs: implications for fisheries and conservation

Climate change is altering the rate and distribution of primary production in the world's oceans. Primary production is critical to maintaining biodiversity and supporting fishery catches, but predicting the response of populations to primary production change is complicated by predation and competition interactions. We simulated the effects of change in primary production on diverse marine ecosystems across a wide latitudinal range in Australia using the marine food web model Ecosim. We link models of primary production of lower trophic levels (phytoplankton and benthic producers) under climate change with Ecosim to predict changes in fishery catch, fishery value, biomass of animals of conservation interest, and indicators of community composition. Under a plausible climate change scenario, primary production will increase around Australia and generally this benefits fisheries catch and value and leads to increased biomass of threatened marine animals such as turtles and sharks. However, community composition is not strongly affected. Sensitivity analyses indicate overall positive linear responses of functional groups to primary production change. Responses are robust to the ecosystem type and the complexity of the model used. However, model formulations with more complex predation and competition interactions can reverse the expected responses for some species, resulting in catch declines for some fished species and localized declines of turtle and marine mammal populations under primary productivity increases. We conclude that climate‐driven primary production change needs to be considered by marine ecosystem managers and more specifically, that production increases can simultaneously benefit fisheries and conservation. Greater focus on incorporating predation and competition interactions into models will significantly improve the ability to identify species and industries most at risk from climate change.     

An evaluation of factors affecting vertical distribution among recruits of Calanus finmarchicus in three adjacent high-latitude localities

The vertical distributions of populations of Calanus finmarchicus are described in three different fjord areas near Tromse, northern Norway during May 1986. These localities (Malangen, Grøtsund and Balsfjorden) had characteristic differences in temperature, phytoplankton and population density of copepods. They probably are representative annual situations during the spring and summer period for coastal and fjord areas in northern Norway. Copepodite stage I and II C. finmarchicus are found in the surface waters (0–30 m) during a 24 h cycle, while the other stages appear to have a different diel depth distribution in Malangen. Pronounced differences in the depth distribution of the various copepodite stages and adult females were found in Grøtsund and Balsfjorden during the same period of the day on 20 and 21 May. The tendency for vertical overlap among CI–CV was clearly less pronounced in an environment with low phytoplankton standing stock and high population density of copepods. The patterns of vertical distribution are analysed by multidimentional scaling (MDS) and it is evident that the distribution pattern of C. finmarchicus is different at each locality. These preliminary results, are discussed in relation to ontogenetic vertical migration and aspects of resource partitioning and the possible importance of vertical separation for reducing competitive interactions between the different life stages of C. finmarchicus.     

High dispersal potential has maintained long-term population stability in the North Atlantic copepod Calanus finmarchicus

The cool-water copepod Calanus finmarchicus is a key species in North Atlantic marine ecosystems since it represents an important food resource for the developmental stages of several fish of major economic value. Over the last 40 years, however, data from the Continuous Plankton Recorder survey have highlighted a 70 per cent reduction in C. finmarchicus biomass, coupled with a gradual northward shift in the species's distribution, which have both been linked with climate change. To determine the potential for C. finmarchicus to track changes in habitat availability and maintain stable effective population sizes, we have assessed levels of gene flow and dispersal in current populations, as well as using a coalescent approach together with palaeodistribution modelling to elucidate the historical population demography of the species over previous changes in Earth's climate. Our findings indicate high levels of dispersal and a constant effective population size over the period 359,000-566,000 BP and suggest that C. finmarchicus possesses the capacity to track changes in available habitat, a feature that may be of crucial importance to the species's ability to cope with the current period of global climate change.     

Incorporating uncertainty in predictive species distribution modelling

Motivated by the need to solve ecological problems (climate change, habitat fragmentation and biological invasions), there has been increasing interest in species distribution models (SDMs). Predictions from these models inform conservation policy, invasive species management and disease-control measures. However, predictions are subject to uncertainty, the degree and source of which is often unrecognized. Here, we review the SDM literature in the context of uncertainty, focusing on three main classes of SDM: niche-based models, demographic models and process-based models. We identify sources of uncertainty for each class and discuss how uncertainty can be minimized or included in the modelling process to give realistic measures of confidence around predictions. Because this has typically not been performed, we conclude that uncertainty in SDMs has often been underestimated and a false precision assigned to predictions of geographical distribution. We identify areas where development of new statistical tools will improve predictions from distribution models, notably the development of hierarchical models that link different types of distribution model and their attendant uncertainties across spatial scales. Finally, we discuss the need to develop more defensible methods for assessing predictive performance, quantifying model goodness-of-fit and for assessing the significance of model covariates.     

Hemispheric‐scale patterns of climate‐related shifts in planktonic diatoms from North American and European lakes

A synthesis of over 200 diatom‐based paleolimnological records from nonacidified/nonenriched lakes reveals remarkably similar taxon‐specific shifts across the Northern Hemisphere since the 19th century. Our data indicate that these diatom shifts occurred in conjunction with changes in freshwater habitat structure and quality, which, in turn, we link to hemispheric warming trends. Significant increases in the relative abundances of planktonic Cyclotella taxa (P<0.01) were concurrent with sharp declines in both heavily silicified Aulacoseira taxa (P<0.01) and benthic Fragilaria taxa (P<0.01). We demonstrate that this trend is not limited to Arctic and alpine environments, but that lakes at temperate latitudes are now showing similar ecological changes. As expected, the onset of biological responses to warming occurred significantly earlier (P<0.05) in climatically sensitive Arctic regions (median age=ad 1870) compared with temperate regions (median age=ad 1970). In a detailed paleolimnological case study, we report strong relationships (P<0.005) between sedimentary diatom data from Whitefish Bay, Lake of the Woods (Ontario, Canada), and long‐term changes in air temperature and ice‐out records. Other potential environmental factors, such as atmospheric nitrogen deposition, could not explain our observations. These data provide clear evidence that unparalleled warming over the last few decades resulted in substantial increases in the length of the ice‐free period that, similar to 19th century changes in high‐latitude lakes, likely triggered a reorganization of diatom community composition. We show that many nonacidified, nutrient‐poor, freshwater ecosystems throughout the Northern Hemisphere have crossed important climatically induced ecological thresholds. These findings are worrisome, as the ecological changes that we report at both mid‐ and high‐latitude sites have occurred with increases in mean annual air temperature that are less than half of what is projected for these regions over the next half century.     

Fire severity mediates climate‐driven shifts in understorey community composition of black spruce stands of interior Alaska

Question: How do pre‐fire conditions (community composition and environmental characteristics) and climate‐driven disturbance characteristics (fire severity) affect post‐fire community composition in black spruce stands? Location: Northern boreal forest, interior Alaska. Methods: We compared plant community composition and environmental stand characteristics in 14 black spruce stands before and after multiple, naturally occurring wildfires. We used a combination of vegetation table sorting, univariate (ANOVA, paired t‐tests), and multivariate (detrended correspondence analysis) statistics to determine the impact of fire severity and site moisture on community composition, dominant species and growth forms. Results: Severe wildfires caused a 50% reduction in number of plant species in our study sites. The largest species loss, and therefore the greatest change in species composition, occurred in severely burned sites. This was due mostly to loss of non‐vascular species (mosses and lichens) and evergreen shrubs. New species recruited most abundantly to severely burned sites, contributing to high species turnover on these sites. As well as the strong effect of fire severity, pre‐fire and post‐fire mineral soil pH had an effect on post‐fire vegetation patterns, suggesting a legacy effect of site acidity. In contrast, pre‐fire site moisture, which was a strong determinant of pre‐fire community composition, showed no relationship with post‐fire community composition. Site moisture was altered by fire, due to changes in permafrost, and therefore post‐fire site moisture overrode pre‐fire site moisture as a strong correlate. Conclusions: In the rapidly warming climate of interior Alaska, changes in fire severity had more effect on post‐fire community composition than did environmental factors (moisture and pH) that govern landscape patterns of unburned vegetation. This suggests that climate change effects on future community composition of black spruce forests may be mediated more strongly by fire severity than by current landscape patterns. Hence, models that represent the effects of climate change on boreal forests could improve their accuracy by including dynamic responses to fire disturbance.     

Modelled spatial distribution of marine fish and projected modifications in the North Atlantic Ocean

The objectives of this work were to examine the past, current and potential influence of global climate change on the spatial distribution of some commercially exploited fish and to evaluate a recently proposed new ecological niche model (ENM) called nonparametric probabilistic ecological niche model (NPPEN). This new technique is based on a modified version of the test called Multiple Response Permutation Procedure (MRPP) using the generalized Mahalanobis distance. The technique was applied in the extratropical regions of the North Atlantic Ocean on eight commercially exploited fish species using three environmental parameters (sea surface temperature, bathymetry and sea surface salinity). The numerical procedure and the model allowed a better characterization of the niche (sensu Hutchinson) and an improved modelling of the spatial distribution of the species. Furthermore, the technique appeared to be robust to incomplete or bimodal training sets. Despite some potential limitations related to the choice of the climatic scenarios (A2 and B2), the type of physical model (ECHAM 4) and the absence of consideration of biotic interactions, modelled changes in species distribution explained some current observed shifts in dominance that occurred in the North Atlantic sector, and particularly in the North Sea. Although projected changes suggest a poleward movement of species, our results indicate that some species may not be able to track their climatic envelope and that climate change may have a prominent influence on fish distribution during this century. The phenomenon is likely to trigger locally major changes in the dominance of species with likely implications for socio‐economical systems. In this way, ENMs might provide a new management tool against which changes in the resource might be better anticipated.     

100 million years of turtle paleoniche dynamics enable the prediction of latitudinal range shifts in a warming world

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Modelling climate‐change‐induced nonlinear thresholds in cephalopod population dynamics

A significant global challenge lies in our current inability to anticipate, and therefore prepare for, critical ecological thresholds (i.e. tipping points in ecosystems). This deficit stems largely from an inadequate understanding of the many complex interactions between species and the environment at the ecosystem level, and the paucity of mechanistic models relating environment to population dynamics at the species level. In marine ecosystems, abundant, short‐lived and fast‐growing species such as anchovies or squids, consistently function as ‘keystone’ groups whose population dynamics affect entire ecosystems. Increasing exploitation coupled with climate change impacts has the potential to affect these ecological groups and consequently, the entire marine ecosystem. There are currently very few models that predict the impact of climate change on these keystone groups. Here we use a combination of individual‐based bioenergetics and stage‐structured population models to characterize the fundamental capacity of cephalopods to respond to climate change. We demonstrate the potential for, and mechanisms behind, two unfavourable climate‐change‐induced thresholds in future population dynamics. Although one threshold was the direct consequence of a decrease in incubation time caused by ocean warming, the other threshold was linked to survivorship, implying the possibility of management through a modification of fishing mortality. Additional substantive changes in phenology were also predicted, with a possible loss in population resilience. Our results demonstrate the feasibility of predicting complex nonlinear dynamics with a reasonably simplistic mechanistic model, and highlight the necessity of developing such approaches for other species if attempts to moderate the impact of climate change on natural resources are to be effective.     

Projected changes in plant species richness and extent of riparian vegetation belts as a result of climate‐driven hydrological change along the Vindel River in Sweden

1. Riparian plant communities are primarily structured by the hydrological regime of the stream. Models of climate change predict increased temperatures and changed patterns of precipitation that will alter the flow of rivers and streams with consequences for riparian communities. In boreal regions of Europe, stream flows will exhibit earlier spring‐flood peaks of lower magnitude, lower summer flows and higher flows in autumn and winter. We quantified the effects of predicted hydrological change on riparian plant species richness, using four different scenarios for the free‐flowing Vindel River in northern Sweden. 2. We calculated the hydrological niche of vegetation belts by relating the occurrence of species and vegetation belts to data on flood duration for 10 years preceding the vegetation survey. We then used the flood duration predicted for 2071–2100 to estimate expected changes in the extent of each vegetation belt. Using species accumulation curves, we then predicted changes in plant species richness as a result of changes in extent. 3. The two most species‐rich vegetation belts, riparian forest and willow shrub, were predicted to decrease most in elevational extent, up to 39 and 32%, respectively. The graminoid belt below the shrub belt will mainly shift upwards in elevation while the amphibious vegetation belt at the bottom of the riparian zone increases in size. 4. In the Vindel River, the riparian forest and willow shrub zone will lose most species, with reductions of 5–12% and 1–13% per site, respectively, depending on the scenario. The predicted loss from the entire riparian zone is lower, 1–9%, since many species occur in more than one vegetation belt. More extensive species losses are expected in the southern boreal zone for which much larger spring‐flood reductions are projected. 5. With an expected reduction in area of the most species‐rich belts, it becomes increasingly important to manage and protect riparian zones to alleviate other threats, thus minimising the risk of species losses. Restoring river and stream reaches degraded by other impacts to gain riparian habitat is another option to avoid species losses.     

Increases in lake phytoplankton biomass caused by future climate‐driven changes to seasonal river flow

For the many lakes world‐wide with short residence times, changes to the rate of water throughput may have important effects on lake ecology. We studied relationships between current and predicted residence times and phytoplankton biomass using a eutrophic lake in the north‐west of England with an annual residence time averaging about 20 days, as a test case. Using 32 years of recent hydrological flow data for Bassenthwaite Lake, multiple sets of scaled flow for each year, and the process‐based phytoplankton response model, PROTECH, we modelled the effects of changing river flow on phytoplankton biomass in the lake. The impact on biomass was shown to depend on seasonal changes in flow rather than annual changes. Furthermore, there was a qualitative difference in impact depending on whether the nutrient loading to the lake came principally from flow‐independent sources, or from flow‐dependent ones. Predictions for changes in river flow under future climate scenarios in the north‐west of England have suggested that, despite little change in the annual flow magnitude, there will be a shift to greater flow in the winter and lesser flow in the summer. Applying these flow predictions to our modelling of Bassenthwaite Lake revealed that, with flow‐independent nutrient loading, and no overall increase in nutrient load, phytoplankton abundance in the summer could increase by up to 70%, including an increased proportion of Cyanobacteria. Conversely, were the loading completely dependent on the flow, the biomass would fall. In many parts of the world, river flow is expected to decrease in the summer even more than in England, suggesting these areas may expect substantial changes to seasonal phytoplankton biomass as a result of climate‐driven changes to seasonal river flow. Such changes would be in addition to any other changes owing to warming effects or eutrophication.     

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 role of climate,habitat, and species co‐occurrence as drivers of change in small mammal distributions over the past century

Species distribution models are commonly used to predict species responses to climate change. However, their usefulness in conservation planning and policy is controversial because they are difficult to validate across time and space. Here we capitalize on small mammal surveys repeated over a century in Yosemite National Park, USA, to assess accuracy of model predictions. Historical (1900–1940) climate, vegetation, and species occurrence data were used to develop single‐ and multi‐species multivariate adaptive regression spline distribution models for three species of chipmunk. Models were projected onto the current (1980–2007) environmental surface and then tested against modern field resurveys of each species. We evaluated models both within and between time periods and found that even with the inclusion of biotic predictors, climate alone is the dominant predictor explaining the distribution of the study species within a time period. However, climate was not consistently an adequate predictor of the distributional change observed in all three species across time. For two of the three species, climate alone or climate and vegetation models showed good predictive performance across time. The stability of the distribution from the past to present observed in the third species, however, was not predicted by our modeling approach. Our results demonstrate that correlative distribution models are useful in understanding species' potential responses to environmental change, but also show how changes in species‐environment correlations through time can limit the predictive performance of models.     

The influence of coarse‐scale environmental features on current and predicted future distributions of narrow‐range endemic crayfish populations

1. A major limitation to effective management of narrow‐range crayfish populations is the paucity of information on the spatial distribution of crayfish species and a general understanding of the interacting environmental variables that drive current and future potential distributional patterns. 2. Maximum Entropy Species Distribution Modeling Software (MaxEnt) was used to predict the current and future potential distributions of four endemic crayfish species in the Ouachita Mountains. Current distributions were modelled using climate, geology, soils, land use, landform and flow variables thought to be important to lotic crayfish. Potential changes in the distribution were forecast by using models trained on current conditions and projecting onto the landscape predicted under climate‐change scenarios. 3. The modelled distribution of the four species closely resembled the perceived distribution of each species but also predicted populations in streams and catchments where they had not previously been collected. Soils, elevation and winter precipitation and temperature most strongly related to current distributions and represented 65–87% of the predictive power of the models. Model accuracy was high for all models, and model predictions of new populations were verified through additional field sampling. 4. Current models created using two spatial resolutions (1 and 4.5 km²) showed that fine‐resolution data more accurately represented current distributions. For three of the four species, the 1‐km² resolution models resulted in more conservative predictions. However, the modelled distributional extent of Orconectes leptogonopodus was similar regardless of data resolution. Field validations indicated 1‐km² resolution models were more accurate than 4.5‐km² resolution models. 5. Future projected (4.5‐km² resolution models) model distributions indicated three of the four endemic species would have truncated ranges with low occurrence probabilities under the low‐emission scenario, whereas two of four species would be severely restricted in range under moderate–high emissions. Discrepancies in the two emission scenarios probably relate to the exclusion of behavioural adaptations from species‐distribution models. 6. These model predictions illustrate possible impacts of climate change on narrow‐range endemic crayfish populations. The predictions do not account for biotic interactions, migration, local habitat conditions or species adaptation. However, we identified the constraining landscape features acting on these populations that provide a framework for addressing habitat needs at a fine scale and developing targeted and systematic monitoring programmes.