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.     

Incorporating spatial autocorrelation into species distribution models alters forecasts of climate‐mediated range shifts

Species distribution models (SDMs) are widely used to forecast changes in the spatial distributions of species and communities in response to climate change. However, spatial autocorrelation (SA) is rarely accounted for in these models, despite its ubiquity in broad‐scale ecological data. While spatial autocorrelation in model residuals is known to result in biased parameter estimates and the inflation of type I errors, the influence of unmodeled SA on species' range forecasts is poorly understood. Here we quantify how accounting for SA in SDMs influences the magnitude of range shift forecasts produced by SDMs for multiple climate change scenarios. SDMs were fitted to simulated data with a known autocorrelation structure, and to field observations of three mangrove communities from northern Australia displaying strong spatial autocorrelation. Three modeling approaches were implemented: environment‐only models (most frequently applied in species' range forecasts), and two approaches that incorporate SA; autologistic models and residuals autocovariate (RAC) models. Differences in forecasts among modeling approaches and climate scenarios were quantified. While all model predictions at the current time closely matched that of the actual current distribution of the mangrove communities, under the climate change scenarios environment‐only models forecast substantially greater range shifts than models incorporating SA. Furthermore, the magnitude of these differences intensified with increasing increments of climate change across the scenarios. When models do not account for SA, forecasts of species' range shifts indicate more extreme impacts of climate change, compared to models that explicitly account for SA. Therefore, where biological or population processes induce substantial autocorrelation in the distribution of organisms, and this is not modeled, model predictions will be inaccurate. These results have global importance for conservation efforts as inaccurate forecasts lead to ineffective prioritization of conservation activities and potentially to avoidable species extinctions.     

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.     

Climate,copepods and seabirds in the boreal Northeast Atlantic – current state and future outlook

The boreal Northeast Atlantic is strongly affected by current climate change, and large shifts in abundance and distribution of many organisms have been observed, including the dominant copepod Calanus finmarchicus, which supports the grazing food web and thus many fish populations. At the same time, large‐scale declines have been observed in many piscivorous seabirds, which depend on abundant small pelagic fish. Here, we combine predictions from a niche model of C. finmarchicus with long‐term data on seabird breeding success to link trophic levels. The niche model shows that environmental suitability for C. finmarchicus has declined in southern areas with large breeding seabird populations (e.g. the North Sea), and predicts that this decline is likely to spread northwards during the 21st century to affect populations in Iceland and the Faroes. In a North Sea colony, breeding success of three common piscivorous seabird species [black‐legged kittiwake (Rissa tridactyla), common guillemot (Uria aalge) and Atlantic puffin (Fratercula arctica)] was strongly positively correlated with local environmental suitability for C. finmarchicus, whereas this was not the case at a more northerly colony in west Norway. Large seabird populations seem only to occur where C. finmarchicus is abundant, and northward distributional shifts of common boreal seabirds are therefore expected over the coming decades. Whether or not population size can be maintained depends on the dispersal ability and inclination of these colonial breeders, and on the carrying capacity of more northerly areas in a warmer climate.     

Reconstructing the demise of Tethyan plants: climate‐driven range dynamics of Laurus since the Pliocene

Aim Climate changes are thought to be responsible for the retreat and eventual extinction of subtropical lauroid species that covered much of Europe and North Africa during the Palaeogene and early Neogene; little is known, however, of the spatial and temporal patterns of this demise. Herein we calibrate ecological niche models to assess the climatic requirements of Laurus L. (Lauraceae), an emblematic relic from the Tethyan subtropical flora, subsequently using these models to infer how the range dynamics of Laurus were affected by Plio‐Pleistocene climate changes. We also provide predictions of likely range changes resulting from future climatic scenarios. Location The Mediterranean Basin and Macaronesian islands (Canaries, Madeira, Azores). Methods We used a maximum‐entropy algorithm (Maxent) to model the relationship between climate and Laurus distribution over time. The models were fitted both to the present and to the middle Pliocene, based on fossil records. We employed climatic reconstructions for the mid‐Pliocene (3 Ma), the Last Glacial Maximum (21 ka) and a CO₂‐doubling future scenario to project putative species distribution in each period. We validated the model projections with Laurus fossil and present occurrences. Results Laurus preferentially occupied warm and moist areas with low seasonality, showing a marked stasis of its climatic niche. Models fitted to Pliocene conditions successfully predicted the current species distribution. Large suitable areas existed during the Pliocene, which were strongly reduced during the Pleistocene, but humid refugia within the Mediterranean Basin and Macaronesian islands enabled long‐term persistence. Future climate conditions are likely to re‐open areas suitable for colonization north of the current range. Main conclusions The climatic requirements of Laurus remained virtually unchanged over the last 3 Myr. This marked niche conservatism imposed largely deterministic range dynamics driven by climate conditions. This species's relatively high drought tolerance might account for the survival of Laurus in continental Europe throughout the Quaternary whilst other Lauraceae became extinct. Climatic scenarios for the end of this century would favour an expansion of the species's range towards northern latitudes, while severely limiting southern populations due to increased water stress.     

Compositional shifts in Costa Rican forests due to climate‐driven species migrations

Species are predicted to shift their distributions upslope or poleward in response to global warming. This prediction is supported by a growing number of studies documenting species migrations in temperate systems but remains poorly tested for tropical species, and especially for tropical plant species. We analyzed changes in tree species composition in a network of 10 annually censused 1‐ha plots spanning an altitudinal gradient of 70–2800 m elevation in Costa Rica. Specifically, we combined plot data with herbarium records (accessed through GBIF) to test if the plots' community temperature scores (CTS, average thermal mean of constituent species weighted by basal area) have increased over the past decade as is predicted by climate‐driven species migrations. In addition, we quantified the contributions of stem growth, recruitment, and mortality to the observed patterns. Supporting our a priori hypothesis of upward species migrations, we found that there have been consistent directional shifts in the composition of the plots, such that the relative abundance of lowland species, and hence CTS, increased in 90% of plots. The rate of the observed compositional shifts corresponds to a mean thermal migration rate (TMR) of 0.0065 °C yr^?1 (95% CI = 0.0005–0.0132 °C yr^?1). While the overall TMR is slower than predicted based on concurrent regional warming of 0.0167 °C yr^?1, migrations were on pace with warming in 4 of the 10 plots. The observed shifts in composition were driven primarily by mortality events (i.e., the disproportionate death of highland vs. lowland species), suggesting that individuals of many tropical tree species will not be able to tolerate future warming and thus their persistence in the face of climate change will depend on successful migrations. Unfortunately, in Costa Rica and elsewhere, land area inevitably decreases at higher elevations; hence, even species that are able to migrate successfully will face heightened risks of extinction.     

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.     

Warming of Subarctic waters accelerates development of a key marine zooplankton Calanus finmarchicus

Recent observations confirm the rising temperatures of Atlantic waters transported into the Arctic Ocean via the West Spitsbergen Current (WSC). We studied the overall abundance and population structure of the North Atlantic keystone zooplankton copepod Calanus finmarchicus, which is the main prey for pelagic fish and some seabirds, in relation to selected environmental variables in this area between 2001 and 2011, when warming in the Arctic and Subarctic was particularly pronounced. Sampling within a 3‐week time window each summer demonstrated that trends in the overall abundance of C. finmarchicus varied between years, with the highest values in “extreme” years, due to high numbers of nauplii and early copepodite stages in colder years (2001, 2004, 2010), and contrary to that, the fifth copepodite stage (C5) peaking in warm years (2006, 2007, 2009). The most influential environmental variable driving C. finmarchicus life cycle was temperature, which promoted an increased C5 abundance when the temperature was above 6°C, indicating earlier spawning and/or accelerated development, and possibly leading to their development to adults later in the summer and spawning for the second time, given adequate food supply. Based on the presented high interannual and spatial variability, we hypothesize that under a warmer climate, C. finmarchicus may annually produce two generations in the southern part of the WSC, what in turn could lead to food web reorganization of important top predators, such as little auks, and induce northward migrations of fish, especially the Norwegian herring.     

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.     

Projected climate‐driven faunal movement routes

Historically, many species moved great distances as climates changed. However, modern movements will be limited by the patterns of human‐dominated landscapes. Here, we use a combination of projected climate‐driven shifts in the distributions of 2903 vertebrate species, estimated current human impacts on the landscape, and movement models, to determine through which areas in the western hemisphere species will likely need to move to track suitable climates. Our results reveal areas with projected high densities of climate‐driven movements – including, the Amazon Basin, the southeastern United States and southeastern Brazil. Some of these regions, such as southern Bolivia and northern Paraguay, contain relatively intact landscapes, whereas others such as the southeastern United States and Brazil are heavily impacted by human activities. Thus, these results highlight both critical areas for protecting lands that will foster movement, and barriers where human land‐use activities will likely impede climate‐driven shifts in species distributions.     

Climate‐driven shifts in the distribution of koala‐browse species from the Last Interglacial to the near future

The koala's Phascolarctos cinereus distribution is currently restricted to eastern and south‐eastern Australia. However, fossil records dating from 70 ± 4 ka (ka = 10³ yr) from south‐western Australia and the Nullarbor Plain are evidence of subpopulation extinctions in the southwest at least after the Last Interglacial (~128–116 ka). We hypothesize that koala sub‐population extinctions resulted from the eastward retraction of the koala's main browse species in response to unsuitable climatic conditions. We further posit a general reduction in the distribution of main koala‐browse trees in the near future in response climate change. We modelled 60 koala‐browse species and constructed a set of correlative species distribution models for five time periods: Last Interglacial (~128–116 ka), Last Glacial Maximum (~23–19 ka), Mid‐Holocene (~7–5 ka), present (interpolations of observed data, representative of 1960–1990), and 2070. We based our projections on five hindcasts and one forecast of climatic variables extracted from WorldClim based on two general circulation models (considering the most pessimistic scenario of high greenhouse‐gas emissions) and topsoil clay fraction. We used 17 dates of koala fossil specimens identified as reliable from 70 (± 4) to 535 (± 49) ka, with the last appearance of koalas at 70 ka in the southwest. The main simulated koala‐browse species were at their greatest modelled extent of suitability during the Last Glacial Maximum, with the greatest loss of koala habitat occurring between the Mid‐Holocene and the present. We predict a similar habitat loss between the present and 2070. The spatial patterns of habitat change support our hypothesis that koala extinctions in the southwest, Nullarbor Plain and central South Australia resulted from the eastward retraction of the dominant koala‐browse species in response to long‐term climate changes. Future climate patterns will likely increase the extinction risk of koalas in their remaining eastern ranges.     

Projected impacts of 21st century climate change on diapause in Calanus finmarchicus

Diapause plays a key role in the life cycle of high latitude zooplankton. During diapause, animals avoid starving in winter by living in deep waters where metabolism is lower and met by lipid reserves. Global warming is therefore expected to shorten the maximum potential diapause duration by increasing metabolic rates and by reducing body size and lipid reserves. This will alter the phenology of zooplankton, impact higher trophic levels and disrupt biological carbon pumps. Here, we project the impacts of climate change on the key North Atlantic copepod Calanus finmarchicus under IPCC RCP 8.5. Potential diapause duration is modelled in relation to body size and overwintering temperature. The projections show pronounced geographic variations. Potential diapause duration reduces by more than 30% in the Western Atlantic, whereas in the key overwintering centre of the Norwegian Sea it changes only marginally. Surface temperature rises, which reduce body size and lipid reserves, will have a similar impact to deep‐water changes on diapause in many regions. Because deep‐water warming lags that at the surface, animals in the Labrador Sea could offset warming impacts by diapausing in deeper waters. However, the ability to control diapause depth may be limited.     

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.     

Evolutionary consequences of climate‐induced range shifts in insects

Range shifts can rapidly create new areas of geographic overlap between formerly allopatric taxa and evidence is accumulating that this can affect species persistence. We review the emerging literature on the short‐ and long‐term consequences of these geographic range shifts. Specifically, we focus on the evolutionary consequences of novel species interactions in newly created sympatric areas by describing the potential (i) short‐term processes acting on reproductive barriers between species and (ii) long‐term consequences of range shifts on the stability of hybrid zones, introgression and ultimately speciation and extinction rates. Subsequently, we (iii) review the empirical literature on insects to evaluate which processes have been studied, and (iv) outline some areas that deserve increased attention in the future, namely the genomics of hybridisation and introgression, our ability to forecast range shifts and the impending threat from insect vectors and pests on biodiversity, human health and crop production. Our review shows that species interactions in de novo sympatric areas can be manifold, sometimes increasing and sometimes decreasing species diversity. A key issue that emerges is that climate‐induced hybridisations in insects are much more widespread than anticipated and that rising temperatures and increased anthropogenic disturbances are accelerating the process of species mixing. The existing evidence only shows the tip of the iceberg and we are likely to see many more cases of species mixing following range shifts in the near future.     

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.     

Potential source and sink locations for climate‐driven species range shifts in Europe since the Last Glacial Maximum

Aim To identify potential source and sink locations for climate‐driven species range shifts in Europe since the Last Glacial Maximum (LGM). Location Europe. Methods We developed a new approach combining past‐climate simulations with the concept of analogous climate space. Our index gives a continuous measure of the potential of a location to have acted as a source or a sink for species that have shifted their ranges since the LGM. High glacial source potential is indicated by LGM climatic conditions that are widespread now; high post‐glacial sink potential is indicated by current climatic conditions that were widespread at the LGM. The degree of isolation of source and sink areas was calculated as the median distance to areas with analogous climate conditions. Results We identified areas of high glacial source potential in the previously recognized refugial areas in the southern European peninsulas, but also in large areas in central‐western Europe. The most climatically isolated source areas were located in northern Spain, in north‐western Europe and in eastern Turkey. From here species would have had to cover substantial distances to find current climate conditions analogous to LGM conditions of these areas. Areas with high post‐glacial sink potential were mainly located in Fennoscandia and in central and south‐eastern Europe. Some of the most isolated sink areas were located in the Spanish highlands and around the Baltic Sea. Main conclusions Our species‐independent approach successfully identified previously recognized glacial refugial areas with high source potential for species range shifts in southern Europe and in addition highlighted other potential source areas in central Europe. This study offers new insights into how the distribution of past and current climatic conditions may have influenced past species range shifts and current large‐scale biodiversity patterns.     

Predicting when climate‐driven phenotypic change affects population dynamics

Species' responses to climate change are variable and diverse, yet our understanding of how different responses (e.g. physiological, behavioural, demographic) relate and how they affect the parameters most relevant for conservation (e.g. population persistence) is lacking. Despite this, studies that observe changes in one type of response typically assume that effects on population dynamics will occur, perhaps fallaciously. We use a hierarchical framework to explain and test when impacts of climate on traits (e.g. phenology) affect demographic rates (e.g. reproduction) and in turn population dynamics. Using this conceptual framework, we distinguish four mechanisms that can prevent lower‐level responses from impacting population dynamics. Testable hypotheses were identified from the literature that suggest life‐history and ecological characteristics which could predict when these mechanisms are likely to be important. A quantitative example on birds illustrates how, even with limited data and without fully‐parameterized population models, new insights can be gained; differences among species in the impacts of climate‐driven phenological changes on population growth were not explained by the number of broods or density dependence. Our approach helps to predict the types of species in which climate sensitivities of phenotypic traits have strong demographic and population consequences, which is crucial for conservation prioritization of data‐deficient species.