Forecasting range shifts of a cold‐adapted species under climate change: are genomic and ecological diversity within species crucial for future resilience?

Cold‐adapted taxa are experiencing severe range shifts due to climate change and are expected to suffer a significant reduction of their climatically suitable habitats in the next few decades. However, it has been proposed that taxa with sufficient standing genetic and ecologic diversity will better withstand climate change. These taxa are typically more broadly distributed in geographic and ecological niche space, therefore they are likely to endure higher levels of populations loss than more restricted, less diverse taxa before the effects of those losses impact their overall diversity and resilience. Here, we explore the potential relationship between intraspecific genetic and ecological diversity and future resilience, using the cold‐adapted plant Primula farinosa. We employ high‐throughput sequencing to assess the genomic diversity of phylogeographic lineages in P. farinosa. Additionally, we use current climatic variables to define niche breadth and niche differentiation across lineages. Finally, we calibrate species distribution models (SDMs) and project the climatic preferences of each lineage on future climate to predict lineage‐specific shifts in climatically suitable habitats. Our study predicts relative persistence of future suitable habitats for the most genetically and ecologically diverse lineages of the cold‐adapted P. farinosa, but significant reduction of them for two out of its four lineages. While we do not provide specific experiments aimed at identifying the causal links between genetic diversity and resilience to climate change, our results indicate that greater genetic diversity and wider ecological breadth may buffer species responses to rapid climatic changes. This study further highlights the importance of integrating knowledge of intraspecific diversity for predicting species fate in response to climate change.     

Has past climate change affected cold‐specialized species differentially through space and time?

Quaternary climate change has been strongly linked to distributional shifts and recent species diversification. Montane species, in particular, have experienced enhanced isolation and rapid genetic divergence during glacial fluctuations, and these processes have resulted in a disproportionate number of neo‐endemic species forming in high‐elevation habitats. In temperate montane environments, a general model of alpine population history is well supported, where cold‐specialized species track favourable climate conditions downslope during glacial episodes and upslope during warmer interglacial periods, which leads to a climate‐driven population or species diversification pump. However, it remains unclear how geography mediates distributional changes and whether certain episodes of glacial history have differentially impacted rates of diversification. We address these questions by examining phylogenomic data in a North American clade of flightless, cold‐specialized insects, the ice crawlers (Insecta: Grylloblattodea: Grylloblattidae: Grylloblatta). These low‐vagility organisms have the potential to reveal highly localized refugia and patterns of spatial recolonization, as well as a longer history of in situ diversification. Using continuous phylogeographic analysis of species groups, we show that all species tend to retreat to nearby low‐elevation habitats across western North America during episodes of glaciation, but species at high latitude exhibit larger distributional shifts. Lineage diversification was examined over the course of the Neogene and Quaternary periods, with statistical analysis supporting a direct association between climate variation and diversification rate. Major increases in lineage diversification appear to be correlated with warm and dry periods, rather than with extreme glacial events. Finally, we identify substantial cryptic diversity among ice crawlers, leading to high endemism across their range. This diversity provides new insights into highly localized glacial refugia for cold‐specialized species across western North America.     

Sources or sinks? The responses of tropical forests to current and future climate and atmospheric composition.

How tropical rainforests are responding to the ongoing global changes in atmospheric composition and climate is little studied and poorly understood. Although rising atmospheric carbon dioxide (CO2) could enhance forest productivity, increased temperatures and drought are likely to diminish it. The limited field data have produced conflicting views of the net impacts of these changes so far. One set of studies has seemed to point to enhanced carbon uptake; however, questions have arisen about these findings, and recent experiments with tropical forest trees indicate carbon saturation of canopy leaves and no biomass increase under enhanced CO2. Other field observations indicate decreased forest productivity and increased tree mortality in recent years of peak temperatures and drought (strong El Niño episodes). To determine current climatic responses of forests around the world tropics will require careful annual monitoring of ecosystem performance in representative forests. To develop the necessary process-level understanding of these responses will require intensified experimentation at the whole-tree and stand levels. Finally, a more complete understanding of tropical rainforest carbon cycling is needed for determining whether these ecosystems are carbon sinks or sources now, and how this status might change during the next century.     

Long‐distance dispersal or postglacial contraction? Insights into disjunction between Himalaya–Hengduan Mountains and Taiwan in a cold‐adapted herbaceous genus,Triplostegia

Current disjunct patterns can result from long‐distance dispersal or postglacial contraction. We herein investigate the evolutionary history of Triplostegia to elucidate the disjunction between the Himalaya–Hengduan Mountain region (HHM) and Taiwan (TW). Genetic structure of Triplostegia was investigated for 48 populations using sequences from five chloroplast loci and the ribosomal nuclear internal transcribed spacer. Divergence time estimation, ancestral area reconstruction, and species distribution modeling (SDM) were employed to examine the biogeographic history of Triplostegia. Substantial genetic differentiation among populations from southwestern China (SW), Central China (CC), and TW was detected. Triplostegia was inferred to have originated in SW, and diversification began during the late Miocene; CC was colonized in the mid‐Pliocene, and TW was finally colonized in the early Pleistocene. SDM suggested an expansion of climatically suitable areas during the Last Glacial Maximum and range contraction during the Last interglacial in Triplostegia. Disjunction between HHM and TW in Triplostegia is most likely the consequence of topographic isolation and postglacial contraction. The potential climatic suitability areas for Triplostegia by 2070s (2061–2080) are predicted to slightly shrink and move northward. With continued global warming and human‐induced deforestation, extinction risk may increase for the cold‐adapted species, and appropriate strategies should be employed for ecosystem conservation.     

Plant extinction risk under climate change: are forecast range shifts alone a good indicator of species vulnerability to global warming?

Models that couple habitat suitability with demographic processes offer a potentially improved approach for estimating spatial distributional shifts and extinction risk under climate change. Applying such an approach to five species of Australian plants with contrasting demographic traits, we show that: (i) predicted climate‐driven changes in range area are sensitive to the underlying habitat model, regardless of whether demographic traits and their interaction with habitat patch configuration are modeled explicitly; and (ii) caution should be exercised when using predicted changes in total habitat suitability or geographic extent to infer extinction risk, because the relationship between these metrics is often weak. Measures of extinction risk, which quantify threats to population persistence, are particularly sensitive to life‐history traits, such as recruitment response to fire, which explained approximately 60% of the deviance in expected minimum abundance. Dispersal dynamics and habitat patch structure have the strongest influence on the amount of movement of the trailing and leading edge of the range margin, explaining roughly 40% of modeled structural deviance. These results underscore the need to consider direct measures of extinction risk (population declines and other measures of stochastic viability), as well as measures of change in habitat area, when assessing climate change impacts on biodiversity. Furthermore, direct estimation of extinction risk incorporates important demographic and ecosystem processes, which potentially influence species’ vulnerability to extinction due to climate change.     

Can we project changes in fish abundance and distribution in response to climate?

Large‐scale and long‐term changes in fish abundance and distribution in response to climate change have been simulated using both statistical and process‐based models. However, national and regional fisheries management requires also shorter term projections on smaller spatial scales, and these need to be validated against fisheries data. A 26‐year time series of fish surveys with high spatial resolution in the North‐East Atlantic provides a unique opportunity to assess the ability of models to correctly simulate the changes in fish distribution and abundance that occurred in response to climate variability and change. We use a dynamic bioclimate envelope model forced by physical–biogeochemical output from eight ocean models to simulate changes in fish abundance and distribution at scales down to a spatial resolution of 0.5°. When comparing with these simulations with annual fish survey data, we found the largest differences at the 0.5° scale. Differences between fishery model runs driven by different biogeochemical models decrease dramatically when results are aggregated to larger scales (e.g. the whole North Sea), to total catches rather than individual species or when the ensemble mean instead of individual simulations are used. Recent improvements in the fidelity of biogeochemical models translate into lower error rates in the fisheries simulations. However, predictions based on different biogeochemical models are often more similar to each other than they are to the survey data, except for some pelagic species. We conclude that model results can be used to guide fisheries management at larger spatial scales, but more caution is needed at smaller scales.     

Life‐history innovation to climate change: can single‐brooded migrant birds become multiple breeders?

When climatic conditions change and become outside the range experienced in the past, species may show life‐history innovations allowing them to adapt in new ways. We report such an innovation for pied flycatchers Ficedula hypoleuca. Decades of breeding biological studies on pied flycatchers have rarely reported multiple breeding in this long‐distance migrant. In two populations, we found 12 recent incidents of females with second broods, all produced by extremely early laying females in warm springs. As such early first broods are a recent phenomenon, because laying dates have gradually advanced over time, this innovation now allows individual females to enhance their reproductive success considerably. If laying dates continue advancing, potentially more females may become multiple breeders and selection for early (and multiple) breeding phenotypes increases, which may accelerate adaptation to climatic change.     

Shifting altitudinal distribution of outbreak zones of winter moth Operophtera brumata in sub‐arctic birch forest: a response to recent climate warming?

Climatic change is expected to affect the extent and severity of geometrid moth outbreaks, a major disturbance factor in sub-arctic birch forests. Previous studies have reported that the two geometrid species involved, autumnal moth and winter moth, differ in their temperature requirements and, consequently, in their altitudinal and latitudinal distribution patterns. In this study, we document the altitudinal distribution of winter moth outbreaks in a large coastal area in northern Norway. We show that, in the present winter moth outbreak, defoliated birch stands were seen as distinct zones with a rather constant width in the uppermost part of the forest and where the upper limit coincided with the forest line. The outbreak zone closely followed the spatially variable forest line as an undulating belt, although some of the variation in outbreak zone width was also related to variation in topographical variables, such as distance from the coast, forest line altitude, and slope of the terrain. A distinct outbreak zone at the altitudinal forest line is the typical picture that has been depicted in more qualitative historical records on previous outbreaks of autumnal moth rather than winter moth. We suggest that the recent documented climate warming in this region may have induced a shift in distribution of the winter moth both relative to topography and geography. Further investigation is, however, required to substantiate these suspicions.     

Can changes in the distributions of resident birds in China over the past 50?years be attributed to climate change?

The distributions of bird species have changed over the past 50 years in China. To evaluate whether the changes can be attributed to the changing climate, we analyzed the distributions of 20 subspecies of resident birds in relation to climate change. Long‐term records of bird distributions, gray relational analysis, fuzzy‐set classification techniques, and attribution methods were used. Among the 20 subspecies of resident birds, the northern limits of over half of the subspecies have shifted northward since the 1960s, and most changes have been related to the thermal index. Driven by climate change over the past 50 years, the suitable range and latitude or longitude of the distribution centers of certain birds have exhibited increased fluctuations. The northern boundaries of over half of the subspecies have shifted northward compared with those in the 1960s. The consistency between the observed and predicted changes in the range limits was quite high for some subspecies. The changes in the northern boundaries or the latitudes of the centers of distribution of nearly half of the subspecies can be attributed to climate change. The results suggest that climate change has affected the distributions of particular birds. The method used to attribute changes in bird distributions to climate change may also be effective for other animals.     

Does a long‐term oscillation in nitrogen concentration reflect climate impact on submerged vegetation and vulnerability to state shifts in a shallow lake?

Various ecosystems, including shallow lakes, are suggested to possess alternative stable state dynamics. The response of such systems to environmental change is non-linear and not fully reversible, which calls for identification of feedback mechanisms and subtle changes connected to structural stability. Here, we used a 25-year data series on water chemistry to make inferences on processes prior to a recent shift from a clear to a turbid state in Lake Tåkern, Sweden. Before the shift, annual concentration of total organic nitrogen (TON) described a cyclic pattern, with a periodicity of eight years. Annual TON was negatively correlated with the magnitude of a summer decline in calcium carbonate, treated as a proxy of the seasonal production of submerged vegetation. Cross-correlations of TON and the north Atlantic oscillation (NAO) indicated a possible connection to climate. The strongest correlation was obtained by a three-year lag of the NAO index. Using a set of linear time series models, the most parsimonious model was a 3^rd order autoregressive model with NAO, delayed three years, as a covariate. Analyses of seasonality indicated that the delayed NAO signal was strongly correlated with summer TON. Also, the autocorrelation function was very similar for these two time series, and autoregressive models including NAO as a covariate were strongly supported (sum of Akaike weights=0.93). These results indicate that climate may have contributed to the regime shift through lowered macrophyte production at the time of the shift, and therefore most likely also a depleted resilience of the clear water state. The delayed effect of climate is suggested to result from indirect and inter-year dependent response of submerged vegetation to fish kills during harsh winters.     

Fish-induced life-history shifts in the cladocerans Daphnia and Simocephalus: are they positive or negative responses?

The cladocerans Daphnia and Simocephalus were exposed to fishsmells under different food conditions, and their life-historyparameters were analysed. They showed similar responses to fishsmells and food shortage: lowered juvenile growth rate, reducedmature size, elongated maturation time, reduced brood size andincreased offspring size. The life-history changes induced bythe fish smell might be a result of deleterious effects of thefactor. Although it has been considered that fish-induced life-historyshifts in Daphnia are positive responses to predators, our resultssuggest that they are negative responses of animals that showsome other positive responses to fish, such as behavioural andmorphological changes to reduce mortality due to predation.Synergism was detected in the effects of the fish smell andfood deficiency, which suggests that fish smell reduces thetolerance of Daphnia to an environmental stress, food shortage.     

Discontinuous or semi‐discontinuous DNA replication in Escherichia coli?

The postulate that a stalled/collapsed replication fork will be generated when the replication complex encounters a UV-induced lesion in the template for leading-strand DNA synthesis is based on the model of semi-discontinuous DNA replication. A review of existing data indicates that the semi-discontinuous DNA replication model is supported by data from in vitro studies, while the discontinuous DNA replication model is supported by in vivo studies in Escherichia coli. Until the question of whether DNA replicates discontinuously in one or both strands is clearly resolved, any model building based on either one of the two DNA replication models should be treated with caution.     

The future of soil invertebrate communities in polar regions: different climate change responses in the Arctic and Antarctic?

The polar regions are experiencing rapid climate change with implications for terrestrial ecosystems. Here, despite limited knowledge, we make some early predictions on soil invertebrate community responses to predicted twenty‐first century climate change. Geographic and environmental differences suggest that climate change responses will differ between the Arctic and Antarctic. We predict significant, but different, belowground community changes in both regions. This change will be driven mainly by vegetation type changes in the Arctic, while communities in Antarctica will respond to climate amelioration directly and indirectly through changes in microbial community composition and activity, and the development of, and/or changes in, plant communities. Climate amelioration is likely to allow a greater influx of non‐native species into both the Arctic and Antarctic promoting landscape scale biodiversity change. Non‐native competitive species could, however, have negative effects on local biodiversity particularly in the Arctic where the communities are already species rich. Species ranges will shift in both areas as the climate changes potentially posing a problem for endemic species in the Arctic where options for northward migration are limited. Greater soil biotic activity may move the Arctic towards a trajectory of being a substantial carbon source, while Antarctica could become a carbon sink.     

Can amphibians take the heat? Vulnerability to climate warming in subtropical and temperate larval amphibian communities

Predicting the biodiversity impacts of global warming implies that we know where and with what magnitude these impacts will be encountered. Amphibians are currently the most threatened vertebrates, mainly due to habitat loss and to emerging infectious diseases. Global warming may further exacerbate their decline in the near future, although the impact might vary geographically. We predicted that subtropical amphibians should be relatively susceptible to warming‐induced extinctions because their upper critical thermal limits (CT_max) might be only slightly higher than maximum pond temperatures (T_max). We tested this prediction by measuring CT_max and T_max for 47 larval amphibian species from two thermally distinct subtropical communities (the warm community of the Gran Chaco and the cool community of Atlantic Forest, northern Argentina), as well as from one European temperate community. Upper thermal tolerances of tadpoles were positively correlated (controlling for phylogeny) with maximum pond temperatures, although the slope was steeper in subtropical than in temperate species. CT_max values were lowest in temperate species and highest in the subtropical warm community, which paradoxically, had very low warming tolerance (CT_max–T_max) and therefore may be prone to future local extinction from acute thermal stress if rising pond T_max soon exceeds their CT_max. Canopy‐protected subtropical cool species have larger warming tolerance and thus should be less impacted by peak temperatures. Temperate species are relatively secure to warming impacts, except for late breeders with low thermal tolerance, which may be exposed to physiological thermal stress in the coming years.     

Are ontogenetic shifts in foliar structure and resource acquisition spatially conditioned in tank‐bromeliads?

The phenotypic plasticity of plants has been explored as a function of either ontogeny (apparent plasticity) or environment (adaptive plasticity), although few studies have analyzed these factors together. In the present study, we take advantage of the dispersal of Aechmea mertensii bromeliads by Camponotus femoratus or Pachycondyla goeldii ants in shaded and sunny environments, respectively, to quantify ontogenetic changes in morphological, foliar, and functional traits, and to analyze ontogenetic and ant species effects on 14 traits. Most of the morphological (plant height, number of leaves), foliar (leaf thickness, leaf mass area, total water content, trichome density), and functional (leaf δ¹³C) traits differed as a function of ontogeny. Conversely, only leaf δ¹⁵N showed an adaptive phenotypic plasticity. On the other hand, plant width, tank width, longest leaf length, stomatal density, and leaf C concentration showed an adaptation to local environment with ontogeny. The exception was leaf N concentration, which showed no trend at all. Aechmea mertensii did not show an abrupt morphological modification such as in heteroblastic bromeliads, although it was characterized by strong, size‐related functional modifications for CO₂ acquisition. The adaptive phenotypic variation found between the two ant species indicates the spatially conditioned plasticity of A. mertensii in the context of insect‐assisted dispersal. However, ant‐mediated effects on phenotypic plasticity in A. mertensii are not obvious because ant species and light environment are confounding variables. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 299–312.     

The spatial distribution of avian relatives: do obligate army‐ant‐following birds roost and feed near family members?

The ocellated antbird ( Phaenostictus mcleannani ) feeds in groups and therefore is an informative species in which to study the biological factors that modulate avian group living. These birds congregate at swarms of army ants to capture fleeing prey, and previous observations suggest that males may be philopatric, feed with close relatives, and defend communal feeding ranges. We assessed whether kin selection could be an important factor maintaining group formation in a population of ocellated antbirds inhabiting continuous forest at La Selva Biological Station, Costa Rica, using radiotelemetry and 15 novel microsatellite markers. We predicted that the roosting areas of closely related adult males should overlap and that adult males feeding simultaneously at the same swarm should be highly related. We banded and genotyped 65 individuals (≥ 88% of the population) and radiotagged 30 of them. The results generally did not conform to our predictions. Little overlap occurred among the roosting areas of same-sex individuals, and nearest roosting neighbours (either same or opposite sex) were generally unrelated. A small proportion of male dyads suggested short-distance dispersal, but in general the distribution of genotypes within the study area approached randomness. We found little evidence of natal philopatry in either sex. Less than half of the feeding groups sampled included highly related males; most consisted of unrelated individuals. Hence, we found limited potential for kin selection to favour group living and suggest that other factors, particularly direct benefits (e.g. food intake), are probably more important than indirect effects (nepotism).