Alien and native plant life‐forms respond differently to human and climate pressures

Aim To investigate whether differences in the elevational trend in native and alien species richness were dependent on climate or human pressures. Specifically we tested whether life‐form and/or alien/native status modifies the response of plant species richness to human population and temperature along: (1) a complete elevational gradient, and (2) within separate elevational bands that, by keeping temperature within a narrow range, elucidate the effects of human pressures more clearly. Location Two provinces (c. 7507 km²) on the southern border of the European Alps (Italy), subdivided into 240 contiguous sampling cells (c. 35.7 km²). Methods We used an extensive dataset on alien and native species richness across an elevation gradient (20–2900 m a.s.l.). Richness of natives and naturalized aliens were separately related to temperature, human population and Raunkiaer life‐form using general linear mixed models. Life‐form describes different plant strategies for survival during seasons with adverse cold/arid conditions. Results The relationship between species richness and temperature for natives was strongly dependent on life‐form, while aliens showed a consistent positive trend. Similar trends across alien and native life‐forms were found for the relationship between species richness and human population along the whole gradient and within separate elevational bands. Main conclusions The absence of life‐form‐dependent responses amongst aliens supports the hypothesis that the distribution of alien plant species richness was more related to propagule pressure and availability of novel niches created by human activities than to climatic filtering. While climate change will potentially contribute to relaxing species thermal constraints, the response of alien species to future warming will also be contingent on changes in anthropogenic pressures.     

Phenology in a warming world: differences between native and non‐native plant species

Phenology is a harbinger of climate change, with many species advancing flowering in response to rising temperatures. However, there is tremendous variation among species in phenological response to warming, and any phenological differences between native and non‐native species may influence invasion outcomes under global warming. We simulated global warming in the field and found that non‐native species flowered earlier and were more phenologically plastic to temperature than natives, which did not accelerate flowering in response to warming. Non‐native species' flowering also became more synchronous with other community members under warming. Earlier flowering was associated with greater geographic spread of non‐native species, implicating phenology as a potential trait associated with the successful establishment of non‐native species across large geographic regions. Such phenological differences in both timing and plasticity between native and non‐natives are hypothesised to promote invasion success and population persistence, potentially benefiting non‐native over native species under climate change.     

Meta‐analyzing the likely cross‐species responses to climate change

Ecological Niche Models (ENMs) have different performances in predicting potential geographic distributions. Here we meta‐analyzed the likely effects of climate change on the potential geographic distribution of 1,205 bird species from the Neotropical region, modeled using eight ENMs and three Atmosphere‐Ocean General Circulation Models (AOGCM). We considered the variability in ENMs performance to estimate a weighted mean difference between potential geographic distributions for baseline and future climates. On average, potential future ranges were projected to be from 25.7% to 44.5% smaller than current potential ranges across species. However, we found that 0.2% to 18.3% of the total variance in range shifts occurred “within species” (i.e., owing to the use of different modeling techniques and climate models) and 81.7% to 99.8% remained between species (i.e., it could be explained by ecological correlates). Using meta‐analytical techniques akin to regression, we also showed that potential range shifts are barely predicted by bird biological traits. We demonstrated that one can combine and reduce species‐specific effects with high uncertainty in ENMs and also explore potential causes of climate change effect on species using meta‐analytical tools. We also highlight that the search for powerful correlates of climate change‐induced range shifts can be a promising line of investigation.     

Ocean cleaning stations under a changing climate: biological responses of tropical and temperate fish‐cleaner shrimp to global warming

Cleaning symbioses play an important role in the health of certain coastal marine communities. These interspecific associations often occur at specific sites (cleaning stations) where a cleaner organism (commonly a fish or shrimp) removes ectoparasites/damaged tissue from a ‘client’ (a larger cooperating fish). At present, the potential impact of climate change on the fitness of cleaner organisms remains unknown. This study investigated the physiological and biochemical responses of tropical (Lysmata amboinensis) and temperate (L. seticaudata) cleaner shrimp to global warming. Specifically, thermal limits (CTMax), metabolic rates, thermal sensitivity, heat shock response (HSR), lipid peroxidation [malondialdehyde (MDA) concentration], lactate levels, antioxidant (GST, SOD and catalase) and digestive enzyme activities (trypsin and alkaline phosphatase) at current and warming (+3 °C) temperature conditions. In contrast to the temperate species, CTMax values decreased significantly from current (24–27 °C) to warming temperature conditions (30 °C) for the tropical shrimp, where metabolic thermal sensitivity was affected and the HSR was significantly reduced. MDA levels in tropical shrimp increased dramatically, indicating extreme cellular lipid peroxidation, which was not observed in the temperate shrimp. Lactate levels, GST and SOD activities were significantly enhanced within the muscle tissue of the tropical species. Digestive enzyme activities in the hepatopancreas of both species were significantly decreased by warmer temperatures. Our data suggest that the tropical cleaner shrimp will be more vulnerable to global warming than the temperate Lysmata seticaudata; the latter evolved in a relatively unstable environment with seasonal thermal variations that may have conferred greater adaptive plasticity. Thus, tropical cleaning symbioses may be challenged at a greater degree by warming‐related anthropogenic forcing, with potential cascading effects on the health and structuring of tropical coastal communities (e.g. coral reefs).     

Trophic niche segregation allows range‐extending coral reef fishes to co‐exist with temperate species under climate change

Changing climate is forcing many terrestrial and marine species to extend their ranges poleward to stay within the bounds of their thermal tolerances. However, when such species enter higher latitude ecosystems, they engage in novel interactions with local species, such as altered predator–prey dynamics and competition for food. Here, we evaluate the trophic overlap between range‐extending and local fish species along the east coast of temperate Australia, a hotspot for ocean warming and species range extensions. Stable isotope ratios (δ¹⁵N and δ¹³C) of muscle tissue and stomach content analysis were used to quantify overlap of trophic niche space between vagrant tropical and local temperate fish communities along a 730 km (6°) latitudinal gradient. Our study shows that in recipient temperate ecosystems, sympatric tropical and temperate species do not overlap significantly in their diet—even though they forage on broadly similar prey groups—and are therefore unlikely to compete for trophic niche space. The tropical and temperate species we studied, which are commonly found in shallow‐water coastal environments, exhibited moderately broad niche breadths and local‐scale dietary plasticity, indicating trophic generalism. We posit that because these species are generalists, they can co‐exist under current climate change, facilitating the existence of novel community structures.     

Adding climate change to the mix: responses of aquatic ectotherms to the combined effects of eutrophication and warming

The threat of excessive nutrient enrichment, or eutrophication, is intensifying across the globe as climate change progresses, presenting a major management challenge. Alterations in precipitation patterns and increases in temperature are increasing nutrient loadings in aquatic habitats and creating conditions that promote the proliferation of cyanobacterial blooms. The exacerbating effects of climate warming on eutrophication are well established, but we lack an in-depth understanding of how aquatic ectotherms respond to eutrophication and warming in tandem. Here, I provide a brief overview and critique of studies exploring the cumulative impacts of eutrophication and warming on aquatic ectotherms, and provide forward direction using mechanistically focused, multi-threat experiments to disentangle complex interactions. Evidence to date suggests that rapid warming will exacerbate the negative effects of eutrophication on aquatic ectotherms, but gradual warming will induce physiological remodelling that provides protection against nutrients and hypoxia. Moving forward, research will benefit from a greater focus on unveiling cause and effect mechanisms behind interactions and designing treatments that better mimic threat dynamics in nature. This approach will enable robust predictions of species responses to ongoing eutrophication and climate warming and enable the integration of climate warming into eutrophication management policies.     

The responses of microbial temperature relationships to seasonal change and winter warming in a temperate grassland

Microorganisms dominate the decomposition of organic matter and their activities are strongly influenced by temperature. As the carbon (C) flux from soil to the atmosphere due to microbial activity is substantial, understanding temperature relationships of microbial processes is critical. It has been shown that microbial temperature relationships in soil correlate with the climate, and microorganisms in field experiments become more warm‐tolerant in response to chronic warming. It is also known that microbial temperature relationships reflect the seasons in aquatic ecosystems, but to date this has not been investigated in soil. Although climate change predictions suggest that temperatures will be mostly affected during winter in temperate ecosystems, no assessments exist of the responses of microbial temperature relationships to winter warming. We investigated the responses of the temperature relationships of bacterial growth, fungal growth, and respiration in a temperate grassland to seasonal change, and to 2 years’ winter warming. The warming treatments increased winter soil temperatures by 5–6°C, corresponding to 3°C warming of the mean annual temperature. Microbial temperature relationships and temperature sensitivities (Q₁₀) could be accurately established, but did not respond to winter warming or to seasonal temperature change, despite significant shifts in the microbial community structure. The lack of response to winter warming that we demonstrate, and the strong response to chronic warming treatments previously shown, together suggest that it is the peak annual soil temperature that influences the microbial temperature relationships, and that temperatures during colder seasons will have little impact. Thus, mean annual temperatures are poor predictors for microbial temperature relationships. Instead, the intensity of summer heat‐spells in temperate systems is likely to shape the microbial temperature relationships that govern the soil‐atmosphere C exchange.     

Water-mediated responses of ecosystem carbon fluxes to climatic change in a temperate steppe

Global warming and a changing precipitation regime could have a profound impact on ecosystem carbon fluxes, especially in arid and semiarid grasslands where water is limited. A field experiment manipulating temperature and precipitation has been conducted in a temperate steppe in northern China since 2005. A paired, nested experimental design was used, with increased precipitation as the primary factor and warming simulated by infrared radiators as the secondary factor. The results for the first 2 yr showed that gross ecosystem productivity (GEP) was higher than ecosystem respiration, leading to net C sink (measured by net ecosystem CO(2) exchange, NEE) over the growing season in the study site. The interannual variation of NEE resulted from the difference in mean annual precipitation. Experimental warming reduced GEP and NEE, whereas increased precipitation stimulated ecosystem C and water fluxes in both years. Increased precipitation also alleviated the negative effect of experimental warming on NEE. The results demonstrate that water availability plays a dominant role in regulating ecosystem C and water fluxes and their responses to climatic change in the temperate steppe of northern China.     

The Mediterranean Sea as a ‘cul‐de‐sac’ for endemic fishes facing climate change

The Mediterranean Sea is a hotspot of biodiversity, and climate warming is expected to have a significant influence on its endemic fish species. However, no previous studies have predicted whether fish species will experience geographic range extensions or contractions as a consequence of warming. Here, we projected the potential future climatic niches of 75 Mediterranean Sea endemic fish species based on a global warming scenario implemented with the Mediterranean model OPAMED8 and a multimodel inference, which included uncertainty. By 2070–2099, the average surface temperature of the Mediterranean Sea was projected to warm by 3.1 °C. Projections for 2041–2060 are that 25 species would qualify for the International Union for the Conservation of Nature and Natural Resources (IUCN) Red List, and six species would become extinct. By 2070–2099, 45 species were expected to qualify for the IUCN Red List whereas 14 were expected to become extinct. By the middle of the 21st century, the coldest areas of the Mediterranean Sea (Adriatic Sea and Gulf of Lion) would act as a refuge for cold‐water species, but by the end of the century, those areas were projected to become a ‘cul‐de‐sac’ that would drive those species towards extinction. In addition, the range size of endemic species was projected to undergo extensive fragmentation, which is a potentially aggravating factor. Since a majority of endemic fishes are specialists, regarding substratum and diet, we may expect a reduced ability to track projected climatic niches. As a whole, 25% of the Mediterranean Sea continental shelf was predicted to experience a total modification of endemic species assemblages by the end of the 21st century. This expected turnover rate could be mitigated by marine protected areas or accelerated by fishing pressure or competition from exotic fishes. It remains a challenge to predict how these assemblage modifications might affect ecosystem function.     

Variable evolutionary response to regional climate change in a polymorphic species

The evolutionary response to regional and global climate change may vary in widespread polymorphic species, so predicting future genetic responses will require careful tracking of genetic variability in local populations. We surveyed chromosomal inversion polymorphisms in 25 populations of Drosophila robusta, many of which have been sampled repeatedly starting in the 1940s, 50s, and 60s up until 2007, across its range in the USA. Frequencies of some northerly, or cold‐adapted, gene arrangements have declined in the face of increasing temperatures, whereas frequencies of several southern, or warm‐adapted, gene arrangements were positively correlated with increasing temperature changes. Over a finer geographic scale, populations from the west‐central part of the species range from the Ozark Plateau, Ouachita mountains, and eastern Oklahoma showed genetic differentiation between south‐central Ozark and western Ozark/Ouachita regions that has persisted in the face of recent shifts in gene arrangement frequencies. Overall, populations of D. robusta exhibited dynamic genetic changes over time, with some populations shifting chromosome frequencies in just 10–15 years. Some temporal genetic shifts were widespread and significantly correlated with temperature increases, but regions of the genome marked by different gene arrangements have responded in different sections of the species range. In some parts of the species range, chromosome frequencies shifted but were not associated with changing temperatures, showed little or no temporal change, or temporal shifts stopped for temperature sensitive gene arrangements near fixation. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 702–718.     

Thermal plasticity of photosynthesis: the role of acclimation in forest responses to a warming climate

The increasing air temperatures central to climate change predictions have the potential to alter forest ecosystem function and structure by exceeding temperatures optimal for carbon gain. Such changes are projected to threaten survival of sensitive species, leading to local extinctions, range migrations, and altered forest composition. This study investigated photosynthetic sensitivity to temperature and the potential for acclimation in relation to the climatic provenance of five species of deciduous trees, Liquidambar styraciflua, Quercus rubra, Quercus falcata, Betula alleghaniensis, and Populus grandidentata. Open‐top chambers supplied three levels of warming (+0, +2, and +4 °C above ambient) over 3 years, tracking natural temperature variability. Optimal temperature for CO₂ assimilation was strongly correlated with daytime temperature in all treatments, but assimilation rates at those optima were comparable. Adjustment of thermal optima was confirmed in all species, whether temperatures varied with season or treatment, and regardless of climate in the species' range or provenance of the plant material. Temperature optima from 17° to 34° were observed. Across species, acclimation potentials varied from 0.55 °C to 1.07 °C per degree change in daytime temperature. Responses to the temperature manipulation were not different from the seasonal acclimation observed in mature indigenous trees, suggesting that photosynthetic responses should not be modeled using static temperature functions, but should incorporate an adjustment to account for acclimation. The high degree of homeostasis observed indicates that direct impacts of climatic warming on forest productivity, species survival, and range limits may be less than predicted by existing models.     

Water‐use efficiency in response to climate change: from leaf to ecosystem in a temperate steppe

Water‐use efficiency (WUE) has been recognized as an important characteristic of ecosystem productivity, which links carbon (C) and water cycling. However, little is known about how WUE responds to climate change at different scales. Here, we investigated WUE at leaf, canopy, and ecosystem levels under increased precipitation and warming from 2005 to 2008 in a temperate steppe in Northern China. We measured gross ecosystem productivity (GEP), net ecosystem CO₂ exchange (NEE), evapotranspiration (ET), evaporation (E), canopy transpiration (T_c), as well as leaf photosynthesis (P_max) and transpiration (T_l) of a dominant species to calculate canopy WUE (WUE_c=GEP/T), ecosystem WUE (WUE_gep=GEP/ET or WUE_nee=NEE/ET) and leaf WUE (WUE_l=P_max/T_l). The results showed that increased precipitation stimulated WUE_c, WUE_gep and WUE_nee by 17.1%, 10.2% and 12.6%, respectively, but decreased WUE_l by 27.4%. Climate warming reduced canopy and ecosystem WUE over the 4 years but did not affect leaf level WUE. Across the 4 years and the measured plots, canopy and ecosystem WUE linearly increased, but leaf level WUE of the dominant species linearly decreased with increasing precipitation. The differential responses of canopy/ecosystem WUE and leaf WUE to climate change suggest that caution should be taken when upscaling WUE from leaf to larger scales. Our findings will also facilitate mechanistic understanding of the C–water relationships across different organism levels and in projecting the effects of climate warming and shifting precipitation regimes on productivity in arid and semiarid ecosystems.     

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.     

Leaf‐trait plasticity and species vulnerability to climate change in a Mongolian steppe

Climate change is expected to modify plant assemblages in ways that will have major consequences for ecosystem functions. How climate change will affect community composition will depend on how individual species respond, which is likely related to interspecific differences in functional traits. The extraordinary plasticity of some plant traits is typically neglected in assessing how climate change will affect different species. In the Mongolian steppe, we examined whether leaf functional traits under ambient conditions and whether plasticity in these traits under altered climate could explain climate‐induced biomass responses in 12 co‐occurring plant species. We experimentally created three probable climate change scenarios and used a model selection procedure to determine the set of baseline traits or plasticity values that best explained biomass response. Under all climate change scenarios, plasticity for at least one leaf trait correlated with change in species performance, while functional leaf‐trait values in ambient conditions did not. We demonstrate that trait plasticity could play a critical role in vulnerability of species to a rapidly changing environment. Plasticity should be considered when examining how climate change will affect plant performance, species' niche spaces, and ecological processes that depend on plant community composition.     

Landscape resistance mediates native fish species distribution shifts and vulnerability to climate change in riverscapes

A broader understanding of how landscape resistance influences climate change vulnerability for many species is needed, as is an understanding of how barriers to dispersal may impact vulnerability. Freshwater biodiversity is at particular risk, but previous studies have focused on popular cold‐water fishes (e.g., salmon, trout, and char) with relatively large body sizes and mobility. Those fishes may be able to track habitat change more adeptly than less mobile species. Smaller, less mobile fishes are rarely represented in studies demonstrating effects of climate change, but depending on their thermal tolerance, they may be particularly vulnerable to environmental change. By revisiting 280 sites over a 20 year interval throughout a warming riverscape, we described changes in occupancy (i.e., site extirpation and colonization probabilities) and assessed the environmental conditions associated with those changes for four fishes spanning a range of body sizes, thermal and habitat preferences. Two larger‐bodied trout species exhibited small changes in site occupancy, with bull trout experiencing a 9.2% (95% CI = 8.3%–10.1%) reduction, mostly in warmer stream reaches, and westslope cutthroat trout experiencing a nonsignificant 1% increase. The small‐bodied cool water slimy sculpin was originally distributed broadly throughout the network and experienced a 48.0% (95% CI = 42.0%–54.0%) reduction in site occupancy with declines common in warmer stream reaches and areas subject to wildfire disturbances. The small‐bodied comparatively warmer water longnose dace primarily occupied larger streams and increased its occurrence in the lower portions of connected tributaries during the study period. Distribution shifts for sculpin and dace were significantly constrained by barriers, which included anthropogenic water diversions, natural step‐pools and cascades in steeper upstream reaches. Our results suggest that aquatic communities exhibit a range of responses to climate change, and that improving passage and fluvial connectivity will be important climate adaptation tactics for conserving aquatic biodiversity.     

Synergistic and antagonistic effects of land use and non‐native species on community responses to climate change

Climate change, land‐use change and introductions of non‐native species are key determinants of biodiversity change worldwide. However, the extent to which anthropogenic drivers of environmental change interact to affect biological communities is largely unknown, especially over longer time periods. Here, we show that plant community composition in 996 Swedish landscapes has consistently shifted to reflect the warmer and wetter climate that the region has experienced during the second half of the 20th century. Using community climatic indices, which reflect the average climatic associations of the species within each landscape at each time period, we found that species compositions in 74% of landscapes now have a higher representation of warm‐associated species than they did previously, while 84% of landscapes now host more species associated with higher levels of precipitation. In addition to a warmer and wetter climate, there have also been large shifts in land use across the region, while the fraction of non‐native species has increased in the majority of landscapes. Climatic warming at the landscape level appeared to favour the colonization of warm‐associated species, while also potentially driving losses in cool‐associated species. However, the resulting increases in community thermal means were apparently buffered by landscape simplification (reduction in habitat heterogeneity within landscapes) in the form of increased forest cover. Increases in non‐native species, which generally originate from warmer climates than Sweden, were a strong driver of community‐level warming. In terms of precipitation, both landscape simplification and increases in non‐natives appeared to favour species associated with drier climatic conditions, to some extent counteracting the climate‐driven shift towards wetter communities. Anthropogenic drivers can act both synergistically and antagonistically to determine trajectories of change in biological communities over time. Therefore, it is important to consider multiple drivers of global change when trying to understand, manage and predict biodiversity in the future.     

Thermodependent bacterial pathogens and mass mortalities in temperate benthic communities: a new case of emerging disease linked to climate change

In the temperate north-western Mediterranean Sea, large-scale disease outbreaks in benthic invertebrate species have recently occurred during climatic anomalies characterized by elevated seawater temperatures. One of the most affected species was the red gorgonian Paramuricea clavata , a key species of highly diverse communities dwelling in dim-lit habitats in the Mediterranean. From diseased P. clavata colonies, we isolated culturable bacteria associated to tissue lesions in order to investigate their potential as pathogens. Inoculation of four bacterial isolates onto healthy P. clavata in aquaria caused disease signs similar to those observed during the 2003 mortality event. The infection process was dependent on elevated seawater temperatures, in a range of values consistent with recordings performed in the field during the climatic anomalies. Among the four isolates, we identified a Vibrio coralliilyticus strain that showed virulence to P. clavata . This strain was re-isolated from diseased colonies during the experimentations. V. coralliilyticus has been previously identified as a thermodependent pathogen of a tropical coral species, emphasizing a causal role of this infectious agent in the P. clavata disease. Taking into consideration predicted global warming over the coming decades, a better understanding of the factors and mechanisms that affect the disease process will be of critical importance in predicting future threats to temperate gorgonians communities in the Mediterranean Sea.     

Species interactions and climate change: How the disruption of species co‐occurrence will impact on an avian forest guild

Interspecific interactions are crucial in determining species occurrence and community assembly. Understanding these interactions is thus essential for correctly predicting species' responses to climate change. We focussed on an avian forest guild of four hole‐nesting species with differing sensitivities to climate that show a range of well‐understood reciprocal interactions, including facilitation, competition and predation. We modelled the potential distributions of black woodpecker and boreal, tawny and Ural owl, and tested whether the spatial patterns of the more widespread species (excluding Ural owl) were shaped by interspecific interactions. We then modelled the potential future distributions of all four species, evaluating how the predicted changes will alter the overlap between the species' ranges, and hence the spatial outcomes of interactions. Forest cover/type and climate were important determinants of habitat suitability for all species. Field data analysed with N‐mixture models revealed effects of interspecific interactions on current species abundance, especially in boreal owl (positive effects of black woodpecker, negative effects of tawny owl). Climate change will impact the assemblage both at species and guild levels, as the potential area of range overlap, relevant for species interactions, will change in both proportion and extent in the future. Boreal owl, the most climate‐sensitive species in the guild, will retreat, and the range overlap with its main predator, tawny owl, will increase in the remaining suitable area: climate change will thus impact on boreal owl both directly and indirectly. Climate change will cause the geographical alteration or disruption of species interaction networks, with different consequences for the species belonging to the guild and a likely spatial increase of competition and/or intraguild predation. Our work shows significant interactions and important potential changes in the overlap of areas suitable for the interacting species, which reinforce the importance of including relevant biotic interactions in predictive climate change models for increasing forecast accuracy.