Drought-induced dynamics of carbon and water use efficiency of global grasslands from 2000 to 2011

Drought is frequently recorded as a result of climate warming and elevated concentration of greenhouse gases, which affect the carbon and water cycles in terrestrial ecosystems, particularly in arid and semi-arid regions. To identify the drought in grassland ecosystems and to determine how such drought affects grassland ecosystems in terms of carbon and water cycles across the globe, this study evaluated the drought conditions of global grassland ecosystems from 2000 to 2011 on the basis of the remotely sensed Drought Severity Index (DSI) data. The temporal dynamics of grassland carbon use efficiency (CUE) and water use efficiency (WUE), as well as their correlations with DSI, were also investigated at the global scale. Results showed that 57.04% of grassland ecosystems experienced a dry trend over this period. In general, most grassland ecosystems in the northern hemisphere (N.H.) were in near normal condition, whereas those in the southern hemisphere (S.H.) experienced a clear drying and wetting trend, with the year 2005 regarded as the turning point. Grassland CUE increased continually despite the varied drought conditions over this period. By contrast, WUE increased in the closed shrublands and woody savannas but decreased in all the other grassland types. The drought conditions affected the carbon and water use mainly by influencing the primary production and evapotranspiration of grass through photosynthesis and transpiration process. The CUE and WUE of savannas was most sensitive to droughts among all the grassland types. The areas of grassland DSI that showed significant correlations with CUE and WUE were 52.92% and 22.11% of the total grassland areas, respectively. Overall, droughts sufficiently explained the dynamics of grassland CUE, especially in the S.H. In comparison with grassland CUE, the grassland WUE was less sensitive to drought conditions at the global scale.     

Long‐term change within a Neotropical forest: assessing differential functional and floristic responses to disturbance and drought

Disentangling the relative roles of biotic and abiotic forces influencing forest structure, function, and local community composition continues to be an important goal in ecology. Here, utilizing two forest surveys 20‐year apart from a Central American dry tropical forest, we assess the relative role of past disturbance and local climatic change in the form of increased drought in driving forest dynamics. We observe: (i) a net decrease in the number of trees; (ii) a decrease in total forest biomass by 7.7 Mg ha^?1 but when calculated on subquadrat basis the biomass per unit area did not change indicating scale sensitivity of forest biomass measures; (iii) that the decrease in the number of stems occurred mainly in the smallest sizes, and in more moist and evergreen habitats; (iv) that there has been an increase in the proportion of trees that are deciduous, compound leaved and are canopy species, and a concomitant reduction in trees that are evergreen, simple‐leaved, and understory species. These changes are opposite to predictions based on recovery from disturbance, and have resulted in (v) a uniform multivariate shift from a more mesic to a more xeric forest. Together, our results show that over relatively short time scales, community composition and the functional dominance may be more responsive to climate change than recovery to past disturbances. Our findings point to the importance of assessing proportional changes in forest composition and not just changes in absolute numbers. Our findings are also consistent with the hypothesis that tropical tree species exhibit differential sensitivity to changes in precipitation. Predicted future decreases in rainfall may result in quick differential shifts in forest function, physiognomy, and species composition. Quantifying proportional functional composition offers a basis for a predictive framework for how the structure, and diversity of tropical forests will respond to global change.     

Long‐term phytoplankton community changes in a deep subalpine lake: responses to nutrient availability and climatic fluctuations

1. In natural lakes, modifications in the species composition and abundance of phytoplankton communities may ultimately be responses to changes in nutrient availability and climatic fluctuations. Phytoplankton and associated environmental factors were collected at monthly intervals from the beginning of the 1990s to 2007 in the large subalpine Lake Garda (z_max = 350 m, V = 49 × 10⁹ m³). In this study period, the lake showed a slight and continuous increase of total phosphorus (TP) in the water column, up to concentrations of 18–20 μg P L^?1. This increase represented the last stage of a long‐term process of enrichment documented since the 1970s, when concentrations of TP were below or around 10 μg P L^?1. 2. At the community level, annual phytoplankton cycles underwent a unidirectional and slow shift mainly due to changes in the species more affected by the nutrient enrichment of the lake. After a first and long period of dominance by conjugatophytes (Mougeotia) and diatoms (Fragilaria), phytoplankton biomass in recent years was sustained by cyanobacteria (Planktothrix). Other important modifications in the development of phytoplankton were superimposed on this pattern due to the effects of annual climate fluctuations principally mediated by the deep mixing events at spring overturn and, secondarily, by temperature and thermal stability of the water column during the growing season. 3. Interannual variations in the stability and temperature of the water column appeared to influence the development of a few subdominant flagellates (dinophytes and cryptophytes). Nevertheless, the major impact of climate on phytoplankton was indirect, and mediated through the effects of winter climatic conditions on deep mixing dynamics. Winter climatic fluctuations proved to be a key element in a linked chain of causal factors including cooling of hypolimnetic waters, deep vertical mixing and epilimnetic nutrient replenishment. The process of fertilisation was measurable both for TP and dissolved inorganic nitrogen, although only the first had a large effect, reinforcing the seasonal growth of a few dominant groups. The degree of nutrient replenishment further increased the spring development of large diatoms and the increase of Planktothrix in summer and autumn. 4. Currently, changes in nutrient concentrations have the greatest effect on the phytoplankton community, while direct effects due to the interannual variations in the thermal regime are of secondary importance compared with the indirect effects mediated through deep water mixing and spring fertilisation. Overall, the results demonstrate that the consequences of climatic fluctuations and climate warming on phytoplankton communities need to be studied at different levels of complexity and integration, from the direct effects of temperature and thermal regime, to the indirect effects mediated by the physiographic characteristics of water bodies.     

Habitat distribution models for intertidal seaweeds: responses to climatic and non‐climatic drivers

Aim Because intertidal organisms often live close to their physiological tolerance limits, they are potentially sensitive indicators of climate‐driven changes in the environment. The goals of this study were to assess the effect of climatic and non‐climatic factors on the geographical distribution of intertidal macroalgae, and to predict future distributions under different climate‐warming scenarios. Location North‐western Iberian Peninsula, southern Europe. Methods We developed distribution models for six ecologically important intertidal seaweed species. Occurrence and microhabitat data were sampled at 1‐km² resolution and analysed with climate variables measured at larger spatial scales. We used generalized linear models and applied the deviance and Bayesian information criterion to model the relationship between environmental variables and the distribution of each target species. We also used hierarchical partitioning (HP) to identify predictor variables with higher independent explanatory power. Results The distributions of Himanthalia elongata and Bifurcaria bifurcata were correlated with measures of terrestrial and marine climate, although in opposite directions. Model projections under two warming scenarios indicated the extinction of the former at a faster rate in the Cantabrian Sea (northern Spain) than in the Atlantic (west). In contrast, these models predicted an increase in the occurrence of B. bifurcata in both areas. The occurrences of Ascophyllum nodosum and Pelvetia canaliculata, species showing rather static historical distributions, were related to specific non‐climatic environmental conditions and locations, such as the location of sheltered sites. At the southernmost distributional limit, these habitats may present favourable microclimatic conditions or provide refuges from competitors or natural enemies. Model performances for Fucus vesiculosus and F. serratus were similar and poor, but several climatic variables influenced the occurrence of the latter in the HP analyses. Main conclusions The correlation between species distributions and climate was evident for two species, whereas the distributions of the others were associated with non‐climatic predictors. We hypothesize that the distribution of F. serratus responds to diverse combinations of factors in different sections of the north‐west Iberian Peninsula. Our study shows how the response of species distributions to climatic and non‐climatic variables may be complex and vary geographically. Our analyses also highlight the difficulty of making predictions based solely on variation in climatic factors measured at coarse spatial scales.     

Distance‐dependent seedling mortality and long‐term spacing dynamics in a neotropical forest community

Negative distance dependence (NDisD), or reduced recruitment near adult conspecifics, is thought to explain the astounding diversity of tropical forests. While many studies show greater mortality at near vs. far distances from adults, these studies do not seek to track changes in the peak seedling curve over time, thus limiting our ability to link NDisD to coexistence. Using census data collected over 12 years from central Panama in conjunction with spatial mark‐connection functions, we show evidence for NDisD for many species, and find that the peak seedling curve shifts away from conspecific adults over time. We find wide variation in the strength of NDisD, which was correlated with seed size and canopy position, but other life‐history traits showed no relationship with variation in NDisD mortality. Our results document shifts in peak seedling densities over time, thus providing evidence for the hypothesized spacing mechanism necessary for diversity maintenance in tropical forests.     

Carbon dynamics in the future forest: the importance of long‐term successional legacy and climate–fire interactions

Understanding how climate change may influence forest carbon (C) budgets requires knowledge of forest growth relationships with regional climate, long‐term forest succession, and past and future disturbances, such as wildfires and timber harvesting events. We used a landscape‐scale model of forest succession, wildfire, and C dynamics (LANDIS‐II) to evaluate the effects of a changing climate (A2 and B1 IPCC emissions; Geophysical Fluid Dynamics Laboratory General Circulation Models) on total forest C, tree species composition, and wildfire dynamics in the Lake Tahoe Basin, California, and Nevada. The independent effects of temperature and precipitation were assessed within and among climate models. Results highlight the importance of modeling forest succession and stand development processes at the landscape scale for understanding the C cycle. Due primarily to landscape legacy effects of historic logging of the Comstock Era in the late 1880s, C sequestration may continue throughout the current century, and the forest will remain a C sink (Net Ecosystem Carbon Balance > 0), regardless of climate regime. Climate change caused increases in temperatures limited simulated C sequestration potential because of augmented fire activity and reduced establishment ability of subalpine and upper montane trees. Higher temperatures influenced forest response more than reduced precipitation. As the forest reached its potential steady state, the forest could become C neutral or a C source, and climate change could accelerate this transition. The future of forest ecosystem C cycling in many forested systems worldwide may depend more on major disturbances and landscape legacies related to land use than on projected climate change alone.     

Long‐term physiological and growth responses of Himalayan fir to environmental change are mediated by mean climate

High‐elevation forests are experiencing high rates of warming, in combination with CO₂ rise and (sometimes) drying trends. In these montane systems, the effects of environmental changes on tree growth are also modified by elevation itself, thus complicating our ability to predict effects of future climate change. Tree‐ring analysis along an elevation gradient allows quantifying effects of gradual and annual environmental changes. Here, we study long‐term physiological (ratio of internal to ambient CO₂, i.e., C_i/C_a and intrinsic water‐use efficiency, iWUE) and growth responses (tree‐ring width) of Himalayan fir (Abies spectabilis) trees in response to warming, drying, and CO₂ rise. Our study was conducted along elevational gradients in a dry and a wet region in the central Himalaya. We combined dendrochronology and stable carbon isotopes (δ¹³C) to quantify long‐term trends in C_i/C_a ratio and iWUE (δ¹³C‐derived), growth (mixed‐effects models), and evaluate climate sensitivity (correlations). We found that iWUE increased over time at all elevations, with stronger increase in the dry region. Climate–growth relations showed growth‐limiting effects of spring moisture (dry region) and summer temperature (wet region), and negative effects of temperature (dry region). We found negative growth trends at lower elevations (dry and wet regions), suggesting that continental‐scale warming and regional drying reduced tree growth. This interpretation is supported by δ¹³C‐derived long‐term physiological responses, which are consistent with responses to reduced moisture and increased vapor pressure deficit. At high elevations (wet region), we found positive growth trends, suggesting that warming has favored tree growth in regions where temperature most strongly limits growth. At lower elevations (dry and wet regions), the positive effects of CO₂ rise did not mitigate the negative effects of warming and drying on tree growth. Our results raise concerns on the productivity of Himalayan fir forests at low and middle (<3,300 m) elevations as climate change progresses.     

Species and community responses to short‐term climate manipulation: Microarthropods in the sub‐Antarctic

Abstract: Both species and community‐level investigations are important for understanding the biotic impacts of climate change, because current evidence suggests that individual species responses are idiosyncratic. However, few studies of climate change impacts have been conducted on entire terrestrial arthropod communities living in the same habitat in the southern Hemisphere, and the effects of precipitation changes on them are particularly poorly understood. Here we investigate the species‐ and community‐level responses of microarthropods inhabiting a keystone plant species, on sub‐Antarctic Marion Island, to experimental reduction in precipitation, warming and shading. These climate manipulations were chosen based on observed climate trends and predicted indirect climate change impacts on this system. The dry‐warm and shade inducing treatments that were imposed effected significant species‐ and community‐level responses after a single year. Although the strongest community‐level trends included a dramatic decline in springtail abundance and total biomass under the dry‐warm and shade treatments, species responses were generally individualistic, that is, springtails responded differently to mites, and particular mite and springtail species responded differently to each other. Our results therefore provide additional support for the dynamic rather than static model for community responses to climate change, in the first such experiment in the sub‐Antarctic. In conclusion, these results show that an ongoing decline in precipitation and increase in temperature is likely to have dramatic direct and indirect effects on this microarthropod community. Moreover, they indicate that while at a broad scale it may be possible to make generalizations regarding species responses to climate change, these generalizations are unlikely to translate into predictable effects at the community level.     

Long‐term shifts in abundance and distribution of a temperate fish fauna: a response to climate change and fishing practices

Aim South‐eastern Australia is a climate change hotspot with well‐documented recent changes in its physical marine environment. The impact on and temporal responses of the biota to change are less well understood, but appear to be due to influences of climate, as well as the non‐climate related past and continuing human impacts. We attempt to resolve the agents of change by examining major temporal and distributional shifts in the fish fauna and making a tentative attribution of causal factors. Location Temperate seas of south‐eastern Australia. Methods Mixed data sources synthesized from published accounts, scientific surveys, spearfishing and angling competitions, commercial catches and underwater photographic records, from the ‘late 1800s’ to the ‘present’, were examined to determine shifts in coastal fish distributions. Results Forty‐five species, representing 27 families (about 30% of the inshore fish families occurring in the region), exhibited major distributional shifts thought to be climate related. These are distributed across the following categories: species previously rare or unlisted (12), with expanded ranges (23) and/or abundance increases (30), expanded populations in south‐eastern Tasmania (16) and extra‐limital vagrants (4). Another 9 species, representing 7 families, experienced longer‐term changes (since the 1800s) probably due to anthropogenic factors, such as habitat alteration and fishing pressure: species now extinct locally (3), recovering (3), threatened (2) or with remnant populations (1). One species is a temporary resident periodically recruited from New Zealand. Of fishes exhibiting an obvious poleward movement, most are reef dwellers from three Australian biogeographic categories: widespread southern, western warm temperate (Flindersian) or eastern warm temperate (Peronian) species. Main conclusions Some of the region's largest predatory reef fishes have become extinct in Tasmanian seas since the ‘late 1800s’, most likely as a result of poor fishing practices. In more recent times, there have been major changes in the distribution patterns of Tasmanian fishes that correspond to dramatic warming observed in the local marine environment.     

Radial growth responses to drought of Pinus sylvestris and Quercus pubescens in an inner‐Alpine dry valley

Question: Lower montane treeline ecotones such as the inner Alpine dry valleys are regarded as sensitive to climate change. In the dry Valais valley (Switzerland) the composition of the widespread, low altitude Pinus forests is shifting towards a mixed deciduous state. The sub‐boreal P. sylvestris shows high mortality rates, whereas the deciduous sub‐mediterranean Quercus pubescens is spreading. These species may act as early indicators of climate change. We evaluate this hypothesis by focusing on their differences in drought tolerance, which are hardly known, but are likely to be crucial in the current forest shift and also for future forest development. Methods: We used dendroecological methods to detect species‐specific patterns in the growth response to drought. The relationship between radial growth of 401 trees from 15 mixed stands and drought was analysed by calculating response functions using yearly tree‐ring indices and monthly drought indices. PCA was applied to the response ratios to discover spatial patterns of drought response. Results: A species‐specific response to moisture as well as a sub‐regional differentiation of the response patterns were found. While Quercus showed a response mainly to the conditions of the previous autumn and those of current spring, Pinus did not start responding before May, but showed responses throughout the whole summer. Quercus may restrict physiological activity to moist periods; growth of Pinus was much more dependent on prior growth. Conclusions: Given that the climate is changing towards (1) longer summer drought periods, (2) higher mean temperatures and (3) shifted seasonally of moisture availability, Quercus may benefit from adapting better to drier conditions. Pinus may increasingly face problems related to drought stress as it depends on summer moisture and has a smaller adaptive capacity due to its long‐lived photosynthetic tissue.     

Seed germination responses to seasonal temperature and drought stress are species‐specific but not related to seed size in a desert steppe: Implications for effect of climate change on community structure

Investigating how seed germination of multiple species in an ecosystem responds to environmental conditions is crucial for understanding the mechanisms for community structure and biodiversity maintenance. However, knowledge of seed germination response of species to environmental conditions is still scarce at the community level. We hypothesized that responses of seed germination to environmental conditions differ among species at the community level, and that germination response is not correlated with seed size. To test this hypothesis, we determined the response of seed germination of 20 common species in the Siziwang Desert Steppe, China, to seasonal temperature regimes (representing April, May, June, and July) and drought stress (0, ?0.003, ?0.027, ?0.155, and ?0.87 MPa). Seed germination percentage increased with increasing temperature regime, but Allium ramosum, Allium tenuissimum, Artemisia annua, Artemisia mongolica, Artemisia scoparia, Artemisia sieversiana, Bassia dasyphylla, Kochia prastrata, and Neopallasia pectinata germinated to >60% in the lowest temperature regime (April). Germination decreased with increasing water stress, but Allium ramosum, Artemisia annua, Artemisia scoparia, Bassia dasyphylla, Heteropappus altaicus, Kochia prastrata, Neopallasia pectinata, and Potentilla tanacetifolia germinated to near 60% at ?0.87 MPa. Among these eight species, germination of six was tolerant to both temperature and water stress. Mean germination percentage in the four temperature regimes and the five water potentials was not significantly correlated with seed mass or seed area, which were highly correlated. Our results suggest that the species‐specific germination responses to environmental conditions are important in structuring the desert steppe community and have implications for predicting community structure under climate change. Thus, the predicted warmer and dryer climate will favor germination of drought‐tolerant species, resulting in altered proportions of germinants of different species and subsequently change in community composition of the desert steppe.     

Dissecting insect responses to climate warming: overwintering and post‐diapause performance in the southern green stink bug,Nezara viridula,under simulated climate‐change conditions

The effect of simulated climate change on overwintering and post‐diapause reproductive performance is studied in Nezara viridula (L.) (Heteroptera: Pentatomidae) close to the species' northern range limit in Japan. Insects are reared from October to June under quasi‐natural (i.e. ambient outdoor) conditions and in a transparent incubator, in which climate warming is simulated by adding 2.5°C to the ambient temperatures. Despite the earlier assumption that females of N. viridula overwinter in diapause, whereas males do so in quiescence, regular dissections show that the two sexes overwinter in a state of true diapause. During winter, both sexes are dark‐coloured and have undeveloped reproductive organs. Resumption of development does not start until late March. During winter, the effect of simulated warming on the dynamics and timing of physiological processes appears to be limited. However, the warming significantly enhances winter survival (from 27–31% to 47–70%), which is a key factor in range expansion of N. viridula. In spring, the effect of simulated warming is complex. It advances the post‐diapause colour change and transition from dormancy to reproduction. The earlier resumption of development is more pronounced in females: in April, significantly more females are already in a reproductive state under the simulated warming than under quasi‐natural conditions. In males, the tendency is similar, although the difference is not significant. Warming significantly enhances spring survival and percentage of copulating adults, although not the percentage of ovipositing females and fecundity. The results suggest that, under the expected climate‐warming conditions, N. viridula will likely benefit mostly as a result of increased winter and spring survival and advanced post‐diapause reproduction. Further warming is likely to allow more adults to survive the critical cold season and contribute (both numerically and by increasing heterogeneity) to the post‐overwintering population growth, thus promoting the establishment of this species in newly‐colonized areas.